id sid tid token lemma pos 10_1101-2020_12_31_424971 1 1 Insights insight NNS 10_1101-2020_12_31_424971 1 2 into into IN 10_1101-2020_12_31_424971 1 3 Genome Genome NNP 10_1101-2020_12_31_424971 1 4 Recoding recode VBG 10_1101-2020_12_31_424971 1 5 from from IN 10_1101-2020_12_31_424971 1 6 the the DT 10_1101-2020_12_31_424971 1 7 Mechanism Mechanism NNP 10_1101-2020_12_31_424971 1 8 of of IN 10_1101-2020_12_31_424971 1 9 a a DT 10_1101-2020_12_31_424971 1 10 Classic Classic NNP 10_1101-2020_12_31_424971 1 11 +1-Frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 1 12 tRNA trna NN 10_1101-2020_12_31_424971 1 13 1 1 CD 10_1101-2020_12_31_424971 1 14 Insights insight NNS 10_1101-2020_12_31_424971 1 15 into into IN 10_1101-2020_12_31_424971 1 16 Genome Genome NNP 10_1101-2020_12_31_424971 1 17 Recoding recode VBG 10_1101-2020_12_31_424971 1 18 1 1 CD 10_1101-2020_12_31_424971 1 19 from from IN 10_1101-2020_12_31_424971 1 20 the the DT 10_1101-2020_12_31_424971 1 21 Mechanism Mechanism NNP 10_1101-2020_12_31_424971 1 22 of of IN 10_1101-2020_12_31_424971 1 23 a a DT 10_1101-2020_12_31_424971 1 24 Classic Classic NNP 10_1101-2020_12_31_424971 1 25 +1-Frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 1 26 tRNA trna CD 10_1101-2020_12_31_424971 1 27 2 2 CD 10_1101-2020_12_31_424971 1 28 3 3 CD 10_1101-2020_12_31_424971 1 29 4 4 CD 10_1101-2020_12_31_424971 1 30 Howard Howard NNP 10_1101-2020_12_31_424971 1 31 Gamper1,5 gamper1,5 CD 10_1101-2020_12_31_424971 1 32 , , , 10_1101-2020_12_31_424971 1 33 Haixing haixe VBG 10_1101-2020_12_31_424971 1 34 Li2,5 li2,5 NN 10_1101-2020_12_31_424971 1 35 , , , 10_1101-2020_12_31_424971 1 36 Isao Isao NNP 10_1101-2020_12_31_424971 1 37 Masuda1 Masuda1 NNP 10_1101-2020_12_31_424971 1 38 , , , 10_1101-2020_12_31_424971 1 39 D. D. NNP 10_1101-2020_12_31_424971 1 40 Miklos Miklos NNP 10_1101-2020_12_31_424971 1 41 Robkis3 Robkis3 NNP 10_1101-2020_12_31_424971 1 42 , , , 10_1101-2020_12_31_424971 1 43 Thomas Thomas NNP 10_1101-2020_12_31_424971 1 44 Christian1 Christian1 NNP 10_1101-2020_12_31_424971 1 45 , , , 10_1101-2020_12_31_424971 1 46 5 5 CD 10_1101-2020_12_31_424971 1 47 Adam Adam NNP 10_1101-2020_12_31_424971 1 48 B. B. NNP 10_1101-2020_12_31_424971 1 49 Conn4 Conn4 NNS 10_1101-2020_12_31_424971 1 50 , , , 10_1101-2020_12_31_424971 1 51 Gregor Gregor NNP 10_1101-2020_12_31_424971 1 52 Blaha4 Blaha4 NNP 10_1101-2020_12_31_424971 1 53 , , , 10_1101-2020_12_31_424971 1 54 E. E. NNP 10_1101-2020_12_31_424971 1 55 James James NNP 10_1101-2020_12_31_424971 1 56 Petersson3 Petersson3 NNP 10_1101-2020_12_31_424971 1 57 , , , 10_1101-2020_12_31_424971 1 58 Ruben Ruben NNP 10_1101-2020_12_31_424971 1 59 L. L. NNP 10_1101-2020_12_31_424971 1 60 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 1 61 , , , 10_1101-2020_12_31_424971 1 62 Jr2 Jr2 NNP 10_1101-2020_12_31_424971 1 63 , , , 10_1101-2020_12_31_424971 1 64 # # $ 10_1101-2020_12_31_424971 1 65 , , , 10_1101-2020_12_31_424971 1 66 6 6 CD 10_1101-2020_12_31_424971 1 67 and and CC 10_1101-2020_12_31_424971 1 68 Ya Ya NNP 10_1101-2020_12_31_424971 1 69 - - HYPH 10_1101-2020_12_31_424971 1 70 Ming Ming NNP 10_1101-2020_12_31_424971 1 71 Hou1 Hou1 NNP 10_1101-2020_12_31_424971 1 72 , , , 10_1101-2020_12_31_424971 1 73 # # $ 10_1101-2020_12_31_424971 1 74 , , , 10_1101-2020_12_31_424971 1 75 * * NFP 10_1101-2020_12_31_424971 1 76 7 7 CD 10_1101-2020_12_31_424971 1 77 8 8 CD 10_1101-2020_12_31_424971 1 78 9 9 CD 10_1101-2020_12_31_424971 1 79 1Department 1department CD 10_1101-2020_12_31_424971 1 80 of of IN 10_1101-2020_12_31_424971 1 81 Biochemistry Biochemistry NNP 10_1101-2020_12_31_424971 1 82 and and CC 10_1101-2020_12_31_424971 1 83 Molecular Molecular NNP 10_1101-2020_12_31_424971 1 84 Biology Biology NNP 10_1101-2020_12_31_424971 1 85 , , , 10_1101-2020_12_31_424971 1 86 Thomas Thomas NNP 10_1101-2020_12_31_424971 1 87 Jefferson Jefferson NNP 10_1101-2020_12_31_424971 1 88 University University NNP 10_1101-2020_12_31_424971 1 89 , , , 10_1101-2020_12_31_424971 1 90 10 10 CD 10_1101-2020_12_31_424971 1 91 Philadelphia Philadelphia NNP 10_1101-2020_12_31_424971 1 92 , , , 10_1101-2020_12_31_424971 1 93 PA pa NN 10_1101-2020_12_31_424971 1 94 19107 19107 CD 10_1101-2020_12_31_424971 1 95 , , , 10_1101-2020_12_31_424971 1 96 USA USA NNP 10_1101-2020_12_31_424971 1 97 11 11 CD 10_1101-2020_12_31_424971 1 98 2Department 2department CD 10_1101-2020_12_31_424971 1 99 of of IN 10_1101-2020_12_31_424971 1 100 Chemistry Chemistry NNP 10_1101-2020_12_31_424971 1 101 , , , 10_1101-2020_12_31_424971 1 102 Columbia Columbia NNP 10_1101-2020_12_31_424971 1 103 University University NNP 10_1101-2020_12_31_424971 1 104 , , , 10_1101-2020_12_31_424971 1 105 New New NNP 10_1101-2020_12_31_424971 1 106 York York NNP 10_1101-2020_12_31_424971 1 107 , , , 10_1101-2020_12_31_424971 1 108 NY NY NNP 10_1101-2020_12_31_424971 1 109 10027 10027 CD 10_1101-2020_12_31_424971 1 110 , , , 10_1101-2020_12_31_424971 1 111 USA USA NNP 10_1101-2020_12_31_424971 1 112 12 12 CD 10_1101-2020_12_31_424971 1 113 3Department 3Department NNP 10_1101-2020_12_31_424971 1 114 of of IN 10_1101-2020_12_31_424971 1 115 Chemistry Chemistry NNP 10_1101-2020_12_31_424971 1 116 , , , 10_1101-2020_12_31_424971 1 117 University University NNP 10_1101-2020_12_31_424971 1 118 of of IN 10_1101-2020_12_31_424971 1 119 Pennsylvania Pennsylvania NNP 10_1101-2020_12_31_424971 1 120 , , , 10_1101-2020_12_31_424971 1 121 Philadelphia Philadelphia NNP 10_1101-2020_12_31_424971 1 122 , , , 10_1101-2020_12_31_424971 1 123 PA PA NNP 10_1101-2020_12_31_424971 1 124 19104 19104 CD 10_1101-2020_12_31_424971 1 125 , , , 10_1101-2020_12_31_424971 1 126 USA USA NNP 10_1101-2020_12_31_424971 1 127 13 13 CD 10_1101-2020_12_31_424971 1 128 4Department 4department NN 10_1101-2020_12_31_424971 1 129 of of IN 10_1101-2020_12_31_424971 1 130 Biochemistry Biochemistry NNP 10_1101-2020_12_31_424971 1 131 , , , 10_1101-2020_12_31_424971 1 132 University University NNP 10_1101-2020_12_31_424971 1 133 of of IN 10_1101-2020_12_31_424971 1 134 California California NNP 10_1101-2020_12_31_424971 1 135 , , , 10_1101-2020_12_31_424971 1 136 Riverside Riverside NNP 10_1101-2020_12_31_424971 1 137 , , , 10_1101-2020_12_31_424971 1 138 CA CA NNP 10_1101-2020_12_31_424971 1 139 92521 92521 CD 10_1101-2020_12_31_424971 1 140 , , , 10_1101-2020_12_31_424971 1 141 USA USA NNP 10_1101-2020_12_31_424971 1 142 14 14 CD 10_1101-2020_12_31_424971 1 143 5These 5these JJ 10_1101-2020_12_31_424971 1 144 authors author NNS 10_1101-2020_12_31_424971 1 145 contributed contribute VBD 10_1101-2020_12_31_424971 1 146 equally equally RB 10_1101-2020_12_31_424971 1 147 to to IN 10_1101-2020_12_31_424971 1 148 this this DT 10_1101-2020_12_31_424971 1 149 work work NN 10_1101-2020_12_31_424971 1 150 . . . 10_1101-2020_12_31_424971 2 1 15 15 CD 10_1101-2020_12_31_424971 2 2 # # $ 10_1101-2020_12_31_424971 2 3 Corresponding Corresponding NNP 10_1101-2020_12_31_424971 2 4 authors author NNS 10_1101-2020_12_31_424971 2 5 : : : 10_1101-2020_12_31_424971 2 6 16 16 CD 10_1101-2020_12_31_424971 2 7 rlg2118@columbia.edu rlg2118@columbia.edu NNS 10_1101-2020_12_31_424971 2 8 ( ( -LRB- 10_1101-2020_12_31_424971 2 9 T t NN 10_1101-2020_12_31_424971 2 10 ) ) -RRB- 10_1101-2020_12_31_424971 2 11 212 212 CD 10_1101-2020_12_31_424971 2 12 - - HYPH 10_1101-2020_12_31_424971 2 13 854 854 CD 10_1101-2020_12_31_424971 2 14 - - HYPH 10_1101-2020_12_31_424971 2 15 1096 1096 CD 10_1101-2020_12_31_424971 2 16 ; ; : 10_1101-2020_12_31_424971 2 17 ( ( -LRB- 10_1101-2020_12_31_424971 2 18 F f NN 10_1101-2020_12_31_424971 2 19 ) ) -RRB- 10_1101-2020_12_31_424971 2 20 212 212 CD 10_1101-2020_12_31_424971 2 21 - - HYPH 10_1101-2020_12_31_424971 2 22 932 932 CD 10_1101-2020_12_31_424971 2 23 - - HYPH 10_1101-2020_12_31_424971 2 24 1289 1289 CD 10_1101-2020_12_31_424971 2 25 ; ; : 10_1101-2020_12_31_424971 2 26 ORCID ORCID NNP 10_1101-2020_12_31_424971 2 27 : : : 10_1101-2020_12_31_424971 2 28 0000 0000 CD 10_1101-2020_12_31_424971 2 29 - - HYPH 10_1101-2020_12_31_424971 2 30 0002 0002 CD 10_1101-2020_12_31_424971 2 31 - - HYPH 10_1101-2020_12_31_424971 2 32 1344 1344 CD 10_1101-2020_12_31_424971 2 33 - - SYM 10_1101-2020_12_31_424971 2 34 5581 5581 CD 10_1101-2020_12_31_424971 2 35 17 17 CD 10_1101-2020_12_31_424971 2 36 ya-ming.hou@jefferson.edu ya-ming.hou@jefferson.edu NN 10_1101-2020_12_31_424971 2 37 ( ( -LRB- 10_1101-2020_12_31_424971 2 38 T t NN 10_1101-2020_12_31_424971 2 39 ) ) -RRB- 10_1101-2020_12_31_424971 2 40 215 215 CD 10_1101-2020_12_31_424971 2 41 - - HYPH 10_1101-2020_12_31_424971 2 42 503 503 CD 10_1101-2020_12_31_424971 2 43 - - HYPH 10_1101-2020_12_31_424971 2 44 4480 4480 CD 10_1101-2020_12_31_424971 2 45 ; ; : 10_1101-2020_12_31_424971 2 46 ( ( -LRB- 10_1101-2020_12_31_424971 2 47 F f NN 10_1101-2020_12_31_424971 2 48 ) ) -RRB- 10_1101-2020_12_31_424971 2 49 215 215 CD 10_1101-2020_12_31_424971 2 50 - - HYPH 10_1101-2020_12_31_424971 2 51 503 503 CD 10_1101-2020_12_31_424971 2 52 - - HYPH 10_1101-2020_12_31_424971 2 53 4954 4954 CD 10_1101-2020_12_31_424971 2 54 ; ; : 10_1101-2020_12_31_424971 2 55 18 18 CD 10_1101-2020_12_31_424971 2 56 ORCID orcid NN 10_1101-2020_12_31_424971 2 57 : : : 10_1101-2020_12_31_424971 2 58 0000 0000 CD 10_1101-2020_12_31_424971 2 59 - - HYPH 10_1101-2020_12_31_424971 2 60 0001 0001 CD 10_1101-2020_12_31_424971 2 61 - - HYPH 10_1101-2020_12_31_424971 2 62 6546 6546 CD 10_1101-2020_12_31_424971 2 63 - - HYPH 10_1101-2020_12_31_424971 2 64 2597 2597 CD 10_1101-2020_12_31_424971 2 65 19 19 CD 10_1101-2020_12_31_424971 2 66 20 20 CD 10_1101-2020_12_31_424971 2 67 * * NFP 10_1101-2020_12_31_424971 2 68 Lead lead JJ 10_1101-2020_12_31_424971 2 69 contact contact NN 10_1101-2020_12_31_424971 2 70 : : : 10_1101-2020_12_31_424971 2 71 Ya Ya NNP 10_1101-2020_12_31_424971 2 72 - - HYPH 10_1101-2020_12_31_424971 2 73 Ming Ming NNP 10_1101-2020_12_31_424971 2 74 Hou Hou NNP 10_1101-2020_12_31_424971 2 75 ( ( -LRB- 10_1101-2020_12_31_424971 2 76 ya-ming.hou@jefferson.edu ya-ming.hou@jefferson.edu NNP 10_1101-2020_12_31_424971 2 77 ) ) -RRB- 10_1101-2020_12_31_424971 2 78 21 21 CD 10_1101-2020_12_31_424971 2 79 22 22 CD 10_1101-2020_12_31_424971 2 80 Running Running NNP 10_1101-2020_12_31_424971 2 81 Title title NN 10_1101-2020_12_31_424971 2 82 : : : 10_1101-2020_12_31_424971 2 83 Mechanism mechanism NN 10_1101-2020_12_31_424971 2 84 of of IN 10_1101-2020_12_31_424971 2 85 SufB2-induced SufB2-induced NNP 10_1101-2020_12_31_424971 2 86 +1 +1 NNS 10_1101-2020_12_31_424971 2 87 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 2 88 23 23 CD 10_1101-2020_12_31_424971 2 89 24 24 CD 10_1101-2020_12_31_424971 2 90 25 25 CD 10_1101-2020_12_31_424971 2 91 .CC .CC : 10_1101-2020_12_31_424971 2 92 - - HYPH 10_1101-2020_12_31_424971 2 93 BY by IN 10_1101-2020_12_31_424971 2 94 - - HYPH 10_1101-2020_12_31_424971 2 95 NC NC NNP 10_1101-2020_12_31_424971 2 96 - - HYPH 10_1101-2020_12_31_424971 2 97 ND ND NNP 10_1101-2020_12_31_424971 2 98 4.0 4.0 CD 10_1101-2020_12_31_424971 2 99 International International NNP 10_1101-2020_12_31_424971 2 100 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 2 101 under under IN 10_1101-2020_12_31_424971 2 102 a a DT 10_1101-2020_12_31_424971 2 103 ( ( -LRB- 10_1101-2020_12_31_424971 2 104 which which WDT 10_1101-2020_12_31_424971 2 105 was be VBD 10_1101-2020_12_31_424971 2 106 not not RB 10_1101-2020_12_31_424971 2 107 certified certify VBN 10_1101-2020_12_31_424971 2 108 by by IN 10_1101-2020_12_31_424971 2 109 peer peer NN 10_1101-2020_12_31_424971 2 110 review review NN 10_1101-2020_12_31_424971 2 111 ) ) -RRB- 10_1101-2020_12_31_424971 2 112 is be VBZ 10_1101-2020_12_31_424971 2 113 the the DT 10_1101-2020_12_31_424971 2 114 author author NN 10_1101-2020_12_31_424971 2 115 / / SYM 10_1101-2020_12_31_424971 2 116 funder funder NN 10_1101-2020_12_31_424971 2 117 , , , 10_1101-2020_12_31_424971 2 118 who who WP 10_1101-2020_12_31_424971 2 119 has have VBZ 10_1101-2020_12_31_424971 2 120 granted grant VBN 10_1101-2020_12_31_424971 2 121 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 2 122 a a DT 10_1101-2020_12_31_424971 2 123 license license NN 10_1101-2020_12_31_424971 2 124 to to TO 10_1101-2020_12_31_424971 2 125 display display VB 10_1101-2020_12_31_424971 2 126 the the DT 10_1101-2020_12_31_424971 2 127 preprint preprint NN 10_1101-2020_12_31_424971 2 128 in in IN 10_1101-2020_12_31_424971 2 129 perpetuity perpetuity NN 10_1101-2020_12_31_424971 2 130 . . . 10_1101-2020_12_31_424971 3 1 It -PRON- PRP 10_1101-2020_12_31_424971 3 2 is be VBZ 10_1101-2020_12_31_424971 3 3 made make VBN 10_1101-2020_12_31_424971 3 4 The the DT 10_1101-2020_12_31_424971 3 5 copyright copyright NN 10_1101-2020_12_31_424971 3 6 holder holder NN 10_1101-2020_12_31_424971 3 7 for for IN 10_1101-2020_12_31_424971 3 8 this this DT 10_1101-2020_12_31_424971 3 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 3 10 version version NN 10_1101-2020_12_31_424971 3 11 posted post VBD 10_1101-2020_12_31_424971 3 12 January January NNP 10_1101-2020_12_31_424971 3 13 2 2 CD 10_1101-2020_12_31_424971 3 14 , , , 10_1101-2020_12_31_424971 3 15 2021 2021 CD 10_1101-2020_12_31_424971 3 16 . . . 10_1101-2020_12_31_424971 3 17 ; ; : 10_1101-2020_12_31_424971 3 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 3 19 : : : 10_1101-2020_12_31_424971 3 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 3 21 preprint preprint NN 10_1101-2020_12_31_424971 3 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 3 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 3 24 2 2 CD 10_1101-2020_12_31_424971 3 25 ABSTRACT ABSTRACT NNP 10_1101-2020_12_31_424971 3 26 26 26 CD 10_1101-2020_12_31_424971 3 27 While while IN 10_1101-2020_12_31_424971 3 28 genome genome JJ 10_1101-2020_12_31_424971 3 29 recoding recode VBG 10_1101-2020_12_31_424971 3 30 using use VBG 10_1101-2020_12_31_424971 3 31 quadruplet quadruplet NN 10_1101-2020_12_31_424971 3 32 codons codon NNS 10_1101-2020_12_31_424971 3 33 to to TO 10_1101-2020_12_31_424971 3 34 incorporate incorporate VB 10_1101-2020_12_31_424971 3 35 non non JJ 10_1101-2020_12_31_424971 3 36 - - JJ 10_1101-2020_12_31_424971 3 37 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 3 38 amino amino NN 10_1101-2020_12_31_424971 3 39 27 27 CD 10_1101-2020_12_31_424971 3 40 acids acid NNS 10_1101-2020_12_31_424971 3 41 is be VBZ 10_1101-2020_12_31_424971 3 42 attractive attractive JJ 10_1101-2020_12_31_424971 3 43 for for IN 10_1101-2020_12_31_424971 3 44 biotechnology biotechnology NN 10_1101-2020_12_31_424971 3 45 and and CC 10_1101-2020_12_31_424971 3 46 bioengineering bioengineering NN 10_1101-2020_12_31_424971 3 47 purposes purpose NNS 10_1101-2020_12_31_424971 3 48 , , , 10_1101-2020_12_31_424971 3 49 the the DT 10_1101-2020_12_31_424971 3 50 mechanism mechanism NN 10_1101-2020_12_31_424971 3 51 through through IN 10_1101-2020_12_31_424971 3 52 which which WDT 10_1101-2020_12_31_424971 3 53 28 28 CD 10_1101-2020_12_31_424971 3 54 such such JJ 10_1101-2020_12_31_424971 3 55 codons codon NNS 10_1101-2020_12_31_424971 3 56 are be VBP 10_1101-2020_12_31_424971 3 57 translated translate VBN 10_1101-2020_12_31_424971 3 58 is be VBZ 10_1101-2020_12_31_424971 3 59 poorly poorly RB 10_1101-2020_12_31_424971 3 60 understood understand VBN 10_1101-2020_12_31_424971 3 61 . . . 10_1101-2020_12_31_424971 4 1 Here here RB 10_1101-2020_12_31_424971 4 2 we -PRON- PRP 10_1101-2020_12_31_424971 4 3 investigate investigate VBP 10_1101-2020_12_31_424971 4 4 translation translation NN 10_1101-2020_12_31_424971 4 5 of of IN 10_1101-2020_12_31_424971 4 6 quadruplet quadruplet NN 10_1101-2020_12_31_424971 4 7 29 29 CD 10_1101-2020_12_31_424971 4 8 codons codon NNS 10_1101-2020_12_31_424971 4 9 by by IN 10_1101-2020_12_31_424971 4 10 a a DT 10_1101-2020_12_31_424971 4 11 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 4 12 tRNA trna NN 10_1101-2020_12_31_424971 4 13 , , , 10_1101-2020_12_31_424971 4 14 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 4 15 , , , 10_1101-2020_12_31_424971 4 16 that that WDT 10_1101-2020_12_31_424971 4 17 contains contain VBZ 10_1101-2020_12_31_424971 4 18 an an DT 10_1101-2020_12_31_424971 4 19 extra extra JJ 10_1101-2020_12_31_424971 4 20 nucleotide nucleotide NN 10_1101-2020_12_31_424971 4 21 in in IN 10_1101-2020_12_31_424971 4 22 its -PRON- PRP$ 10_1101-2020_12_31_424971 4 23 anticodon anticodon NN 10_1101-2020_12_31_424971 4 24 loop loop NN 10_1101-2020_12_31_424971 4 25 . . . 10_1101-2020_12_31_424971 5 1 30 30 CD 10_1101-2020_12_31_424971 5 2 Natural natural JJ 10_1101-2020_12_31_424971 5 3 post post JJ 10_1101-2020_12_31_424971 5 4 - - JJ 10_1101-2020_12_31_424971 5 5 transcriptional transcriptional JJ 10_1101-2020_12_31_424971 5 6 modification modification NN 10_1101-2020_12_31_424971 5 7 of of IN 10_1101-2020_12_31_424971 5 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 5 9 in in IN 10_1101-2020_12_31_424971 5 10 cells cell NNS 10_1101-2020_12_31_424971 5 11 prevents prevent VBZ 10_1101-2020_12_31_424971 5 12 it -PRON- PRP 10_1101-2020_12_31_424971 5 13 from from IN 10_1101-2020_12_31_424971 5 14 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 5 15 using use VBG 10_1101-2020_12_31_424971 5 16 a a DT 10_1101-2020_12_31_424971 5 17 31 31 CD 10_1101-2020_12_31_424971 5 18 quadruplet quadruplet NN 10_1101-2020_12_31_424971 5 19 - - HYPH 10_1101-2020_12_31_424971 5 20 pairing pair VBG 10_1101-2020_12_31_424971 5 21 mechanism mechanism NN 10_1101-2020_12_31_424971 5 22 such such JJ 10_1101-2020_12_31_424971 5 23 that that IN 10_1101-2020_12_31_424971 5 24 it -PRON- PRP 10_1101-2020_12_31_424971 5 25 preferentially preferentially RB 10_1101-2020_12_31_424971 5 26 employs employ VBZ 10_1101-2020_12_31_424971 5 27 a a DT 10_1101-2020_12_31_424971 5 28 triplet triplet NN 10_1101-2020_12_31_424971 5 29 - - HYPH 10_1101-2020_12_31_424971 5 30 slippage slippage NN 10_1101-2020_12_31_424971 5 31 mechanism mechanism NN 10_1101-2020_12_31_424971 5 32 . . . 10_1101-2020_12_31_424971 6 1 32 32 CD 10_1101-2020_12_31_424971 6 2 We -PRON- PRP 10_1101-2020_12_31_424971 6 3 show show VBP 10_1101-2020_12_31_424971 6 4 that that IN 10_1101-2020_12_31_424971 6 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 6 6 uses use VBZ 10_1101-2020_12_31_424971 6 7 triplet triplet NN 10_1101-2020_12_31_424971 6 8 anticodon anticodon NN 10_1101-2020_12_31_424971 6 9 - - HYPH 10_1101-2020_12_31_424971 6 10 codon codon NNP 10_1101-2020_12_31_424971 6 11 pairing pair VBG 10_1101-2020_12_31_424971 6 12 in in IN 10_1101-2020_12_31_424971 6 13 the the DT 10_1101-2020_12_31_424971 6 14 0-frame 0-frame CD 10_1101-2020_12_31_424971 6 15 to to TO 10_1101-2020_12_31_424971 6 16 initially initially RB 10_1101-2020_12_31_424971 6 17 decode decode VB 10_1101-2020_12_31_424971 6 18 the the DT 10_1101-2020_12_31_424971 6 19 33 33 CD 10_1101-2020_12_31_424971 6 20 quadruplet quadruplet NN 10_1101-2020_12_31_424971 6 21 codon codon NN 10_1101-2020_12_31_424971 6 22 , , , 10_1101-2020_12_31_424971 6 23 but but CC 10_1101-2020_12_31_424971 6 24 subsequently subsequently RB 10_1101-2020_12_31_424971 6 25 shifts shift VBZ 10_1101-2020_12_31_424971 6 26 to to IN 10_1101-2020_12_31_424971 6 27 the the DT 10_1101-2020_12_31_424971 6 28 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 6 29 during during IN 10_1101-2020_12_31_424971 6 30 tRNA tRNA NNP 10_1101-2020_12_31_424971 6 31 - - HYPH 10_1101-2020_12_31_424971 6 32 mRNA mrna NN 10_1101-2020_12_31_424971 6 33 translocation translocation NN 10_1101-2020_12_31_424971 6 34 . . . 10_1101-2020_12_31_424971 7 1 34 34 CD 10_1101-2020_12_31_424971 7 2 SufB2 sufb2 NN 10_1101-2020_12_31_424971 7 3 frameshifting frameshifting NN 10_1101-2020_12_31_424971 7 4 involves involve VBZ 10_1101-2020_12_31_424971 7 5 perturbation perturbation NN 10_1101-2020_12_31_424971 7 6 of of IN 10_1101-2020_12_31_424971 7 7 an an DT 10_1101-2020_12_31_424971 7 8 essential essential JJ 10_1101-2020_12_31_424971 7 9 ribosome ribosome JJ 10_1101-2020_12_31_424971 7 10 conformational conformational JJ 10_1101-2020_12_31_424971 7 11 change change NN 10_1101-2020_12_31_424971 7 12 that that IN 10_1101-2020_12_31_424971 7 13 35 35 CD 10_1101-2020_12_31_424971 7 14 facilitates facilitate NNS 10_1101-2020_12_31_424971 7 15 tRNA trna NN 10_1101-2020_12_31_424971 7 16 - - HYPH 10_1101-2020_12_31_424971 7 17 mRNA mRNA NNS 10_1101-2020_12_31_424971 7 18 movements movement NNS 10_1101-2020_12_31_424971 7 19 at at IN 10_1101-2020_12_31_424971 7 20 a a DT 10_1101-2020_12_31_424971 7 21 late late JJ 10_1101-2020_12_31_424971 7 22 stage stage NN 10_1101-2020_12_31_424971 7 23 of of IN 10_1101-2020_12_31_424971 7 24 the the DT 10_1101-2020_12_31_424971 7 25 translocation translocation NN 10_1101-2020_12_31_424971 7 26 reaction reaction NN 10_1101-2020_12_31_424971 7 27 . . . 10_1101-2020_12_31_424971 8 1 Our -PRON- PRP$ 10_1101-2020_12_31_424971 8 2 results result NNS 10_1101-2020_12_31_424971 8 3 36 36 CD 10_1101-2020_12_31_424971 8 4 provide provide VBP 10_1101-2020_12_31_424971 8 5 a a DT 10_1101-2020_12_31_424971 8 6 molecular molecular JJ 10_1101-2020_12_31_424971 8 7 mechanism mechanism NN 10_1101-2020_12_31_424971 8 8 for for IN 10_1101-2020_12_31_424971 8 9 SufB2-induced SufB2-induced NNP 10_1101-2020_12_31_424971 8 10 +1 +1 NNS 10_1101-2020_12_31_424971 8 11 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 8 12 and and CC 10_1101-2020_12_31_424971 8 13 suggest suggest VBP 10_1101-2020_12_31_424971 8 14 that that IN 10_1101-2020_12_31_424971 8 15 engineering engineer VBG 10_1101-2020_12_31_424971 8 16 37 37 CD 10_1101-2020_12_31_424971 8 17 of of IN 10_1101-2020_12_31_424971 8 18 a a DT 10_1101-2020_12_31_424971 8 19 specific specific JJ 10_1101-2020_12_31_424971 8 20 ribosome ribosome JJ 10_1101-2020_12_31_424971 8 21 conformational conformational JJ 10_1101-2020_12_31_424971 8 22 change change NN 10_1101-2020_12_31_424971 8 23 can can MD 10_1101-2020_12_31_424971 8 24 improve improve VB 10_1101-2020_12_31_424971 8 25 the the DT 10_1101-2020_12_31_424971 8 26 efficiency efficiency NN 10_1101-2020_12_31_424971 8 27 of of IN 10_1101-2020_12_31_424971 8 28 genome genome JJ 10_1101-2020_12_31_424971 8 29 recoding recoding NN 10_1101-2020_12_31_424971 8 30 . . . 10_1101-2020_12_31_424971 9 1 38 38 CD 10_1101-2020_12_31_424971 9 2 39 39 CD 10_1101-2020_12_31_424971 9 3 Key key JJ 10_1101-2020_12_31_424971 9 4 words word NNS 10_1101-2020_12_31_424971 9 5 : : : 10_1101-2020_12_31_424971 9 6 SufB2 sufb2 JJ 10_1101-2020_12_31_424971 9 7 frameshift frameshift NN 10_1101-2020_12_31_424971 9 8 suppressor suppressor NN 10_1101-2020_12_31_424971 9 9 tRNA trna NN 10_1101-2020_12_31_424971 9 10 , , , 10_1101-2020_12_31_424971 9 11 +1 +1 : 10_1101-2020_12_31_424971 9 12 ribosomal ribosomal NN 10_1101-2020_12_31_424971 9 13 frameshifting frameshifting NN 10_1101-2020_12_31_424971 9 14 , , , 10_1101-2020_12_31_424971 9 15 quadruplet quadruplet NN 10_1101-2020_12_31_424971 9 16 codon codon NNP 10_1101-2020_12_31_424971 9 17 , , , 10_1101-2020_12_31_424971 9 18 40 40 CD 10_1101-2020_12_31_424971 9 19 genome genome JJ 10_1101-2020_12_31_424971 9 20 expansion expansion NN 10_1101-2020_12_31_424971 9 21 , , , 10_1101-2020_12_31_424971 9 22 m1G37 m1G37 NNP 10_1101-2020_12_31_424971 9 23 methylation methylation CD 10_1101-2020_12_31_424971 9 24 41 41 CD 10_1101-2020_12_31_424971 9 25 42 42 CD 10_1101-2020_12_31_424971 9 26 43 43 CD 10_1101-2020_12_31_424971 9 27 44 44 CD 10_1101-2020_12_31_424971 9 28 45 45 CD 10_1101-2020_12_31_424971 9 29 46 46 CD 10_1101-2020_12_31_424971 9 30 47 47 CD 10_1101-2020_12_31_424971 9 31 48 48 CD 10_1101-2020_12_31_424971 9 32 .CC .cc CD 10_1101-2020_12_31_424971 9 33 - - HYPH 10_1101-2020_12_31_424971 9 34 BY by IN 10_1101-2020_12_31_424971 9 35 - - HYPH 10_1101-2020_12_31_424971 9 36 NC NC NNP 10_1101-2020_12_31_424971 9 37 - - HYPH 10_1101-2020_12_31_424971 9 38 ND ND NNP 10_1101-2020_12_31_424971 9 39 4.0 4.0 CD 10_1101-2020_12_31_424971 9 40 International International NNP 10_1101-2020_12_31_424971 9 41 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 9 42 under under IN 10_1101-2020_12_31_424971 9 43 a a DT 10_1101-2020_12_31_424971 9 44 ( ( -LRB- 10_1101-2020_12_31_424971 9 45 which which WDT 10_1101-2020_12_31_424971 9 46 was be VBD 10_1101-2020_12_31_424971 9 47 not not RB 10_1101-2020_12_31_424971 9 48 certified certify VBN 10_1101-2020_12_31_424971 9 49 by by IN 10_1101-2020_12_31_424971 9 50 peer peer NN 10_1101-2020_12_31_424971 9 51 review review NN 10_1101-2020_12_31_424971 9 52 ) ) -RRB- 10_1101-2020_12_31_424971 9 53 is be VBZ 10_1101-2020_12_31_424971 9 54 the the DT 10_1101-2020_12_31_424971 9 55 author author NN 10_1101-2020_12_31_424971 9 56 / / SYM 10_1101-2020_12_31_424971 9 57 funder funder NN 10_1101-2020_12_31_424971 9 58 , , , 10_1101-2020_12_31_424971 9 59 who who WP 10_1101-2020_12_31_424971 9 60 has have VBZ 10_1101-2020_12_31_424971 9 61 granted grant VBN 10_1101-2020_12_31_424971 9 62 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 9 63 a a DT 10_1101-2020_12_31_424971 9 64 license license NN 10_1101-2020_12_31_424971 9 65 to to TO 10_1101-2020_12_31_424971 9 66 display display VB 10_1101-2020_12_31_424971 9 67 the the DT 10_1101-2020_12_31_424971 9 68 preprint preprint NN 10_1101-2020_12_31_424971 9 69 in in IN 10_1101-2020_12_31_424971 9 70 perpetuity perpetuity NN 10_1101-2020_12_31_424971 9 71 . . . 10_1101-2020_12_31_424971 10 1 It -PRON- PRP 10_1101-2020_12_31_424971 10 2 is be VBZ 10_1101-2020_12_31_424971 10 3 made make VBN 10_1101-2020_12_31_424971 10 4 The the DT 10_1101-2020_12_31_424971 10 5 copyright copyright NN 10_1101-2020_12_31_424971 10 6 holder holder NN 10_1101-2020_12_31_424971 10 7 for for IN 10_1101-2020_12_31_424971 10 8 this this DT 10_1101-2020_12_31_424971 10 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 10 10 version version NN 10_1101-2020_12_31_424971 10 11 posted post VBD 10_1101-2020_12_31_424971 10 12 January January NNP 10_1101-2020_12_31_424971 10 13 2 2 CD 10_1101-2020_12_31_424971 10 14 , , , 10_1101-2020_12_31_424971 10 15 2021 2021 CD 10_1101-2020_12_31_424971 10 16 . . . 10_1101-2020_12_31_424971 10 17 ; ; : 10_1101-2020_12_31_424971 10 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 10 19 : : : 10_1101-2020_12_31_424971 10 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 10 21 preprint preprint NN 10_1101-2020_12_31_424971 10 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 10 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 10 24 3 3 CD 10_1101-2020_12_31_424971 10 25 INTRODUCTION introduction NN 10_1101-2020_12_31_424971 10 26 49 49 CD 10_1101-2020_12_31_424971 10 27 The the DT 10_1101-2020_12_31_424971 10 28 ability ability NN 10_1101-2020_12_31_424971 10 29 to to TO 10_1101-2020_12_31_424971 10 30 recode recode VB 10_1101-2020_12_31_424971 10 31 the the DT 10_1101-2020_12_31_424971 10 32 genome genome NN 10_1101-2020_12_31_424971 10 33 and and CC 10_1101-2020_12_31_424971 10 34 expand expand VB 10_1101-2020_12_31_424971 10 35 the the DT 10_1101-2020_12_31_424971 10 36 chemical chemical NN 10_1101-2020_12_31_424971 10 37 repertoire repertoire NN 10_1101-2020_12_31_424971 10 38 of of IN 10_1101-2020_12_31_424971 10 39 proteins protein NNS 10_1101-2020_12_31_424971 10 40 to to TO 10_1101-2020_12_31_424971 10 41 include include VB 10_1101-2020_12_31_424971 10 42 50 50 CD 10_1101-2020_12_31_424971 10 43 non non JJ 10_1101-2020_12_31_424971 10 44 - - JJ 10_1101-2020_12_31_424971 10 45 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 10 46 amino amino JJ 10_1101-2020_12_31_424971 10 47 acids acid NNS 10_1101-2020_12_31_424971 10 48 promises promise VBZ 10_1101-2020_12_31_424971 10 49 novel novel JJ 10_1101-2020_12_31_424971 10 50 tools tool NNS 10_1101-2020_12_31_424971 10 51 for for IN 10_1101-2020_12_31_424971 10 52 probing probe VBG 10_1101-2020_12_31_424971 10 53 protein protein NN 10_1101-2020_12_31_424971 10 54 structure structure NN 10_1101-2020_12_31_424971 10 55 and and CC 10_1101-2020_12_31_424971 10 56 function function NN 10_1101-2020_12_31_424971 10 57 . . . 10_1101-2020_12_31_424971 11 1 51 51 CD 10_1101-2020_12_31_424971 11 2 While while IN 10_1101-2020_12_31_424971 11 3 most most JJS 10_1101-2020_12_31_424971 11 4 recoding recoding JJ 10_1101-2020_12_31_424971 11 5 employs employ NNS 10_1101-2020_12_31_424971 11 6 stop stop VBP 10_1101-2020_12_31_424971 11 7 codons codon NNS 10_1101-2020_12_31_424971 11 8 as as IN 10_1101-2020_12_31_424971 11 9 sites site NNS 10_1101-2020_12_31_424971 11 10 for for IN 10_1101-2020_12_31_424971 11 11 incorporating incorporate VBG 10_1101-2020_12_31_424971 11 12 non non JJ 10_1101-2020_12_31_424971 11 13 - - JJ 10_1101-2020_12_31_424971 11 14 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 11 15 amino amino NN 10_1101-2020_12_31_424971 11 16 52 52 CD 10_1101-2020_12_31_424971 11 17 acids acid NNS 10_1101-2020_12_31_424971 11 18 , , , 10_1101-2020_12_31_424971 11 19 only only RB 10_1101-2020_12_31_424971 11 20 two two CD 10_1101-2020_12_31_424971 11 21 stop stop NN 10_1101-2020_12_31_424971 11 22 codons codon NNS 10_1101-2020_12_31_424971 11 23 can can MD 10_1101-2020_12_31_424971 11 24 be be VB 10_1101-2020_12_31_424971 11 25 simultaneously simultaneously RB 10_1101-2020_12_31_424971 11 26 recoded recode VBN 10_1101-2020_12_31_424971 11 27 due due IN 10_1101-2020_12_31_424971 11 28 to to IN 10_1101-2020_12_31_424971 11 29 the the DT 10_1101-2020_12_31_424971 11 30 cellular cellular JJ 10_1101-2020_12_31_424971 11 31 need need NN 10_1101-2020_12_31_424971 11 32 to to TO 10_1101-2020_12_31_424971 11 33 reserve reserve VB 10_1101-2020_12_31_424971 11 34 53 53 CD 10_1101-2020_12_31_424971 11 35 the the DT 10_1101-2020_12_31_424971 11 36 third third JJ 10_1101-2020_12_31_424971 11 37 stop stop NN 10_1101-2020_12_31_424971 11 38 codon codon NNP 10_1101-2020_12_31_424971 11 39 for for IN 10_1101-2020_12_31_424971 11 40 termination termination NN 10_1101-2020_12_31_424971 11 41 of of IN 10_1101-2020_12_31_424971 11 42 protein protein NN 10_1101-2020_12_31_424971 11 43 synthesis synthesis NN 10_1101-2020_12_31_424971 11 44 . . . 10_1101-2020_12_31_424971 12 1 The the DT 10_1101-2020_12_31_424971 12 2 use use NN 10_1101-2020_12_31_424971 12 3 of of IN 10_1101-2020_12_31_424971 12 4 quadruplet quadruplet NN 10_1101-2020_12_31_424971 12 5 codons codon NNS 10_1101-2020_12_31_424971 12 6 as as IN 10_1101-2020_12_31_424971 12 7 54 54 CD 10_1101-2020_12_31_424971 12 8 additional additional JJ 10_1101-2020_12_31_424971 12 9 sites site NNS 10_1101-2020_12_31_424971 12 10 for for IN 10_1101-2020_12_31_424971 12 11 incorporating incorporate VBG 10_1101-2020_12_31_424971 12 12 non non JJ 10_1101-2020_12_31_424971 12 13 - - JJ 10_1101-2020_12_31_424971 12 14 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 12 15 amino amino JJ 10_1101-2020_12_31_424971 12 16 acids acid NNS 10_1101-2020_12_31_424971 12 17 has have VBZ 10_1101-2020_12_31_424971 12 18 thus thus RB 10_1101-2020_12_31_424971 12 19 emerged emerge VBN 10_1101-2020_12_31_424971 12 20 as as IN 10_1101-2020_12_31_424971 12 21 an an DT 10_1101-2020_12_31_424971 12 22 attractive attractive JJ 10_1101-2020_12_31_424971 12 23 55 55 CD 10_1101-2020_12_31_424971 12 24 alternative1,2 alternative1,2 CD 10_1101-2020_12_31_424971 12 25 . . . 10_1101-2020_12_31_424971 13 1 Recoding recode VBG 10_1101-2020_12_31_424971 13 2 at at IN 10_1101-2020_12_31_424971 13 3 a a DT 10_1101-2020_12_31_424971 13 4 quadruplet quadruplet NN 10_1101-2020_12_31_424971 13 5 codon codon NN 10_1101-2020_12_31_424971 13 6 requires require VBZ 10_1101-2020_12_31_424971 13 7 a a DT 10_1101-2020_12_31_424971 13 8 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 13 9 tRNA trna NN 10_1101-2020_12_31_424971 13 10 that that WDT 10_1101-2020_12_31_424971 13 11 is be VBZ 10_1101-2020_12_31_424971 13 12 56 56 CD 10_1101-2020_12_31_424971 13 13 aminoacylated aminoacylate VBN 10_1101-2020_12_31_424971 13 14 with with IN 10_1101-2020_12_31_424971 13 15 the the DT 10_1101-2020_12_31_424971 13 16 non non JJ 10_1101-2020_12_31_424971 13 17 - - JJ 10_1101-2020_12_31_424971 13 18 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 13 19 amino amino NN 10_1101-2020_12_31_424971 13 20 acid acid NN 10_1101-2020_12_31_424971 13 21 of of IN 10_1101-2020_12_31_424971 13 22 interest interest NN 10_1101-2020_12_31_424971 13 23 . . . 10_1101-2020_12_31_424971 14 1 The the DT 10_1101-2020_12_31_424971 14 2 primary primary JJ 10_1101-2020_12_31_424971 14 3 challenge challenge NN 10_1101-2020_12_31_424971 14 4 faced face VBN 10_1101-2020_12_31_424971 14 5 by by IN 10_1101-2020_12_31_424971 14 6 57 57 CD 10_1101-2020_12_31_424971 14 7 this this DT 10_1101-2020_12_31_424971 14 8 technology technology NN 10_1101-2020_12_31_424971 14 9 has have VBZ 10_1101-2020_12_31_424971 14 10 been be VBN 10_1101-2020_12_31_424971 14 11 the the DT 10_1101-2020_12_31_424971 14 12 low low JJ 10_1101-2020_12_31_424971 14 13 efficiency efficiency NN 10_1101-2020_12_31_424971 14 14 with with IN 10_1101-2020_12_31_424971 14 15 which which WDT 10_1101-2020_12_31_424971 14 16 the the DT 10_1101-2020_12_31_424971 14 17 full full JJ 10_1101-2020_12_31_424971 14 18 - - HYPH 10_1101-2020_12_31_424971 14 19 length length NN 10_1101-2020_12_31_424971 14 20 protein protein NN 10_1101-2020_12_31_424971 14 21 carrying carry VBG 10_1101-2020_12_31_424971 14 22 the the DT 10_1101-2020_12_31_424971 14 23 non-58 non-58 JJ 10_1101-2020_12_31_424971 14 24 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 14 25 amino amino NN 10_1101-2020_12_31_424971 14 26 acid acid NN 10_1101-2020_12_31_424971 14 27 can can MD 10_1101-2020_12_31_424971 14 28 be be VB 10_1101-2020_12_31_424971 14 29 synthesized synthesize VBN 10_1101-2020_12_31_424971 14 30 . . . 10_1101-2020_12_31_424971 15 1 One one CD 10_1101-2020_12_31_424971 15 2 reason reason NN 10_1101-2020_12_31_424971 15 3 for for IN 10_1101-2020_12_31_424971 15 4 this this DT 10_1101-2020_12_31_424971 15 5 is be VBZ 10_1101-2020_12_31_424971 15 6 the the DT 10_1101-2020_12_31_424971 15 7 poor poor JJ 10_1101-2020_12_31_424971 15 8 recoding recode VBG 10_1101-2020_12_31_424971 15 9 efficiency efficiency NN 10_1101-2020_12_31_424971 15 10 59 59 CD 10_1101-2020_12_31_424971 15 11 of of IN 10_1101-2020_12_31_424971 15 12 the the DT 10_1101-2020_12_31_424971 15 13 +1-frameshifting +1-frameshifting NNP 10_1101-2020_12_31_424971 15 14 aminoacyl aminoacyl NN 10_1101-2020_12_31_424971 15 15 ( ( -LRB- 10_1101-2020_12_31_424971 15 16 aa)-tRNA aa)-tRNA NNP 10_1101-2020_12_31_424971 15 17 , , , 10_1101-2020_12_31_424971 15 18 and and CC 10_1101-2020_12_31_424971 15 19 the the DT 10_1101-2020_12_31_424971 15 20 second second JJ 10_1101-2020_12_31_424971 15 21 is be VBZ 10_1101-2020_12_31_424971 15 22 the the DT 10_1101-2020_12_31_424971 15 23 failure failure NN 10_1101-2020_12_31_424971 15 24 of of IN 10_1101-2020_12_31_424971 15 25 the the DT 10_1101-2020_12_31_424971 15 26 +1-frameshifting +1-frameshifting NNP 10_1101-2020_12_31_424971 15 27 60 60 CD 10_1101-2020_12_31_424971 15 28 aa aa NNP 10_1101-2020_12_31_424971 15 29 - - HYPH 10_1101-2020_12_31_424971 15 30 tRNA trna NN 10_1101-2020_12_31_424971 15 31 to to TO 10_1101-2020_12_31_424971 15 32 compete compete VB 10_1101-2020_12_31_424971 15 33 with with IN 10_1101-2020_12_31_424971 15 34 canonical canonical JJ 10_1101-2020_12_31_424971 15 35 aa aa NNP 10_1101-2020_12_31_424971 15 36 - - HYPH 10_1101-2020_12_31_424971 15 37 tRNAs tRNAs NNP 10_1101-2020_12_31_424971 15 38 that that WDT 10_1101-2020_12_31_424971 15 39 read read VBP 10_1101-2020_12_31_424971 15 40 the the DT 10_1101-2020_12_31_424971 15 41 first first JJ 10_1101-2020_12_31_424971 15 42 three three CD 10_1101-2020_12_31_424971 15 43 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 15 44 of of IN 10_1101-2020_12_31_424971 15 45 the the DT 10_1101-2020_12_31_424971 15 46 61 61 CD 10_1101-2020_12_31_424971 15 47 quadruplet quadruplet NN 10_1101-2020_12_31_424971 15 48 codon codon NNP 10_1101-2020_12_31_424971 15 49 at at IN 10_1101-2020_12_31_424971 15 50 the the DT 10_1101-2020_12_31_424971 15 51 ribosomal ribosomal NNP 10_1101-2020_12_31_424971 15 52 aa aa NNP 10_1101-2020_12_31_424971 15 53 - - HYPH 10_1101-2020_12_31_424971 15 54 tRNA tRNA NNP 10_1101-2020_12_31_424971 15 55 binding binding NN 10_1101-2020_12_31_424971 15 56 ( ( -LRB- 10_1101-2020_12_31_424971 15 57 A a NN 10_1101-2020_12_31_424971 15 58 ) ) -RRB- 10_1101-2020_12_31_424971 15 59 site site NN 10_1101-2020_12_31_424971 15 60 during during IN 10_1101-2020_12_31_424971 15 61 the the DT 10_1101-2020_12_31_424971 15 62 aa aa NNP 10_1101-2020_12_31_424971 15 63 - - HYPH 10_1101-2020_12_31_424971 15 64 tRNA tRNA NNP 10_1101-2020_12_31_424971 15 65 selection selection NN 10_1101-2020_12_31_424971 15 66 step step NN 10_1101-2020_12_31_424971 15 67 of of IN 10_1101-2020_12_31_424971 15 68 62 62 CD 10_1101-2020_12_31_424971 15 69 the the DT 10_1101-2020_12_31_424971 15 70 translation translation NN 10_1101-2020_12_31_424971 15 71 elongation elongation NN 10_1101-2020_12_31_424971 15 72 cycle cycle NN 10_1101-2020_12_31_424971 15 73 . . . 10_1101-2020_12_31_424971 16 1 While while IN 10_1101-2020_12_31_424971 16 2 directed direct VBN 10_1101-2020_12_31_424971 16 3 evolution evolution NN 10_1101-2020_12_31_424971 16 4 by by IN 10_1101-2020_12_31_424971 16 5 synthetic synthetic JJ 10_1101-2020_12_31_424971 16 6 biologists biologist NNS 10_1101-2020_12_31_424971 16 7 has have VBZ 10_1101-2020_12_31_424971 16 8 yielded yield VBN 10_1101-2020_12_31_424971 16 9 +1 +1 NNPS 10_1101-2020_12_31_424971 16 10 - - : 10_1101-2020_12_31_424971 16 11 63 63 CD 10_1101-2020_12_31_424971 16 12 frameshifting frameshifting NN 10_1101-2020_12_31_424971 16 13 tRNAs trnas CD 10_1101-2020_12_31_424971 16 14 , , , 10_1101-2020_12_31_424971 16 15 efficient efficient JJ 10_1101-2020_12_31_424971 16 16 recoding recoding NN 10_1101-2020_12_31_424971 16 17 requires require VBZ 10_1101-2020_12_31_424971 16 18 cell cell NN 10_1101-2020_12_31_424971 16 19 lines line NNS 10_1101-2020_12_31_424971 16 20 that that WDT 10_1101-2020_12_31_424971 16 21 have have VBP 10_1101-2020_12_31_424971 16 22 been be VBN 10_1101-2020_12_31_424971 16 23 engineered engineer VBN 10_1101-2020_12_31_424971 16 24 to to TO 10_1101-2020_12_31_424971 16 25 deplete deplete VB 10_1101-2020_12_31_424971 16 26 64 64 CD 10_1101-2020_12_31_424971 16 27 potential potential JJ 10_1101-2020_12_31_424971 16 28 competitor competitor NN 10_1101-2020_12_31_424971 16 29 tRNAs3 tRNAs3 '' 10_1101-2020_12_31_424971 16 30 - - SYM 10_1101-2020_12_31_424971 16 31 8 8 CD 10_1101-2020_12_31_424971 16 32 . . . 10_1101-2020_12_31_424971 17 1 These these DT 10_1101-2020_12_31_424971 17 2 problems problem NNS 10_1101-2020_12_31_424971 17 3 emphasize emphasize VBP 10_1101-2020_12_31_424971 17 4 the the DT 10_1101-2020_12_31_424971 17 5 need need NN 10_1101-2020_12_31_424971 17 6 to to TO 10_1101-2020_12_31_424971 17 7 better well RBR 10_1101-2020_12_31_424971 17 8 understand understand VB 10_1101-2020_12_31_424971 17 9 the the DT 10_1101-2020_12_31_424971 17 10 65 65 CD 10_1101-2020_12_31_424971 17 11 mechanism mechanism NN 10_1101-2020_12_31_424971 17 12 through through IN 10_1101-2020_12_31_424971 17 13 which which WDT 10_1101-2020_12_31_424971 17 14 quadruplet quadruplet NN 10_1101-2020_12_31_424971 17 15 codons codon NNS 10_1101-2020_12_31_424971 17 16 are be VBP 10_1101-2020_12_31_424971 17 17 translated translate VBN 10_1101-2020_12_31_424971 17 18 by by IN 10_1101-2020_12_31_424971 17 19 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 17 20 tRNAs trnas CD 10_1101-2020_12_31_424971 17 21 . . . 10_1101-2020_12_31_424971 18 1 66 66 CD 10_1101-2020_12_31_424971 18 2 In in IN 10_1101-2020_12_31_424971 18 3 bacteria bacteria NNS 10_1101-2020_12_31_424971 18 4 , , , 10_1101-2020_12_31_424971 18 5 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 18 6 tRNAs trnas CD 10_1101-2020_12_31_424971 18 7 that that WDT 10_1101-2020_12_31_424971 18 8 suppress suppress VBP 10_1101-2020_12_31_424971 18 9 single single JJ 10_1101-2020_12_31_424971 18 10 - - HYPH 10_1101-2020_12_31_424971 18 11 nucleotide nucleotide JJ 10_1101-2020_12_31_424971 18 12 insertion insertion NN 10_1101-2020_12_31_424971 18 13 mutations mutation NNS 10_1101-2020_12_31_424971 18 14 that that IN 10_1101-2020_12_31_424971 18 15 67 67 CD 10_1101-2020_12_31_424971 18 16 shift shift NN 10_1101-2020_12_31_424971 18 17 the the DT 10_1101-2020_12_31_424971 18 18 translational translational JJ 10_1101-2020_12_31_424971 18 19 reading reading NN 10_1101-2020_12_31_424971 18 20 frame frame NN 10_1101-2020_12_31_424971 18 21 to to IN 10_1101-2020_12_31_424971 18 22 the the DT 10_1101-2020_12_31_424971 18 23 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 18 24 have have VBP 10_1101-2020_12_31_424971 18 25 been be VBN 10_1101-2020_12_31_424971 18 26 isolated isolate VBN 10_1101-2020_12_31_424971 18 27 from from IN 10_1101-2020_12_31_424971 18 28 genetic genetic JJ 10_1101-2020_12_31_424971 18 29 studies9,10 studies9,10 NNP 10_1101-2020_12_31_424971 18 30 . . . 10_1101-2020_12_31_424971 19 1 68 68 CD 10_1101-2020_12_31_424971 19 2 These these DT 10_1101-2020_12_31_424971 19 3 +1-frameshifting +1-frameshifting NN 10_1101-2020_12_31_424971 19 4 tRNAs trnas CD 10_1101-2020_12_31_424971 19 5 typically typically RB 10_1101-2020_12_31_424971 19 6 contain contain VBP 10_1101-2020_12_31_424971 19 7 an an DT 10_1101-2020_12_31_424971 19 8 extra extra JJ 10_1101-2020_12_31_424971 19 9 nucleotide nucleotide NN 10_1101-2020_12_31_424971 19 10 in in IN 10_1101-2020_12_31_424971 19 11 the the DT 10_1101-2020_12_31_424971 19 12 anticodon anticodon NNP 10_1101-2020_12_31_424971 19 13 loop loop NN 10_1101-2020_12_31_424971 19 14 – – : 10_1101-2020_12_31_424971 19 15 a a DT 10_1101-2020_12_31_424971 19 16 69 69 CD 10_1101-2020_12_31_424971 19 17 property property NN 10_1101-2020_12_31_424971 19 18 that that WDT 10_1101-2020_12_31_424971 19 19 has have VBZ 10_1101-2020_12_31_424971 19 20 led lead VBN 10_1101-2020_12_31_424971 19 21 to to IN 10_1101-2020_12_31_424971 19 22 the the DT 10_1101-2020_12_31_424971 19 23 proposal proposal NN 10_1101-2020_12_31_424971 19 24 of of IN 10_1101-2020_12_31_424971 19 25 two two CD 10_1101-2020_12_31_424971 19 26 competing compete VBG 10_1101-2020_12_31_424971 19 27 models model NNS 10_1101-2020_12_31_424971 19 28 for for IN 10_1101-2020_12_31_424971 19 29 their -PRON- PRP$ 10_1101-2020_12_31_424971 19 30 mechanism mechanism NN 10_1101-2020_12_31_424971 19 31 of of IN 10_1101-2020_12_31_424971 19 32 action action NN 10_1101-2020_12_31_424971 19 33 . . . 10_1101-2020_12_31_424971 20 1 In in IN 10_1101-2020_12_31_424971 20 2 70 70 CD 10_1101-2020_12_31_424971 20 3 the the DT 10_1101-2020_12_31_424971 20 4 quadruplet quadruplet NN 10_1101-2020_12_31_424971 20 5 - - HYPH 10_1101-2020_12_31_424971 20 6 pairing pair VBG 10_1101-2020_12_31_424971 20 7 model model NN 10_1101-2020_12_31_424971 20 8 , , , 10_1101-2020_12_31_424971 20 9 the the DT 10_1101-2020_12_31_424971 20 10 inserted insert VBN 10_1101-2020_12_31_424971 20 11 nucleotide nucleotide NNP 10_1101-2020_12_31_424971 20 12 joins join VBZ 10_1101-2020_12_31_424971 20 13 the the DT 10_1101-2020_12_31_424971 20 14 triplet triplet NN 10_1101-2020_12_31_424971 20 15 anticodon anticodon NN 10_1101-2020_12_31_424971 20 16 in in IN 10_1101-2020_12_31_424971 20 17 pairing pair VBG 10_1101-2020_12_31_424971 20 18 with with IN 10_1101-2020_12_31_424971 20 19 the the DT 10_1101-2020_12_31_424971 20 20 71 71 CD 10_1101-2020_12_31_424971 20 21 quadruplet quadruplet NN 10_1101-2020_12_31_424971 20 22 codon codon NN 10_1101-2020_12_31_424971 20 23 in in IN 10_1101-2020_12_31_424971 20 24 the the DT 10_1101-2020_12_31_424971 20 25 A A NNP 10_1101-2020_12_31_424971 20 26 site site NN 10_1101-2020_12_31_424971 20 27 and and CC 10_1101-2020_12_31_424971 20 28 this this DT 10_1101-2020_12_31_424971 20 29 quadruplet quadruplet NN 10_1101-2020_12_31_424971 20 30 anticodon anticodon NN 10_1101-2020_12_31_424971 20 31 - - HYPH 10_1101-2020_12_31_424971 20 32 codon codon NNP 10_1101-2020_12_31_424971 20 33 pair pair NN 10_1101-2020_12_31_424971 20 34 is be VBZ 10_1101-2020_12_31_424971 20 35 translocated translocate VBN 10_1101-2020_12_31_424971 20 36 to to IN 10_1101-2020_12_31_424971 20 37 the the DT 10_1101-2020_12_31_424971 20 38 72 72 CD 10_1101-2020_12_31_424971 20 39 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 20 40 peptidyl peptidyl NN 10_1101-2020_12_31_424971 20 41 - - HYPH 10_1101-2020_12_31_424971 20 42 tRNA trna NN 10_1101-2020_12_31_424971 20 43 binding binding NN 10_1101-2020_12_31_424971 20 44 ( ( -LRB- 10_1101-2020_12_31_424971 20 45 P p NN 10_1101-2020_12_31_424971 20 46 ) ) -RRB- 10_1101-2020_12_31_424971 20 47 site11 site11 FW 10_1101-2020_12_31_424971 20 48 . . . 10_1101-2020_12_31_424971 21 1 In in IN 10_1101-2020_12_31_424971 21 2 the the DT 10_1101-2020_12_31_424971 21 3 triplet triplet NN 10_1101-2020_12_31_424971 21 4 - - HYPH 10_1101-2020_12_31_424971 21 5 slippage slippage NN 10_1101-2020_12_31_424971 21 6 model model NN 10_1101-2020_12_31_424971 21 7 , , , 10_1101-2020_12_31_424971 21 8 the the DT 10_1101-2020_12_31_424971 21 9 expanded expand VBN 10_1101-2020_12_31_424971 21 10 anticodon anticodon NNP 10_1101-2020_12_31_424971 21 11 73 73 CD 10_1101-2020_12_31_424971 21 12 loop loop NN 10_1101-2020_12_31_424971 21 13 forms form VBZ 10_1101-2020_12_31_424971 21 14 an an DT 10_1101-2020_12_31_424971 21 15 in in IN 10_1101-2020_12_31_424971 21 16 - - HYPH 10_1101-2020_12_31_424971 21 17 frame frame NN 10_1101-2020_12_31_424971 21 18 ( ( -LRB- 10_1101-2020_12_31_424971 21 19 0-frame 0-frame CD 10_1101-2020_12_31_424971 21 20 ) ) -RRB- 10_1101-2020_12_31_424971 21 21 triplet triplet NN 10_1101-2020_12_31_424971 21 22 anticodon anticodon NN 10_1101-2020_12_31_424971 21 23 - - HYPH 10_1101-2020_12_31_424971 21 24 codon codon NNP 10_1101-2020_12_31_424971 21 25 pair pair NN 10_1101-2020_12_31_424971 21 26 in in IN 10_1101-2020_12_31_424971 21 27 the the DT 10_1101-2020_12_31_424971 21 28 A A NNP 10_1101-2020_12_31_424971 21 29 site site NN 10_1101-2020_12_31_424971 21 30 and and CC 10_1101-2020_12_31_424971 21 31 subsequently subsequently RB 10_1101-2020_12_31_424971 21 32 shifts shift VBZ 10_1101-2020_12_31_424971 21 33 74 74 CD 10_1101-2020_12_31_424971 21 34 .CC .CC , 10_1101-2020_12_31_424971 21 35 - - HYPH 10_1101-2020_12_31_424971 21 36 BY by IN 10_1101-2020_12_31_424971 21 37 - - HYPH 10_1101-2020_12_31_424971 21 38 NC NC NNP 10_1101-2020_12_31_424971 21 39 - - HYPH 10_1101-2020_12_31_424971 21 40 ND ND NNP 10_1101-2020_12_31_424971 21 41 4.0 4.0 CD 10_1101-2020_12_31_424971 21 42 International International NNP 10_1101-2020_12_31_424971 21 43 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 21 44 under under IN 10_1101-2020_12_31_424971 21 45 a a DT 10_1101-2020_12_31_424971 21 46 ( ( -LRB- 10_1101-2020_12_31_424971 21 47 which which WDT 10_1101-2020_12_31_424971 21 48 was be VBD 10_1101-2020_12_31_424971 21 49 not not RB 10_1101-2020_12_31_424971 21 50 certified certify VBN 10_1101-2020_12_31_424971 21 51 by by IN 10_1101-2020_12_31_424971 21 52 peer peer NN 10_1101-2020_12_31_424971 21 53 review review NN 10_1101-2020_12_31_424971 21 54 ) ) -RRB- 10_1101-2020_12_31_424971 21 55 is be VBZ 10_1101-2020_12_31_424971 21 56 the the DT 10_1101-2020_12_31_424971 21 57 author author NN 10_1101-2020_12_31_424971 21 58 / / SYM 10_1101-2020_12_31_424971 21 59 funder funder NN 10_1101-2020_12_31_424971 21 60 , , , 10_1101-2020_12_31_424971 21 61 who who WP 10_1101-2020_12_31_424971 21 62 has have VBZ 10_1101-2020_12_31_424971 21 63 granted grant VBN 10_1101-2020_12_31_424971 21 64 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 21 65 a a DT 10_1101-2020_12_31_424971 21 66 license license NN 10_1101-2020_12_31_424971 21 67 to to TO 10_1101-2020_12_31_424971 21 68 display display VB 10_1101-2020_12_31_424971 21 69 the the DT 10_1101-2020_12_31_424971 21 70 preprint preprint NN 10_1101-2020_12_31_424971 21 71 in in IN 10_1101-2020_12_31_424971 21 72 perpetuity perpetuity NN 10_1101-2020_12_31_424971 21 73 . . . 10_1101-2020_12_31_424971 22 1 It -PRON- PRP 10_1101-2020_12_31_424971 22 2 is be VBZ 10_1101-2020_12_31_424971 22 3 made make VBN 10_1101-2020_12_31_424971 22 4 The the DT 10_1101-2020_12_31_424971 22 5 copyright copyright NN 10_1101-2020_12_31_424971 22 6 holder holder NN 10_1101-2020_12_31_424971 22 7 for for IN 10_1101-2020_12_31_424971 22 8 this this DT 10_1101-2020_12_31_424971 22 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 22 10 version version NN 10_1101-2020_12_31_424971 22 11 posted post VBD 10_1101-2020_12_31_424971 22 12 January January NNP 10_1101-2020_12_31_424971 22 13 2 2 CD 10_1101-2020_12_31_424971 22 14 , , , 10_1101-2020_12_31_424971 22 15 2021 2021 CD 10_1101-2020_12_31_424971 22 16 . . . 10_1101-2020_12_31_424971 22 17 ; ; : 10_1101-2020_12_31_424971 22 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 22 19 : : : 10_1101-2020_12_31_424971 22 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 22 21 preprint preprint NN 10_1101-2020_12_31_424971 22 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 22 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 22 24 4 4 CD 10_1101-2020_12_31_424971 22 25 to to IN 10_1101-2020_12_31_424971 22 26 the the DT 10_1101-2020_12_31_424971 22 27 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 22 28 at at IN 10_1101-2020_12_31_424971 22 29 some some DT 10_1101-2020_12_31_424971 22 30 point point NN 10_1101-2020_12_31_424971 22 31 later later RB 10_1101-2020_12_31_424971 22 32 in in IN 10_1101-2020_12_31_424971 22 33 the the DT 10_1101-2020_12_31_424971 22 34 elongation elongation NN 10_1101-2020_12_31_424971 22 35 cycle12,13 cycle12,13 NNP 10_1101-2020_12_31_424971 22 36 , , , 10_1101-2020_12_31_424971 22 37 possibly possibly RB 10_1101-2020_12_31_424971 22 38 during during IN 10_1101-2020_12_31_424971 22 39 translocation translocation NN 10_1101-2020_12_31_424971 22 40 of of IN 10_1101-2020_12_31_424971 22 41 the the DT 10_1101-2020_12_31_424971 22 42 75 75 CD 10_1101-2020_12_31_424971 22 43 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 22 44 tRNA trna NN 10_1101-2020_12_31_424971 22 45 from from IN 10_1101-2020_12_31_424971 22 46 the the DT 10_1101-2020_12_31_424971 22 47 A a NN 10_1101-2020_12_31_424971 22 48 to to IN 10_1101-2020_12_31_424971 22 49 P p NN 10_1101-2020_12_31_424971 22 50 sites14 sites14 NNS 10_1101-2020_12_31_424971 22 51 or or CC 10_1101-2020_12_31_424971 22 52 within within IN 10_1101-2020_12_31_424971 22 53 the the DT 10_1101-2020_12_31_424971 22 54 P p NN 10_1101-2020_12_31_424971 22 55 site15 site15 NN 10_1101-2020_12_31_424971 22 56 . . . 10_1101-2020_12_31_424971 23 1 The the DT 10_1101-2020_12_31_424971 23 2 triplet triplet NN 10_1101-2020_12_31_424971 23 3 - - HYPH 10_1101-2020_12_31_424971 23 4 slippage slippage NN 10_1101-2020_12_31_424971 23 5 model model NN 10_1101-2020_12_31_424971 23 6 is be VBZ 10_1101-2020_12_31_424971 23 7 76 76 CD 10_1101-2020_12_31_424971 23 8 supported support VBN 10_1101-2020_12_31_424971 23 9 by by IN 10_1101-2020_12_31_424971 23 10 structural structural JJ 10_1101-2020_12_31_424971 23 11 studies study NNS 10_1101-2020_12_31_424971 23 12 of of IN 10_1101-2020_12_31_424971 23 13 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 23 14 complexes complex NNS 10_1101-2020_12_31_424971 23 15 in in IN 10_1101-2020_12_31_424971 23 16 which which WDT 10_1101-2020_12_31_424971 23 17 the the DT 10_1101-2020_12_31_424971 23 18 expanded expand VBN 10_1101-2020_12_31_424971 23 19 anticodon anticodon NNP 10_1101-2020_12_31_424971 23 20 - - HYPH 10_1101-2020_12_31_424971 23 21 stem-77 stem-77 NN 10_1101-2020_12_31_424971 23 22 loops loop NNS 10_1101-2020_12_31_424971 23 23 ( ( -LRB- 10_1101-2020_12_31_424971 23 24 ASLs ASLs NNP 10_1101-2020_12_31_424971 23 25 ) ) -RRB- 10_1101-2020_12_31_424971 23 26 of of IN 10_1101-2020_12_31_424971 23 27 +1-frameshifting +1-frameshifting NN 10_1101-2020_12_31_424971 23 28 tRNAs trnas CD 10_1101-2020_12_31_424971 23 29 have have VBP 10_1101-2020_12_31_424971 23 30 been be VBN 10_1101-2020_12_31_424971 23 31 found find VBN 10_1101-2020_12_31_424971 23 32 to to TO 10_1101-2020_12_31_424971 23 33 use use VB 10_1101-2020_12_31_424971 23 34 triplet triplet NN 10_1101-2020_12_31_424971 23 35 anticodon anticodon NN 10_1101-2020_12_31_424971 23 36 - - HYPH 10_1101-2020_12_31_424971 23 37 codon codon NNP 10_1101-2020_12_31_424971 23 38 pairing pair VBG 10_1101-2020_12_31_424971 23 39 78 78 CD 10_1101-2020_12_31_424971 23 40 in in IN 10_1101-2020_12_31_424971 23 41 the the DT 10_1101-2020_12_31_424971 23 42 0-frame 0-frame CD 10_1101-2020_12_31_424971 23 43 at at IN 10_1101-2020_12_31_424971 23 44 the the DT 10_1101-2020_12_31_424971 23 45 A A NNP 10_1101-2020_12_31_424971 23 46 site16 site16 NNP 10_1101-2020_12_31_424971 23 47 - - HYPH 10_1101-2020_12_31_424971 23 48 18 18 CD 10_1101-2020_12_31_424971 23 49 and and CC 10_1101-2020_12_31_424971 23 50 in in IN 10_1101-2020_12_31_424971 23 51 the the DT 10_1101-2020_12_31_424971 23 52 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 23 53 at at IN 10_1101-2020_12_31_424971 23 54 the the DT 10_1101-2020_12_31_424971 23 55 P p NN 10_1101-2020_12_31_424971 23 56 site19 site19 NN 10_1101-2020_12_31_424971 23 57 . . . 10_1101-2020_12_31_424971 24 1 Nonetheless nonetheless RB 10_1101-2020_12_31_424971 24 2 , , , 10_1101-2020_12_31_424971 24 3 these these DT 10_1101-2020_12_31_424971 24 4 structures structure NNS 10_1101-2020_12_31_424971 24 5 79 79 CD 10_1101-2020_12_31_424971 24 6 do do VBP 10_1101-2020_12_31_424971 24 7 not not RB 10_1101-2020_12_31_424971 24 8 eliminate eliminate VB 10_1101-2020_12_31_424971 24 9 the the DT 10_1101-2020_12_31_424971 24 10 possibility possibility NN 10_1101-2020_12_31_424971 24 11 that that IN 10_1101-2020_12_31_424971 24 12 two two CD 10_1101-2020_12_31_424971 24 13 competing compete VBG 10_1101-2020_12_31_424971 24 14 triplet triplet NN 10_1101-2020_12_31_424971 24 15 pairing pairing NN 10_1101-2020_12_31_424971 24 16 schemes scheme NNS 10_1101-2020_12_31_424971 24 17 ( ( -LRB- 10_1101-2020_12_31_424971 24 18 0-frame 0-frame CD 10_1101-2020_12_31_424971 24 19 and and CC 10_1101-2020_12_31_424971 24 20 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 24 21 ) ) -RRB- 10_1101-2020_12_31_424971 24 22 80 80 CD 10_1101-2020_12_31_424971 24 23 can can MD 10_1101-2020_12_31_424971 24 24 co co VB 10_1101-2020_12_31_424971 24 25 - - VB 10_1101-2020_12_31_424971 24 26 exist exist VB 10_1101-2020_12_31_424971 24 27 when when WRB 10_1101-2020_12_31_424971 24 28 a a DT 10_1101-2020_12_31_424971 24 29 quadruplet quadruplet NN 10_1101-2020_12_31_424971 24 30 codon codon NN 10_1101-2020_12_31_424971 24 31 motif motif NN 10_1101-2020_12_31_424971 24 32 occupies occupy VBZ 10_1101-2020_12_31_424971 24 33 the the DT 10_1101-2020_12_31_424971 24 34 A a NN 10_1101-2020_12_31_424971 24 35 site15 site15 NN 10_1101-2020_12_31_424971 24 36 , , , 10_1101-2020_12_31_424971 24 37 that that IN 10_1101-2020_12_31_424971 24 38 some some DT 10_1101-2020_12_31_424971 24 39 amount amount NN 10_1101-2020_12_31_424971 24 40 of of IN 10_1101-2020_12_31_424971 24 41 +1 +1 NNS 10_1101-2020_12_31_424971 24 42 81 81 CD 10_1101-2020_12_31_424971 24 43 frameshifting frameshifting NN 10_1101-2020_12_31_424971 24 44 can can MD 10_1101-2020_12_31_424971 24 45 occur occur VB 10_1101-2020_12_31_424971 24 46 via via IN 10_1101-2020_12_31_424971 24 47 the the DT 10_1101-2020_12_31_424971 24 48 quadruplet quadruplet NN 10_1101-2020_12_31_424971 24 49 - - HYPH 10_1101-2020_12_31_424971 24 50 pairing pair VBG 10_1101-2020_12_31_424971 24 51 model model NN 10_1101-2020_12_31_424971 24 52 , , , 10_1101-2020_12_31_424971 24 53 and and CC 10_1101-2020_12_31_424971 24 54 that that IN 10_1101-2020_12_31_424971 24 55 the the DT 10_1101-2020_12_31_424971 24 56 quadruplet quadruplet NN 10_1101-2020_12_31_424971 24 57 - - HYPH 10_1101-2020_12_31_424971 24 58 pairing pair VBG 10_1101-2020_12_31_424971 24 59 model model NN 10_1101-2020_12_31_424971 24 60 82 82 CD 10_1101-2020_12_31_424971 24 61 may may MD 10_1101-2020_12_31_424971 24 62 even even RB 10_1101-2020_12_31_424971 24 63 dominate dominate VB 10_1101-2020_12_31_424971 24 64 for for IN 10_1101-2020_12_31_424971 24 65 particular particular JJ 10_1101-2020_12_31_424971 24 66 +1-frameshifting +1-frameshifting NN 10_1101-2020_12_31_424971 24 67 tRNAs trna NNS 10_1101-2020_12_31_424971 24 68 , , , 10_1101-2020_12_31_424971 24 69 codon codon NNP 10_1101-2020_12_31_424971 24 70 sequences sequences NNP 10_1101-2020_12_31_424971 24 71 , , , 10_1101-2020_12_31_424971 24 72 and/or and/or CC 10_1101-2020_12_31_424971 24 73 reaction reaction NN 10_1101-2020_12_31_424971 24 74 83 83 CD 10_1101-2020_12_31_424971 24 75 conditions10 conditions10 NN 10_1101-2020_12_31_424971 24 76 . . . 10_1101-2020_12_31_424971 25 1 We -PRON- PRP 10_1101-2020_12_31_424971 25 2 also also RB 10_1101-2020_12_31_424971 25 3 do do VBP 10_1101-2020_12_31_424971 25 4 not not RB 10_1101-2020_12_31_424971 25 5 know know VB 10_1101-2020_12_31_424971 25 6 how how WRB 10_1101-2020_12_31_424971 25 7 each each DT 10_1101-2020_12_31_424971 25 8 model model NN 10_1101-2020_12_31_424971 25 9 determines determine VBZ 10_1101-2020_12_31_424971 25 10 the the DT 10_1101-2020_12_31_424971 25 11 efficiency efficiency NN 10_1101-2020_12_31_424971 25 12 of of IN 10_1101-2020_12_31_424971 25 13 +1 +1 NNS 10_1101-2020_12_31_424971 25 14 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 25 15 84 84 CD 10_1101-2020_12_31_424971 25 16 or or CC 10_1101-2020_12_31_424971 25 17 whether whether IN 10_1101-2020_12_31_424971 25 18 any any DT 10_1101-2020_12_31_424971 25 19 competition competition NN 10_1101-2020_12_31_424971 25 20 between between IN 10_1101-2020_12_31_424971 25 21 the the DT 10_1101-2020_12_31_424971 25 22 two two CD 10_1101-2020_12_31_424971 25 23 models model NNS 10_1101-2020_12_31_424971 25 24 is be VBZ 10_1101-2020_12_31_424971 25 25 driven drive VBN 10_1101-2020_12_31_424971 25 26 by by IN 10_1101-2020_12_31_424971 25 27 the the DT 10_1101-2020_12_31_424971 25 28 kinetics kinetic NNS 10_1101-2020_12_31_424971 25 29 of of IN 10_1101-2020_12_31_424971 25 30 frameshifting frameshifting NN 10_1101-2020_12_31_424971 25 31 or or CC 10_1101-2020_12_31_424971 25 32 85 85 CD 10_1101-2020_12_31_424971 25 33 the the DT 10_1101-2020_12_31_424971 25 34 thermodynamics thermodynamic NNS 10_1101-2020_12_31_424971 25 35 of of IN 10_1101-2020_12_31_424971 25 36 base base NN 10_1101-2020_12_31_424971 25 37 pairing pairing NN 10_1101-2020_12_31_424971 25 38 . . . 10_1101-2020_12_31_424971 26 1 In in IN 10_1101-2020_12_31_424971 26 2 addition addition NN 10_1101-2020_12_31_424971 26 3 , , , 10_1101-2020_12_31_424971 26 4 virtually virtually RB 10_1101-2020_12_31_424971 26 5 all all DT 10_1101-2020_12_31_424971 26 6 natural natural JJ 10_1101-2020_12_31_424971 26 7 tRNAs trnas CD 10_1101-2020_12_31_424971 26 8 contain contain VB 10_1101-2020_12_31_424971 26 9 a a DT 10_1101-2020_12_31_424971 26 10 purine purine NN 10_1101-2020_12_31_424971 26 11 at at IN 10_1101-2020_12_31_424971 26 12 86 86 CD 10_1101-2020_12_31_424971 26 13 nucleotide nucleotide JJ 10_1101-2020_12_31_424971 26 14 position position NN 10_1101-2020_12_31_424971 26 15 37 37 CD 10_1101-2020_12_31_424971 26 16 on on IN 10_1101-2020_12_31_424971 26 17 the the DT 10_1101-2020_12_31_424971 26 18 3'-side 3'-side NNP 10_1101-2020_12_31_424971 26 19 of of IN 10_1101-2020_12_31_424971 26 20 the the DT 10_1101-2020_12_31_424971 26 21 anticodon anticodon NN 10_1101-2020_12_31_424971 26 22 ( ( -LRB- 10_1101-2020_12_31_424971 26 23 http://trna.bioinf.uni-leipzig.de/ http://trna.bioinf.uni-leipzig.de/ NNP 10_1101-2020_12_31_424971 26 24 ) ) -RRB- 10_1101-2020_12_31_424971 26 25 , , , 10_1101-2020_12_31_424971 26 26 which which WDT 10_1101-2020_12_31_424971 26 27 is be VBZ 10_1101-2020_12_31_424971 26 28 87 87 CD 10_1101-2020_12_31_424971 26 29 invariably invariably RB 10_1101-2020_12_31_424971 26 30 post post JJ 10_1101-2020_12_31_424971 26 31 - - JJ 10_1101-2020_12_31_424971 26 32 transcriptionally transcriptionally RB 10_1101-2020_12_31_424971 26 33 modified modify VBN 10_1101-2020_12_31_424971 26 34 and and CC 10_1101-2020_12_31_424971 26 35 is be VBZ 10_1101-2020_12_31_424971 26 36 important important JJ 10_1101-2020_12_31_424971 26 37 for for IN 10_1101-2020_12_31_424971 26 38 maintaining maintain VBG 10_1101-2020_12_31_424971 26 39 the the DT 10_1101-2020_12_31_424971 26 40 translational translational JJ 10_1101-2020_12_31_424971 26 41 reading reading NN 10_1101-2020_12_31_424971 26 42 88 88 CD 10_1101-2020_12_31_424971 26 43 frame frame NN 10_1101-2020_12_31_424971 26 44 in in IN 10_1101-2020_12_31_424971 26 45 the the DT 10_1101-2020_12_31_424971 26 46 P p NN 10_1101-2020_12_31_424971 26 47 site15 site15 NN 10_1101-2020_12_31_424971 26 48 . . . 10_1101-2020_12_31_424971 27 1 While while IN 10_1101-2020_12_31_424971 27 2 most most JJS 10_1101-2020_12_31_424971 27 3 +1-frameshifting +1-frameshifting JJ 10_1101-2020_12_31_424971 27 4 tRNAs trnas CD 10_1101-2020_12_31_424971 27 5 sequenced sequence VBN 10_1101-2020_12_31_424971 27 6 to to IN 10_1101-2020_12_31_424971 27 7 date date NN 10_1101-2020_12_31_424971 27 8 also also RB 10_1101-2020_12_31_424971 27 9 contain contain VBP 10_1101-2020_12_31_424971 27 10 a a DT 10_1101-2020_12_31_424971 27 11 purine purine NN 10_1101-2020_12_31_424971 27 12 89 89 CD 10_1101-2020_12_31_424971 27 13 nucleotide nucleotide RB 10_1101-2020_12_31_424971 27 14 at at IN 10_1101-2020_12_31_424971 27 15 position position NN 10_1101-2020_12_31_424971 27 16 378 378 CD 10_1101-2020_12_31_424971 27 17 , , , 10_1101-2020_12_31_424971 27 18 we -PRON- PRP 10_1101-2020_12_31_424971 27 19 do do VBP 10_1101-2020_12_31_424971 27 20 not not RB 10_1101-2020_12_31_424971 27 21 know know VB 10_1101-2020_12_31_424971 27 22 whether whether IN 10_1101-2020_12_31_424971 27 23 it -PRON- PRP 10_1101-2020_12_31_424971 27 24 is be VBZ 10_1101-2020_12_31_424971 27 25 post post JJ 10_1101-2020_12_31_424971 27 26 - - JJ 10_1101-2020_12_31_424971 27 27 transcriptionally transcriptionally RB 10_1101-2020_12_31_424971 27 28 modified modify VBN 10_1101-2020_12_31_424971 27 29 or or CC 10_1101-2020_12_31_424971 27 30 how how WRB 10_1101-2020_12_31_424971 27 31 the the DT 10_1101-2020_12_31_424971 27 32 90 90 CD 10_1101-2020_12_31_424971 27 33 modification modification NN 10_1101-2020_12_31_424971 27 34 affects affect VBZ 10_1101-2020_12_31_424971 27 35 +1 +1 : 10_1101-2020_12_31_424971 27 36 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 27 37 . . . 10_1101-2020_12_31_424971 28 1 Perhaps perhaps RB 10_1101-2020_12_31_424971 28 2 most most RBS 10_1101-2020_12_31_424971 28 3 importantly importantly RB 10_1101-2020_12_31_424971 28 4 , , , 10_1101-2020_12_31_424971 28 5 while while IN 10_1101-2020_12_31_424971 28 6 the the DT 10_1101-2020_12_31_424971 28 7 structural structural JJ 10_1101-2020_12_31_424971 28 8 studies study NNS 10_1101-2020_12_31_424971 28 9 91 91 CD 10_1101-2020_12_31_424971 28 10 described describe VBN 10_1101-2020_12_31_424971 28 11 above above IN 10_1101-2020_12_31_424971 28 12 provide provide VB 10_1101-2020_12_31_424971 28 13 snapshots snapshot NNS 10_1101-2020_12_31_424971 28 14 of of IN 10_1101-2020_12_31_424971 28 15 the the DT 10_1101-2020_12_31_424971 28 16 initial initial JJ 10_1101-2020_12_31_424971 28 17 and and CC 10_1101-2020_12_31_424971 28 18 final final JJ 10_1101-2020_12_31_424971 28 19 states state NNS 10_1101-2020_12_31_424971 28 20 of of IN 10_1101-2020_12_31_424971 28 21 +1 +1 NNS 10_1101-2020_12_31_424971 28 22 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 28 23 , , , 10_1101-2020_12_31_424971 28 24 they -PRON- PRP 10_1101-2020_12_31_424971 28 25 do do VBP 10_1101-2020_12_31_424971 28 26 not not RB 10_1101-2020_12_31_424971 28 27 92 92 CD 10_1101-2020_12_31_424971 28 28 reveal reveal VB 10_1101-2020_12_31_424971 28 29 where where WRB 10_1101-2020_12_31_424971 28 30 , , , 10_1101-2020_12_31_424971 28 31 when when WRB 10_1101-2020_12_31_424971 28 32 , , , 10_1101-2020_12_31_424971 28 33 or or CC 10_1101-2020_12_31_424971 28 34 how how WRB 10_1101-2020_12_31_424971 28 35 the the DT 10_1101-2020_12_31_424971 28 36 shift shift NN 10_1101-2020_12_31_424971 28 37 occurs occur VBZ 10_1101-2020_12_31_424971 28 38 , , , 10_1101-2020_12_31_424971 28 39 thereby thereby RB 10_1101-2020_12_31_424971 28 40 precluding preclude VBG 10_1101-2020_12_31_424971 28 41 an an DT 10_1101-2020_12_31_424971 28 42 understanding understanding NN 10_1101-2020_12_31_424971 28 43 of of IN 10_1101-2020_12_31_424971 28 44 the the DT 10_1101-2020_12_31_424971 28 45 structural structural JJ 10_1101-2020_12_31_424971 28 46 93 93 CD 10_1101-2020_12_31_424971 28 47 basis basis NN 10_1101-2020_12_31_424971 28 48 and and CC 10_1101-2020_12_31_424971 28 49 mechanism mechanism NN 10_1101-2020_12_31_424971 28 50 of of IN 10_1101-2020_12_31_424971 28 51 +1 +1 NNS 10_1101-2020_12_31_424971 28 52 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 28 53 . . . 10_1101-2020_12_31_424971 29 1 These these DT 10_1101-2020_12_31_424971 29 2 open open JJ 10_1101-2020_12_31_424971 29 3 questions question NNS 10_1101-2020_12_31_424971 29 4 have have VBP 10_1101-2020_12_31_424971 29 5 limited limit VBN 10_1101-2020_12_31_424971 29 6 our -PRON- PRP$ 10_1101-2020_12_31_424971 29 7 ability ability NN 10_1101-2020_12_31_424971 29 8 to to IN 10_1101-2020_12_31_424971 29 9 94 94 CD 10_1101-2020_12_31_424971 29 10 increase increase VB 10_1101-2020_12_31_424971 29 11 the the DT 10_1101-2020_12_31_424971 29 12 efficiency efficiency NN 10_1101-2020_12_31_424971 29 13 of of IN 10_1101-2020_12_31_424971 29 14 genome genome JJ 10_1101-2020_12_31_424971 29 15 recoding recode VBG 10_1101-2020_12_31_424971 29 16 at at IN 10_1101-2020_12_31_424971 29 17 quadruplet quadruplet NN 10_1101-2020_12_31_424971 29 18 codons codon NNS 10_1101-2020_12_31_424971 29 19 . . . 10_1101-2020_12_31_424971 30 1 95 95 CD 10_1101-2020_12_31_424971 30 2 To to TO 10_1101-2020_12_31_424971 30 3 address address VB 10_1101-2020_12_31_424971 30 4 these these DT 10_1101-2020_12_31_424971 30 5 questions question NNS 10_1101-2020_12_31_424971 30 6 , , , 10_1101-2020_12_31_424971 30 7 we -PRON- PRP 10_1101-2020_12_31_424971 30 8 have have VBP 10_1101-2020_12_31_424971 30 9 investigated investigate VBN 10_1101-2020_12_31_424971 30 10 the the DT 10_1101-2020_12_31_424971 30 11 mechanism mechanism NN 10_1101-2020_12_31_424971 30 12 of of IN 10_1101-2020_12_31_424971 30 13 +1 +1 NNS 10_1101-2020_12_31_424971 30 14 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 30 15 by by IN 10_1101-2020_12_31_424971 30 16 96 96 CD 10_1101-2020_12_31_424971 30 17 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 30 18 ( ( -LRB- 10_1101-2020_12_31_424971 30 19 Figure Figure NNP 10_1101-2020_12_31_424971 30 20 1a 1a NNP 10_1101-2020_12_31_424971 30 21 ) ) -RRB- 10_1101-2020_12_31_424971 30 22 , , , 10_1101-2020_12_31_424971 30 23 a a DT 10_1101-2020_12_31_424971 30 24 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 30 25 tRNA trna NN 10_1101-2020_12_31_424971 30 26 that that WDT 10_1101-2020_12_31_424971 30 27 was be VBD 10_1101-2020_12_31_424971 30 28 isolated isolate VBN 10_1101-2020_12_31_424971 30 29 from from IN 10_1101-2020_12_31_424971 30 30 Salmonella Salmonella NNP 10_1101-2020_12_31_424971 30 31 typhimurium typhimurium NN 10_1101-2020_12_31_424971 30 32 as as IN 10_1101-2020_12_31_424971 30 33 a a DT 10_1101-2020_12_31_424971 30 34 97 97 CD 10_1101-2020_12_31_424971 30 35 suppressor suppressor NN 10_1101-2020_12_31_424971 30 36 of of IN 10_1101-2020_12_31_424971 30 37 a a DT 10_1101-2020_12_31_424971 30 38 single single JJ 10_1101-2020_12_31_424971 30 39 C C NNP 10_1101-2020_12_31_424971 30 40 insertion insertion NN 10_1101-2020_12_31_424971 30 41 into into IN 10_1101-2020_12_31_424971 30 42 a a DT 10_1101-2020_12_31_424971 30 43 proline proline NN 10_1101-2020_12_31_424971 30 44 ( ( -LRB- 10_1101-2020_12_31_424971 30 45 Pro Pro NNP 10_1101-2020_12_31_424971 30 46 ) ) -RRB- 10_1101-2020_12_31_424971 30 47 CCC CCC NNP 10_1101-2020_12_31_424971 30 48 codon20 codon20 NNP 10_1101-2020_12_31_424971 30 49 . . . 10_1101-2020_12_31_424971 31 1 The the DT 10_1101-2020_12_31_424971 31 2 observed observed JJ 10_1101-2020_12_31_424971 31 3 high high JJ 10_1101-2020_12_31_424971 31 4 +1 +1 NNS 10_1101-2020_12_31_424971 31 5 - - SYM 10_1101-2020_12_31_424971 31 6 98 98 CD 10_1101-2020_12_31_424971 31 7 frameshifting frameshifting NN 10_1101-2020_12_31_424971 31 8 efficiency efficiency NN 10_1101-2020_12_31_424971 31 9 of of IN 10_1101-2020_12_31_424971 31 10 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 31 11 at at IN 10_1101-2020_12_31_424971 31 12 the the DT 10_1101-2020_12_31_424971 31 13 CCC CCC NNP 10_1101-2020_12_31_424971 31 14 - - HYPH 10_1101-2020_12_31_424971 31 15 C C NNP 10_1101-2020_12_31_424971 31 16 motif motif NN 10_1101-2020_12_31_424971 31 17 , , , 10_1101-2020_12_31_424971 31 18 nearly nearly RB 10_1101-2020_12_31_424971 31 19 80-fold 80-fold CD 10_1101-2020_12_31_424971 31 20 above above IN 10_1101-2020_12_31_424971 31 21 background20 background20 NNP 10_1101-2020_12_31_424971 31 22 , , , 10_1101-2020_12_31_424971 31 23 99 99 CD 10_1101-2020_12_31_424971 31 24 demonstrates demonstrate VBZ 10_1101-2020_12_31_424971 31 25 its -PRON- PRP$ 10_1101-2020_12_31_424971 31 26 ability ability NN 10_1101-2020_12_31_424971 31 27 to to TO 10_1101-2020_12_31_424971 31 28 successfully successfully RB 10_1101-2020_12_31_424971 31 29 compete compete VB 10_1101-2020_12_31_424971 31 30 with with IN 10_1101-2020_12_31_424971 31 31 the the DT 10_1101-2020_12_31_424971 31 32 naturally naturally RB 10_1101-2020_12_31_424971 31 33 occurring occur VBG 10_1101-2020_12_31_424971 31 34 ProL ProL NNP 10_1101-2020_12_31_424971 31 35 and and CC 10_1101-2020_12_31_424971 31 36 ProM ProM NNP 10_1101-2020_12_31_424971 31 37 100 100 CD 10_1101-2020_12_31_424971 31 38 .CC .CC : 10_1101-2020_12_31_424971 31 39 - - HYPH 10_1101-2020_12_31_424971 31 40 BY by IN 10_1101-2020_12_31_424971 31 41 - - HYPH 10_1101-2020_12_31_424971 31 42 NC NC NNP 10_1101-2020_12_31_424971 31 43 - - HYPH 10_1101-2020_12_31_424971 31 44 ND ND NNP 10_1101-2020_12_31_424971 31 45 4.0 4.0 CD 10_1101-2020_12_31_424971 31 46 International International NNP 10_1101-2020_12_31_424971 31 47 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 31 48 under under IN 10_1101-2020_12_31_424971 31 49 a a DT 10_1101-2020_12_31_424971 31 50 ( ( -LRB- 10_1101-2020_12_31_424971 31 51 which which WDT 10_1101-2020_12_31_424971 31 52 was be VBD 10_1101-2020_12_31_424971 31 53 not not RB 10_1101-2020_12_31_424971 31 54 certified certify VBN 10_1101-2020_12_31_424971 31 55 by by IN 10_1101-2020_12_31_424971 31 56 peer peer NN 10_1101-2020_12_31_424971 31 57 review review NN 10_1101-2020_12_31_424971 31 58 ) ) -RRB- 10_1101-2020_12_31_424971 31 59 is be VBZ 10_1101-2020_12_31_424971 31 60 the the DT 10_1101-2020_12_31_424971 31 61 author author NN 10_1101-2020_12_31_424971 31 62 / / SYM 10_1101-2020_12_31_424971 31 63 funder funder NN 10_1101-2020_12_31_424971 31 64 , , , 10_1101-2020_12_31_424971 31 65 who who WP 10_1101-2020_12_31_424971 31 66 has have VBZ 10_1101-2020_12_31_424971 31 67 granted grant VBN 10_1101-2020_12_31_424971 31 68 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 31 69 a a DT 10_1101-2020_12_31_424971 31 70 license license NN 10_1101-2020_12_31_424971 31 71 to to TO 10_1101-2020_12_31_424971 31 72 display display VB 10_1101-2020_12_31_424971 31 73 the the DT 10_1101-2020_12_31_424971 31 74 preprint preprint NN 10_1101-2020_12_31_424971 31 75 in in IN 10_1101-2020_12_31_424971 31 76 perpetuity perpetuity NN 10_1101-2020_12_31_424971 31 77 . . . 10_1101-2020_12_31_424971 32 1 It -PRON- PRP 10_1101-2020_12_31_424971 32 2 is be VBZ 10_1101-2020_12_31_424971 32 3 made make VBN 10_1101-2020_12_31_424971 32 4 The the DT 10_1101-2020_12_31_424971 32 5 copyright copyright NN 10_1101-2020_12_31_424971 32 6 holder holder NN 10_1101-2020_12_31_424971 32 7 for for IN 10_1101-2020_12_31_424971 32 8 this this DT 10_1101-2020_12_31_424971 32 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 32 10 version version NN 10_1101-2020_12_31_424971 32 11 posted post VBD 10_1101-2020_12_31_424971 32 12 January January NNP 10_1101-2020_12_31_424971 32 13 2 2 CD 10_1101-2020_12_31_424971 32 14 , , , 10_1101-2020_12_31_424971 32 15 2021 2021 CD 10_1101-2020_12_31_424971 32 16 . . . 10_1101-2020_12_31_424971 32 17 ; ; : 10_1101-2020_12_31_424971 32 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 32 19 : : : 10_1101-2020_12_31_424971 32 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 32 21 preprint preprint NN 10_1101-2020_12_31_424971 32 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 32 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 32 24 5 5 CD 10_1101-2020_12_31_424971 32 25 isoacceptor isoacceptor NN 10_1101-2020_12_31_424971 32 26 tRNAs trnas CD 10_1101-2020_12_31_424971 32 27 that that WDT 10_1101-2020_12_31_424971 32 28 read read VBP 10_1101-2020_12_31_424971 32 29 the the DT 10_1101-2020_12_31_424971 32 30 CCC CCC NNP 10_1101-2020_12_31_424971 32 31 codon codon NN 10_1101-2020_12_31_424971 32 32 . . . 10_1101-2020_12_31_424971 33 1 Using use VBG 10_1101-2020_12_31_424971 33 2 the the DT 10_1101-2020_12_31_424971 33 3 ensemble ensemble JJ 10_1101-2020_12_31_424971 33 4 ‘ ' `` 10_1101-2020_12_31_424971 33 5 codon codon NN 10_1101-2020_12_31_424971 33 6 - - HYPH 10_1101-2020_12_31_424971 33 7 walk walk NN 10_1101-2020_12_31_424971 33 8 ’ ' '' 10_1101-2020_12_31_424971 33 9 methodology21 methodology21 CD 10_1101-2020_12_31_424971 33 10 101 101 CD 10_1101-2020_12_31_424971 33 11 and and CC 10_1101-2020_12_31_424971 33 12 single single JJ 10_1101-2020_12_31_424971 33 13 - - HYPH 10_1101-2020_12_31_424971 33 14 molecule molecule NN 10_1101-2020_12_31_424971 33 15 fluorescence fluorescence NN 10_1101-2020_12_31_424971 33 16 resonance resonance NN 10_1101-2020_12_31_424971 33 17 energy energy NN 10_1101-2020_12_31_424971 33 18 transfer transfer NN 10_1101-2020_12_31_424971 33 19 ( ( -LRB- 10_1101-2020_12_31_424971 33 20 smFRET smFRET NNP 10_1101-2020_12_31_424971 33 21 ) ) -RRB- 10_1101-2020_12_31_424971 33 22 , , , 10_1101-2020_12_31_424971 33 23 we -PRON- PRP 10_1101-2020_12_31_424971 33 24 have have VBP 10_1101-2020_12_31_424971 33 25 compared compare VBN 10_1101-2020_12_31_424971 33 26 the the DT 10_1101-2020_12_31_424971 33 27 102 102 CD 10_1101-2020_12_31_424971 33 28 +1 +1 NNS 10_1101-2020_12_31_424971 33 29 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 33 30 activity activity NN 10_1101-2020_12_31_424971 33 31 of of IN 10_1101-2020_12_31_424971 33 32 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 33 33 relative relative JJ 10_1101-2020_12_31_424971 33 34 to to IN 10_1101-2020_12_31_424971 33 35 its -PRON- PRP$ 10_1101-2020_12_31_424971 33 36 closest close JJS 10_1101-2020_12_31_424971 33 37 counterpart counterpart NN 10_1101-2020_12_31_424971 33 38 , , , 10_1101-2020_12_31_424971 33 39 ProL ProL NNP 10_1101-2020_12_31_424971 33 40 , , , 10_1101-2020_12_31_424971 33 41 at at IN 10_1101-2020_12_31_424971 33 42 a a DT 10_1101-2020_12_31_424971 33 43 CCC CCC NNP 10_1101-2020_12_31_424971 33 44 - - HYPH 10_1101-2020_12_31_424971 33 45 C C NNP 10_1101-2020_12_31_424971 33 46 motif motif NN 10_1101-2020_12_31_424971 33 47 , , , 10_1101-2020_12_31_424971 33 48 and and CC 10_1101-2020_12_31_424971 33 49 103 103 CD 10_1101-2020_12_31_424971 33 50 determined determine VBD 10_1101-2020_12_31_424971 33 51 the the DT 10_1101-2020_12_31_424971 33 52 position position NN 10_1101-2020_12_31_424971 33 53 and and CC 10_1101-2020_12_31_424971 33 54 timing timing NN 10_1101-2020_12_31_424971 33 55 of of IN 10_1101-2020_12_31_424971 33 56 the the DT 10_1101-2020_12_31_424971 33 57 shift shift NN 10_1101-2020_12_31_424971 33 58 . . . 10_1101-2020_12_31_424971 34 1 Our -PRON- PRP$ 10_1101-2020_12_31_424971 34 2 results result NNS 10_1101-2020_12_31_424971 34 3 show show VBP 10_1101-2020_12_31_424971 34 4 that that IN 10_1101-2020_12_31_424971 34 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 34 6 is be VBZ 10_1101-2020_12_31_424971 34 7 naturally naturally RB 10_1101-2020_12_31_424971 34 8 N1 N1 NNP 10_1101-2020_12_31_424971 34 9 - - HYPH 10_1101-2020_12_31_424971 34 10 104 104 NNP 10_1101-2020_12_31_424971 34 11 methylated methylate VBD 10_1101-2020_12_31_424971 34 12 at at IN 10_1101-2020_12_31_424971 34 13 G37 G37 NNP 10_1101-2020_12_31_424971 34 14 in in IN 10_1101-2020_12_31_424971 34 15 cells cell NNS 10_1101-2020_12_31_424971 34 16 , , , 10_1101-2020_12_31_424971 34 17 generating generate VBG 10_1101-2020_12_31_424971 34 18 an an DT 10_1101-2020_12_31_424971 34 19 m1G37 m1g37 NN 10_1101-2020_12_31_424971 34 20 that that WDT 10_1101-2020_12_31_424971 34 21 blocks blocks NNP 10_1101-2020_12_31_424971 34 22 quadruplet quadruplet NN 10_1101-2020_12_31_424971 34 23 pairing pairing NN 10_1101-2020_12_31_424971 34 24 and and CC 10_1101-2020_12_31_424971 34 25 forces force VBZ 10_1101-2020_12_31_424971 34 26 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 34 27 105 105 CD 10_1101-2020_12_31_424971 34 28 to to TO 10_1101-2020_12_31_424971 34 29 use use VB 10_1101-2020_12_31_424971 34 30 0-frame 0-frame CD 10_1101-2020_12_31_424971 34 31 triplet triplet NN 10_1101-2020_12_31_424971 34 32 anticodon anticodon NN 10_1101-2020_12_31_424971 34 33 - - HYPH 10_1101-2020_12_31_424971 34 34 codon codon NNP 10_1101-2020_12_31_424971 34 35 pairing pair VBG 10_1101-2020_12_31_424971 34 36 to to TO 10_1101-2020_12_31_424971 34 37 decode decode VB 10_1101-2020_12_31_424971 34 38 the the DT 10_1101-2020_12_31_424971 34 39 quadruplet quadruplet NN 10_1101-2020_12_31_424971 34 40 codon codon NNP 10_1101-2020_12_31_424971 34 41 at at IN 10_1101-2020_12_31_424971 34 42 the the DT 10_1101-2020_12_31_424971 34 43 A A NNP 10_1101-2020_12_31_424971 34 44 site site NN 10_1101-2020_12_31_424971 34 45 . . . 10_1101-2020_12_31_424971 35 1 106 106 CD 10_1101-2020_12_31_424971 35 2 Additionally additionally RB 10_1101-2020_12_31_424971 35 3 , , , 10_1101-2020_12_31_424971 35 4 we -PRON- PRP 10_1101-2020_12_31_424971 35 5 find find VBP 10_1101-2020_12_31_424971 35 6 that that IN 10_1101-2020_12_31_424971 35 7 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 35 8 , , , 10_1101-2020_12_31_424971 35 9 and and CC 10_1101-2020_12_31_424971 35 10 likely likely RB 10_1101-2020_12_31_424971 35 11 all all DT 10_1101-2020_12_31_424971 35 12 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 35 13 tRNAs trnas CD 10_1101-2020_12_31_424971 35 14 , , , 10_1101-2020_12_31_424971 35 15 shifts shift NNS 10_1101-2020_12_31_424971 35 16 to to IN 10_1101-2020_12_31_424971 35 17 the the DT 10_1101-2020_12_31_424971 35 18 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 35 19 during during IN 10_1101-2020_12_31_424971 35 20 107 107 CD 10_1101-2020_12_31_424971 35 21 the the DT 10_1101-2020_12_31_424971 35 22 subsequent subsequent JJ 10_1101-2020_12_31_424971 35 23 translocation translocation NN 10_1101-2020_12_31_424971 35 24 reaction reaction NN 10_1101-2020_12_31_424971 35 25 in in IN 10_1101-2020_12_31_424971 35 26 which which WDT 10_1101-2020_12_31_424971 35 27 the the DT 10_1101-2020_12_31_424971 35 28 translational translational JJ 10_1101-2020_12_31_424971 35 29 GTPase GTPase NNP 10_1101-2020_12_31_424971 35 30 elongation elongation NN 10_1101-2020_12_31_424971 35 31 factor factor NN 10_1101-2020_12_31_424971 35 32 ( ( -LRB- 10_1101-2020_12_31_424971 35 33 EF)-G EF)-G NNP 10_1101-2020_12_31_424971 35 34 108 108 CD 10_1101-2020_12_31_424971 35 35 catalyzes catalyze VBZ 10_1101-2020_12_31_424971 35 36 the the DT 10_1101-2020_12_31_424971 35 37 movement movement NN 10_1101-2020_12_31_424971 35 38 of of IN 10_1101-2020_12_31_424971 35 39 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 35 40 from from IN 10_1101-2020_12_31_424971 35 41 the the DT 10_1101-2020_12_31_424971 35 42 A a NN 10_1101-2020_12_31_424971 35 43 to to IN 10_1101-2020_12_31_424971 35 44 P p NN 10_1101-2020_12_31_424971 35 45 sites site NNS 10_1101-2020_12_31_424971 35 46 ( ( -LRB- 10_1101-2020_12_31_424971 35 47 i.e. i.e. FW 10_1101-2020_12_31_424971 35 48 , , , 10_1101-2020_12_31_424971 35 49 a a DT 10_1101-2020_12_31_424971 35 50 triplet triplet NN 10_1101-2020_12_31_424971 35 51 - - HYPH 10_1101-2020_12_31_424971 35 52 slippage slippage NN 10_1101-2020_12_31_424971 35 53 mechanism mechanism NN 10_1101-2020_12_31_424971 35 54 ) ) -RRB- 10_1101-2020_12_31_424971 35 55 . . . 10_1101-2020_12_31_424971 36 1 More More JJR 10_1101-2020_12_31_424971 36 2 109 109 CD 10_1101-2020_12_31_424971 36 3 specifically specifically RB 10_1101-2020_12_31_424971 36 4 , , , 10_1101-2020_12_31_424971 36 5 we -PRON- PRP 10_1101-2020_12_31_424971 36 6 show show VBP 10_1101-2020_12_31_424971 36 7 that that IN 10_1101-2020_12_31_424971 36 8 this this DT 10_1101-2020_12_31_424971 36 9 frameshift frameshift NN 10_1101-2020_12_31_424971 36 10 occurs occur VBZ 10_1101-2020_12_31_424971 36 11 in in IN 10_1101-2020_12_31_424971 36 12 the the DT 10_1101-2020_12_31_424971 36 13 later later JJ 10_1101-2020_12_31_424971 36 14 steps step NNS 10_1101-2020_12_31_424971 36 15 of of IN 10_1101-2020_12_31_424971 36 16 translocation translocation NN 10_1101-2020_12_31_424971 36 17 , , , 10_1101-2020_12_31_424971 36 18 during during IN 10_1101-2020_12_31_424971 36 19 which which WDT 10_1101-2020_12_31_424971 36 20 110 110 CD 10_1101-2020_12_31_424971 36 21 EF EF NNP 10_1101-2020_12_31_424971 36 22 - - HYPH 10_1101-2020_12_31_424971 36 23 G G NNP 10_1101-2020_12_31_424971 36 24 catalyzes catalyze VBZ 10_1101-2020_12_31_424971 36 25 a a DT 10_1101-2020_12_31_424971 36 26 series series NN 10_1101-2020_12_31_424971 36 27 of of IN 10_1101-2020_12_31_424971 36 28 conformational conformational JJ 10_1101-2020_12_31_424971 36 29 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 36 30 of of IN 10_1101-2020_12_31_424971 36 31 the the DT 10_1101-2020_12_31_424971 36 32 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 36 33 pre pre JJ 10_1101-2020_12_31_424971 36 34 - - JJ 10_1101-2020_12_31_424971 36 35 translocation translocation JJ 10_1101-2020_12_31_424971 36 36 111 111 CD 10_1101-2020_12_31_424971 36 37 ( ( -LRB- 10_1101-2020_12_31_424971 36 38 PRE PRE NNP 10_1101-2020_12_31_424971 36 39 ) ) -RRB- 10_1101-2020_12_31_424971 36 40 complex complex NN 10_1101-2020_12_31_424971 36 41 that that WDT 10_1101-2020_12_31_424971 36 42 enable enable VBP 10_1101-2020_12_31_424971 36 43 the the DT 10_1101-2020_12_31_424971 36 44 tRNA trna NN 10_1101-2020_12_31_424971 36 45 ASLs asl NNS 10_1101-2020_12_31_424971 36 46 and and CC 10_1101-2020_12_31_424971 36 47 their -PRON- PRP$ 10_1101-2020_12_31_424971 36 48 associated associated JJ 10_1101-2020_12_31_424971 36 49 codons codon NNS 10_1101-2020_12_31_424971 36 50 to to TO 10_1101-2020_12_31_424971 36 51 move move VB 10_1101-2020_12_31_424971 36 52 to to IN 10_1101-2020_12_31_424971 36 53 their -PRON- PRP$ 10_1101-2020_12_31_424971 36 54 112 112 CD 10_1101-2020_12_31_424971 36 55 respective respective JJ 10_1101-2020_12_31_424971 36 56 post post JJ 10_1101-2020_12_31_424971 36 57 - - JJ 10_1101-2020_12_31_424971 36 58 translocation translocation JJ 10_1101-2020_12_31_424971 36 59 positions position NNS 10_1101-2020_12_31_424971 36 60 within within IN 10_1101-2020_12_31_424971 36 61 the the DT 10_1101-2020_12_31_424971 36 62 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 36 63 small small NN 10_1101-2020_12_31_424971 36 64 ( ( -LRB- 10_1101-2020_12_31_424971 36 65 30S 30s CD 10_1101-2020_12_31_424971 36 66 in in IN 10_1101-2020_12_31_424971 36 67 bacteria bacteria NNS 10_1101-2020_12_31_424971 36 68 ) ) -RRB- 10_1101-2020_12_31_424971 36 69 subunit22 subunit22 NN 10_1101-2020_12_31_424971 36 70 - - SYM 10_1101-2020_12_31_424971 36 71 28 28 CD 10_1101-2020_12_31_424971 36 72 . . . 10_1101-2020_12_31_424971 37 1 113 113 CD 10_1101-2020_12_31_424971 37 2 Thus thus RB 10_1101-2020_12_31_424971 37 3 , , , 10_1101-2020_12_31_424971 37 4 efforts effort NNS 10_1101-2020_12_31_424971 37 5 to to TO 10_1101-2020_12_31_424971 37 6 increase increase VB 10_1101-2020_12_31_424971 37 7 the the DT 10_1101-2020_12_31_424971 37 8 recoding recode VBG 10_1101-2020_12_31_424971 37 9 efficiency efficiency NN 10_1101-2020_12_31_424971 37 10 of of IN 10_1101-2020_12_31_424971 37 11 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 37 12 tRNAs trnas CD 10_1101-2020_12_31_424971 37 13 should should MD 10_1101-2020_12_31_424971 37 14 focus focus VB 10_1101-2020_12_31_424971 37 15 on on IN 10_1101-2020_12_31_424971 37 16 114 114 CD 10_1101-2020_12_31_424971 37 17 enforcing enforce VBG 10_1101-2020_12_31_424971 37 18 a a DT 10_1101-2020_12_31_424971 37 19 triplet triplet NN 10_1101-2020_12_31_424971 37 20 anticodon anticodon NN 10_1101-2020_12_31_424971 37 21 - - HYPH 10_1101-2020_12_31_424971 37 22 codon codon NNP 10_1101-2020_12_31_424971 37 23 pairing pair VBG 10_1101-2020_12_31_424971 37 24 in in IN 10_1101-2020_12_31_424971 37 25 the the DT 10_1101-2020_12_31_424971 37 26 0-frame 0-frame CD 10_1101-2020_12_31_424971 37 27 at at IN 10_1101-2020_12_31_424971 37 28 the the DT 10_1101-2020_12_31_424971 37 29 A A NNP 10_1101-2020_12_31_424971 37 30 site site NN 10_1101-2020_12_31_424971 37 31 and and CC 10_1101-2020_12_31_424971 37 32 directed direct VBN 10_1101-2020_12_31_424971 37 33 evolution evolution NN 10_1101-2020_12_31_424971 37 34 to to IN 10_1101-2020_12_31_424971 37 35 115 115 CD 10_1101-2020_12_31_424971 37 36 optimize optimize NN 10_1101-2020_12_31_424971 37 37 conformational conformational JJ 10_1101-2020_12_31_424971 37 38 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 37 39 of of IN 10_1101-2020_12_31_424971 37 40 the the DT 10_1101-2020_12_31_424971 37 41 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 37 42 PRE PRE NNP 10_1101-2020_12_31_424971 37 43 complex complex NN 10_1101-2020_12_31_424971 37 44 during during IN 10_1101-2020_12_31_424971 37 45 the the DT 10_1101-2020_12_31_424971 37 46 late late JJ 10_1101-2020_12_31_424971 37 47 stages stage NNS 10_1101-2020_12_31_424971 37 48 of of IN 10_1101-2020_12_31_424971 37 49 116 116 CD 10_1101-2020_12_31_424971 37 50 translocation translocation NN 10_1101-2020_12_31_424971 37 51 . . . 10_1101-2020_12_31_424971 38 1 117 117 CD 10_1101-2020_12_31_424971 38 2 118 118 CD 10_1101-2020_12_31_424971 38 3 RESULTS results NN 10_1101-2020_12_31_424971 38 4 119 119 CD 10_1101-2020_12_31_424971 38 5 Native native JJ 10_1101-2020_12_31_424971 38 6 - - HYPH 10_1101-2020_12_31_424971 38 7 state state NN 10_1101-2020_12_31_424971 38 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 38 9 is be VBZ 10_1101-2020_12_31_424971 38 10 N1-methylated n1-methylate VBN 10_1101-2020_12_31_424971 38 11 at at IN 10_1101-2020_12_31_424971 38 12 G37 G37 NNP 10_1101-2020_12_31_424971 38 13 and and CC 10_1101-2020_12_31_424971 38 14 is be VBZ 10_1101-2020_12_31_424971 38 15 readily readily RB 10_1101-2020_12_31_424971 38 16 aminoacylated aminoacylate VBN 10_1101-2020_12_31_424971 38 17 with with IN 10_1101-2020_12_31_424971 38 18 Pro Pro NNP 10_1101-2020_12_31_424971 38 19 120 120 CD 10_1101-2020_12_31_424971 38 20 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 38 21 contains contain VBZ 10_1101-2020_12_31_424971 38 22 an an DT 10_1101-2020_12_31_424971 38 23 extra extra JJ 10_1101-2020_12_31_424971 38 24 G37a g37a NN 10_1101-2020_12_31_424971 38 25 nucleotide nucleotide NN 10_1101-2020_12_31_424971 38 26 inserted insert VBN 10_1101-2020_12_31_424971 38 27 between between IN 10_1101-2020_12_31_424971 38 28 G37 G37 NNP 10_1101-2020_12_31_424971 38 29 and and CC 10_1101-2020_12_31_424971 38 30 U38 U38 NNP 10_1101-2020_12_31_424971 38 31 of of IN 10_1101-2020_12_31_424971 38 32 ProL20 ProL20 NNP 10_1101-2020_12_31_424971 38 33 ( ( -LRB- 10_1101-2020_12_31_424971 38 34 Figure Figure NNP 10_1101-2020_12_31_424971 38 35 121 121 CD 10_1101-2020_12_31_424971 38 36 1a 1a CD 10_1101-2020_12_31_424971 38 37 ) ) -RRB- 10_1101-2020_12_31_424971 38 38 . . . 10_1101-2020_12_31_424971 39 1 Whether whether IN 10_1101-2020_12_31_424971 39 2 the the DT 10_1101-2020_12_31_424971 39 3 extra extra JJ 10_1101-2020_12_31_424971 39 4 G37a g37a NN 10_1101-2020_12_31_424971 39 5 is be VBZ 10_1101-2020_12_31_424971 39 6 methylated methylate VBN 10_1101-2020_12_31_424971 39 7 and and CC 10_1101-2020_12_31_424971 39 8 how how WRB 10_1101-2020_12_31_424971 39 9 it -PRON- PRP 10_1101-2020_12_31_424971 39 10 affects affect VBZ 10_1101-2020_12_31_424971 39 11 methylation methylation NN 10_1101-2020_12_31_424971 39 12 of of IN 10_1101-2020_12_31_424971 39 13 G37 G37 NNP 10_1101-2020_12_31_424971 39 14 is be VBZ 10_1101-2020_12_31_424971 39 15 unknown unknown JJ 10_1101-2020_12_31_424971 39 16 . . . 10_1101-2020_12_31_424971 40 1 We -PRON- PRP 10_1101-2020_12_31_424971 40 2 122 122 CD 10_1101-2020_12_31_424971 40 3 thus thus RB 10_1101-2020_12_31_424971 40 4 determined determine VBD 10_1101-2020_12_31_424971 40 5 the the DT 10_1101-2020_12_31_424971 40 6 methylation methylation NN 10_1101-2020_12_31_424971 40 7 status status NN 10_1101-2020_12_31_424971 40 8 of of IN 10_1101-2020_12_31_424971 40 9 the the DT 10_1101-2020_12_31_424971 40 10 G37-G37a G37-G37a NNP 10_1101-2020_12_31_424971 40 11 motif motif NN 10_1101-2020_12_31_424971 40 12 using use VBG 10_1101-2020_12_31_424971 40 13 RNase RNase NNP 10_1101-2020_12_31_424971 40 14 T1 T1 NNP 10_1101-2020_12_31_424971 40 15 cleavage cleavage NN 10_1101-2020_12_31_424971 40 16 inhibition inhibition NN 10_1101-2020_12_31_424971 40 17 123 123 CD 10_1101-2020_12_31_424971 40 18 assays assay NNS 10_1101-2020_12_31_424971 40 19 and and CC 10_1101-2020_12_31_424971 40 20 primer primer JJ 10_1101-2020_12_31_424971 40 21 extension extension NN 10_1101-2020_12_31_424971 40 22 inhibition inhibition NN 10_1101-2020_12_31_424971 40 23 assays assays RB 10_1101-2020_12_31_424971 40 24 . . . 10_1101-2020_12_31_424971 41 1 We -PRON- PRP 10_1101-2020_12_31_424971 41 2 first first RB 10_1101-2020_12_31_424971 41 3 generated generate VBD 10_1101-2020_12_31_424971 41 4 a a DT 10_1101-2020_12_31_424971 41 5 plasmid plasmid NN 10_1101-2020_12_31_424971 41 6 - - HYPH 10_1101-2020_12_31_424971 41 7 encoded encode VBN 10_1101-2020_12_31_424971 41 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 41 9 by by IN 10_1101-2020_12_31_424971 41 10 124 124 CD 10_1101-2020_12_31_424971 41 11 inserting inserting NN 10_1101-2020_12_31_424971 41 12 G37a g37a NN 10_1101-2020_12_31_424971 41 13 into into IN 10_1101-2020_12_31_424971 41 14 an an DT 10_1101-2020_12_31_424971 41 15 existing exist VBG 10_1101-2020_12_31_424971 41 16 Tac Tac NNP 10_1101-2020_12_31_424971 41 17 - - HYPH 10_1101-2020_12_31_424971 41 18 inducible inducible JJ 10_1101-2020_12_31_424971 41 19 plasmid plasmid NN 10_1101-2020_12_31_424971 41 20 encoding encode VBG 10_1101-2020_12_31_424971 41 21 Escherichia Escherichia NNP 10_1101-2020_12_31_424971 41 22 coli coli NNS 10_1101-2020_12_31_424971 41 23 ProL29 ProL29 NNP 10_1101-2020_12_31_424971 41 24 , , , 10_1101-2020_12_31_424971 41 25 which which WDT 10_1101-2020_12_31_424971 41 26 has have VBZ 10_1101-2020_12_31_424971 41 27 125 125 CD 10_1101-2020_12_31_424971 41 28 an an DT 10_1101-2020_12_31_424971 41 29 identical identical JJ 10_1101-2020_12_31_424971 41 30 sequence sequence NN 10_1101-2020_12_31_424971 41 31 to to IN 10_1101-2020_12_31_424971 41 32 S. S. NNP 10_1101-2020_12_31_424971 41 33 typhimurium typhimurium NN 10_1101-2020_12_31_424971 41 34 ProL. ProL. . 10_1101-2020_12_31_424971 42 1 We -PRON- PRP 10_1101-2020_12_31_424971 42 2 then then RB 10_1101-2020_12_31_424971 42 3 expressed express VBD 10_1101-2020_12_31_424971 42 4 and and CC 10_1101-2020_12_31_424971 42 5 purified purify VBD 10_1101-2020_12_31_424971 42 6 the the DT 10_1101-2020_12_31_424971 42 7 plasmid-126 plasmid-126 NN 10_1101-2020_12_31_424971 42 8 .CC .CC NFP 10_1101-2020_12_31_424971 42 9 - - : 10_1101-2020_12_31_424971 42 10 BY by IN 10_1101-2020_12_31_424971 42 11 - - HYPH 10_1101-2020_12_31_424971 42 12 NC NC NNP 10_1101-2020_12_31_424971 42 13 - - HYPH 10_1101-2020_12_31_424971 42 14 ND ND NNP 10_1101-2020_12_31_424971 42 15 4.0 4.0 CD 10_1101-2020_12_31_424971 42 16 International International NNP 10_1101-2020_12_31_424971 42 17 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 42 18 under under IN 10_1101-2020_12_31_424971 42 19 a a DT 10_1101-2020_12_31_424971 42 20 ( ( -LRB- 10_1101-2020_12_31_424971 42 21 which which WDT 10_1101-2020_12_31_424971 42 22 was be VBD 10_1101-2020_12_31_424971 42 23 not not RB 10_1101-2020_12_31_424971 42 24 certified certify VBN 10_1101-2020_12_31_424971 42 25 by by IN 10_1101-2020_12_31_424971 42 26 peer peer NN 10_1101-2020_12_31_424971 42 27 review review NN 10_1101-2020_12_31_424971 42 28 ) ) -RRB- 10_1101-2020_12_31_424971 42 29 is be VBZ 10_1101-2020_12_31_424971 42 30 the the DT 10_1101-2020_12_31_424971 42 31 author author NN 10_1101-2020_12_31_424971 42 32 / / SYM 10_1101-2020_12_31_424971 42 33 funder funder NN 10_1101-2020_12_31_424971 42 34 , , , 10_1101-2020_12_31_424971 42 35 who who WP 10_1101-2020_12_31_424971 42 36 has have VBZ 10_1101-2020_12_31_424971 42 37 granted grant VBN 10_1101-2020_12_31_424971 42 38 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 42 39 a a DT 10_1101-2020_12_31_424971 42 40 license license NN 10_1101-2020_12_31_424971 42 41 to to TO 10_1101-2020_12_31_424971 42 42 display display VB 10_1101-2020_12_31_424971 42 43 the the DT 10_1101-2020_12_31_424971 42 44 preprint preprint NN 10_1101-2020_12_31_424971 42 45 in in IN 10_1101-2020_12_31_424971 42 46 perpetuity perpetuity NN 10_1101-2020_12_31_424971 42 47 . . . 10_1101-2020_12_31_424971 43 1 It -PRON- PRP 10_1101-2020_12_31_424971 43 2 is be VBZ 10_1101-2020_12_31_424971 43 3 made make VBN 10_1101-2020_12_31_424971 43 4 The the DT 10_1101-2020_12_31_424971 43 5 copyright copyright NN 10_1101-2020_12_31_424971 43 6 holder holder NN 10_1101-2020_12_31_424971 43 7 for for IN 10_1101-2020_12_31_424971 43 8 this this DT 10_1101-2020_12_31_424971 43 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 43 10 version version NN 10_1101-2020_12_31_424971 43 11 posted post VBD 10_1101-2020_12_31_424971 43 12 January January NNP 10_1101-2020_12_31_424971 43 13 2 2 CD 10_1101-2020_12_31_424971 43 14 , , , 10_1101-2020_12_31_424971 43 15 2021 2021 CD 10_1101-2020_12_31_424971 43 16 . . . 10_1101-2020_12_31_424971 43 17 ; ; : 10_1101-2020_12_31_424971 43 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 43 19 : : : 10_1101-2020_12_31_424971 43 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 43 21 preprint preprint NN 10_1101-2020_12_31_424971 43 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 43 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 43 24 6 6 CD 10_1101-2020_12_31_424971 43 25 encoded encode VBN 10_1101-2020_12_31_424971 43 26 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 43 27 and and CC 10_1101-2020_12_31_424971 43 28 ProL ProL NNP 10_1101-2020_12_31_424971 43 29 from from IN 10_1101-2020_12_31_424971 43 30 an an DT 10_1101-2020_12_31_424971 43 31 E. E. NNP 10_1101-2020_12_31_424971 43 32 coli coli VBZ 10_1101-2020_12_31_424971 43 33 ProL ProL NNP 10_1101-2020_12_31_424971 43 34 knock knock NN 10_1101-2020_12_31_424971 43 35 - - HYPH 10_1101-2020_12_31_424971 43 36 out out NN 10_1101-2020_12_31_424971 43 37 ( ( -LRB- 10_1101-2020_12_31_424971 43 38 ProL ProL NNP 10_1101-2020_12_31_424971 43 39 - - HYPH 10_1101-2020_12_31_424971 43 40 KO KO NNP 10_1101-2020_12_31_424971 43 41 ) ) -RRB- 10_1101-2020_12_31_424971 43 42 strain30 strain30 VB 10_1101-2020_12_31_424971 43 43 containing contain VBG 10_1101-2020_12_31_424971 43 44 all all PDT 10_1101-2020_12_31_424971 43 45 the the DT 10_1101-2020_12_31_424971 43 46 127 127 CD 10_1101-2020_12_31_424971 43 47 endogenous endogenous JJ 10_1101-2020_12_31_424971 43 48 enzymes enzyme NNS 10_1101-2020_12_31_424971 43 49 necessary necessary JJ 10_1101-2020_12_31_424971 43 50 for for IN 10_1101-2020_12_31_424971 43 51 processing process VBG 10_1101-2020_12_31_424971 43 52 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 43 53 and and CC 10_1101-2020_12_31_424971 43 54 ProL ProL NNP 10_1101-2020_12_31_424971 43 55 to to IN 10_1101-2020_12_31_424971 43 56 their -PRON- PRP$ 10_1101-2020_12_31_424971 43 57 S. S. NNP 10_1101-2020_12_31_424971 43 58 typhimurium typhimurium NN 10_1101-2020_12_31_424971 43 59 native native JJ 10_1101-2020_12_31_424971 43 60 128 128 CD 10_1101-2020_12_31_424971 43 61 states state NNS 10_1101-2020_12_31_424971 43 62 such such JJ 10_1101-2020_12_31_424971 43 63 that that IN 10_1101-2020_12_31_424971 43 64 they -PRON- PRP 10_1101-2020_12_31_424971 43 65 possess possess VBP 10_1101-2020_12_31_424971 43 66 the the DT 10_1101-2020_12_31_424971 43 67 full full JJ 10_1101-2020_12_31_424971 43 68 complement complement NN 10_1101-2020_12_31_424971 43 69 of of IN 10_1101-2020_12_31_424971 43 70 naturally naturally RB 10_1101-2020_12_31_424971 43 71 occurring occur VBG 10_1101-2020_12_31_424971 43 72 post post JJ 10_1101-2020_12_31_424971 43 73 - - JJ 10_1101-2020_12_31_424971 43 74 transcriptional transcriptional JJ 10_1101-2020_12_31_424971 43 75 129 129 CD 10_1101-2020_12_31_424971 43 76 modifications modification NNS 10_1101-2020_12_31_424971 43 77 ( ( -LRB- 10_1101-2020_12_31_424971 43 78 termed term VBN 10_1101-2020_12_31_424971 43 79 the the DT 10_1101-2020_12_31_424971 43 80 native native JJ 10_1101-2020_12_31_424971 43 81 - - HYPH 10_1101-2020_12_31_424971 43 82 state state NN 10_1101-2020_12_31_424971 43 83 tRNAs trna NNS 10_1101-2020_12_31_424971 43 84 ) ) -RRB- 10_1101-2020_12_31_424971 43 85 . . . 10_1101-2020_12_31_424971 44 1 In in IN 10_1101-2020_12_31_424971 44 2 addition addition NN 10_1101-2020_12_31_424971 44 3 , , , 10_1101-2020_12_31_424971 44 4 we -PRON- PRP 10_1101-2020_12_31_424971 44 5 prepared prepare VBD 10_1101-2020_12_31_424971 44 6 in in IN 10_1101-2020_12_31_424971 44 7 vitro vitro FW 10_1101-2020_12_31_424971 44 8 transcripts transcript NNS 10_1101-2020_12_31_424971 44 9 of of IN 10_1101-2020_12_31_424971 44 10 130 130 CD 10_1101-2020_12_31_424971 44 11 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 44 12 and and CC 10_1101-2020_12_31_424971 44 13 ProL ProL NNP 10_1101-2020_12_31_424971 44 14 lacking lack VBG 10_1101-2020_12_31_424971 44 15 all all DT 10_1101-2020_12_31_424971 44 16 post post JJ 10_1101-2020_12_31_424971 44 17 - - JJ 10_1101-2020_12_31_424971 44 18 transcriptional transcriptional JJ 10_1101-2020_12_31_424971 44 19 modifications modification NNS 10_1101-2020_12_31_424971 44 20 ( ( -LRB- 10_1101-2020_12_31_424971 44 21 termed term VBN 10_1101-2020_12_31_424971 44 22 the the DT 10_1101-2020_12_31_424971 44 23 G37-state G37-state NNP 10_1101-2020_12_31_424971 44 24 tRNAs trnas CD 10_1101-2020_12_31_424971 44 25 ) ) -RRB- 10_1101-2020_12_31_424971 44 26 , , , 10_1101-2020_12_31_424971 44 27 or or CC 10_1101-2020_12_31_424971 44 28 131 131 CD 10_1101-2020_12_31_424971 44 29 enzymatically enzymatically RB 10_1101-2020_12_31_424971 44 30 methylated methylate VBN 10_1101-2020_12_31_424971 44 31 with with IN 10_1101-2020_12_31_424971 44 32 purified purify VBN 10_1101-2020_12_31_424971 44 33 E. E. NNP 10_1101-2020_12_31_424971 44 34 coli coli NNS 10_1101-2020_12_31_424971 44 35 TrmD30,31 trmd30,31 CD 10_1101-2020_12_31_424971 44 36 such such JJ 10_1101-2020_12_31_424971 44 37 that that IN 10_1101-2020_12_31_424971 44 38 they -PRON- PRP 10_1101-2020_12_31_424971 44 39 possess possess VBP 10_1101-2020_12_31_424971 44 40 only only RB 10_1101-2020_12_31_424971 44 41 the the DT 10_1101-2020_12_31_424971 44 42 N1 N1 NNP 10_1101-2020_12_31_424971 44 43 - - HYPH 10_1101-2020_12_31_424971 44 44 132 132 CD 10_1101-2020_12_31_424971 44 45 methylation methylation NN 10_1101-2020_12_31_424971 44 46 at at IN 10_1101-2020_12_31_424971 44 47 G37 G37 NNP 10_1101-2020_12_31_424971 44 48 and and CC 10_1101-2020_12_31_424971 44 49 no no DT 10_1101-2020_12_31_424971 44 50 other other JJ 10_1101-2020_12_31_424971 44 51 post post JJ 10_1101-2020_12_31_424971 44 52 - - JJ 10_1101-2020_12_31_424971 44 53 transcriptional transcriptional JJ 10_1101-2020_12_31_424971 44 54 modifications modification NNS 10_1101-2020_12_31_424971 44 55 ( ( -LRB- 10_1101-2020_12_31_424971 44 56 termed term VBN 10_1101-2020_12_31_424971 44 57 the the DT 10_1101-2020_12_31_424971 44 58 m1G37-state m1G37-state NNP 10_1101-2020_12_31_424971 44 59 133 133 CD 10_1101-2020_12_31_424971 44 60 tRNAs trnas CD 10_1101-2020_12_31_424971 44 61 ) ) -RRB- 10_1101-2020_12_31_424971 44 62 . . . 10_1101-2020_12_31_424971 45 1 In in IN 10_1101-2020_12_31_424971 45 2 the the DT 10_1101-2020_12_31_424971 45 3 case case NN 10_1101-2020_12_31_424971 45 4 of of IN 10_1101-2020_12_31_424971 45 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 45 6 , , , 10_1101-2020_12_31_424971 45 7 RNase RNase NNP 10_1101-2020_12_31_424971 45 8 T1 T1 NNP 10_1101-2020_12_31_424971 45 9 cleavage cleavage NN 10_1101-2020_12_31_424971 45 10 inhibition inhibition NN 10_1101-2020_12_31_424971 45 11 assays assays RB 10_1101-2020_12_31_424971 45 12 demonstrated demonstrate VBD 10_1101-2020_12_31_424971 45 13 cleavage cleavage NN 10_1101-2020_12_31_424971 45 14 at at IN 10_1101-2020_12_31_424971 45 15 134 134 CD 10_1101-2020_12_31_424971 45 16 G37 G37 NNP 10_1101-2020_12_31_424971 45 17 and and CC 10_1101-2020_12_31_424971 45 18 G37a g37a NN 10_1101-2020_12_31_424971 45 19 of of IN 10_1101-2020_12_31_424971 45 20 the the DT 10_1101-2020_12_31_424971 45 21 G37-state G37-state NNP 10_1101-2020_12_31_424971 45 22 tRNA trna NN 10_1101-2020_12_31_424971 45 23 , , , 10_1101-2020_12_31_424971 45 24 but but CC 10_1101-2020_12_31_424971 45 25 inhibition inhibition NN 10_1101-2020_12_31_424971 45 26 of of IN 10_1101-2020_12_31_424971 45 27 cleavage cleavage NN 10_1101-2020_12_31_424971 45 28 at at IN 10_1101-2020_12_31_424971 45 29 either either DT 10_1101-2020_12_31_424971 45 30 position position NN 10_1101-2020_12_31_424971 45 31 upon upon IN 10_1101-2020_12_31_424971 45 32 treatment treatment NN 10_1101-2020_12_31_424971 45 33 135 135 CD 10_1101-2020_12_31_424971 45 34 with with IN 10_1101-2020_12_31_424971 45 35 TrmD TrmD NNP 10_1101-2020_12_31_424971 45 36 ( ( -LRB- 10_1101-2020_12_31_424971 45 37 Figure Figure NNP 10_1101-2020_12_31_424971 45 38 1b 1b NNP 10_1101-2020_12_31_424971 45 39 ) ) -RRB- 10_1101-2020_12_31_424971 45 40 , , , 10_1101-2020_12_31_424971 45 41 indicating indicate VBG 10_1101-2020_12_31_424971 45 42 that that IN 10_1101-2020_12_31_424971 45 43 both both DT 10_1101-2020_12_31_424971 45 44 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 45 45 are be VBP 10_1101-2020_12_31_424971 45 46 N1-methylated n1-methylate VBN 10_1101-2020_12_31_424971 45 47 in in IN 10_1101-2020_12_31_424971 45 48 the the DT 10_1101-2020_12_31_424971 45 49 m1G37-state m1G37-state NNP 10_1101-2020_12_31_424971 45 50 136 136 CD 10_1101-2020_12_31_424971 45 51 tRNA trna NN 10_1101-2020_12_31_424971 45 52 . . . 10_1101-2020_12_31_424971 46 1 137 137 CD 10_1101-2020_12_31_424971 46 2 Primer Primer NNP 10_1101-2020_12_31_424971 46 3 extension extension NN 10_1101-2020_12_31_424971 46 4 inhibition inhibition NN 10_1101-2020_12_31_424971 46 5 assays assays RB 10_1101-2020_12_31_424971 46 6 , , , 10_1101-2020_12_31_424971 46 7 which which WDT 10_1101-2020_12_31_424971 46 8 were be VBD 10_1101-2020_12_31_424971 46 9 previously previously RB 10_1101-2020_12_31_424971 46 10 validated validate VBN 10_1101-2020_12_31_424971 46 11 by by IN 10_1101-2020_12_31_424971 46 12 mass mass NNP 10_1101-2020_12_31_424971 46 13 spectrometry spectrometry NNP 10_1101-2020_12_31_424971 46 14 138 138 CD 10_1101-2020_12_31_424971 46 15 analysis30 analysis30 NNS 10_1101-2020_12_31_424971 46 16 , , , 10_1101-2020_12_31_424971 46 17 showed show VBD 10_1101-2020_12_31_424971 46 18 inhibition inhibition NN 10_1101-2020_12_31_424971 46 19 of of IN 10_1101-2020_12_31_424971 46 20 extension extension NN 10_1101-2020_12_31_424971 46 21 at at IN 10_1101-2020_12_31_424971 46 22 G37 G37 NNP 10_1101-2020_12_31_424971 46 23 and and CC 10_1101-2020_12_31_424971 46 24 G37a G37a NNS 10_1101-2020_12_31_424971 46 25 in in IN 10_1101-2020_12_31_424971 46 26 m1G37- m1g37- CD 10_1101-2020_12_31_424971 46 27 and and CC 10_1101-2020_12_31_424971 46 28 native native JJ 10_1101-2020_12_31_424971 46 29 - - HYPH 10_1101-2020_12_31_424971 46 30 state state NN 10_1101-2020_12_31_424971 46 31 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 46 32 139 139 CD 10_1101-2020_12_31_424971 46 33 ( ( -LRB- 10_1101-2020_12_31_424971 46 34 Figure Figure NNP 10_1101-2020_12_31_424971 46 35 1c 1c NNP 10_1101-2020_12_31_424971 46 36 ) ) -RRB- 10_1101-2020_12_31_424971 46 37 , , , 10_1101-2020_12_31_424971 46 38 confirming confirm VBG 10_1101-2020_12_31_424971 46 39 that that IN 10_1101-2020_12_31_424971 46 40 both both DT 10_1101-2020_12_31_424971 46 41 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 46 42 are be VBP 10_1101-2020_12_31_424971 46 43 N1-methylated n1-methylate VBN 10_1101-2020_12_31_424971 46 44 in in IN 10_1101-2020_12_31_424971 46 45 these these DT 10_1101-2020_12_31_424971 46 46 species specie NNS 10_1101-2020_12_31_424971 46 47 . . . 10_1101-2020_12_31_424971 47 1 Notably notably RB 10_1101-2020_12_31_424971 47 2 , , , 10_1101-2020_12_31_424971 47 3 N1 N1 NNP 10_1101-2020_12_31_424971 47 4 140 140 CD 10_1101-2020_12_31_424971 47 5 methylation methylation NN 10_1101-2020_12_31_424971 47 6 shifted shift VBD 10_1101-2020_12_31_424971 47 7 almost almost RB 10_1101-2020_12_31_424971 47 8 entirely entirely RB 10_1101-2020_12_31_424971 47 9 to to IN 10_1101-2020_12_31_424971 47 10 G37 G37 NNP 10_1101-2020_12_31_424971 47 11 in in IN 10_1101-2020_12_31_424971 47 12 native native JJ 10_1101-2020_12_31_424971 47 13 - - HYPH 10_1101-2020_12_31_424971 47 14 state state NN 10_1101-2020_12_31_424971 47 15 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 47 16 , , , 10_1101-2020_12_31_424971 47 17 indicating indicate VBG 10_1101-2020_12_31_424971 47 18 that that IN 10_1101-2020_12_31_424971 47 19 m1G37 m1g37 NN 10_1101-2020_12_31_424971 47 20 is be VBZ 10_1101-2020_12_31_424971 47 21 the the DT 10_1101-2020_12_31_424971 47 22 141 141 CD 10_1101-2020_12_31_424971 47 23 dominant dominant JJ 10_1101-2020_12_31_424971 47 24 methylation methylation NN 10_1101-2020_12_31_424971 47 25 product product NN 10_1101-2020_12_31_424971 47 26 in in IN 10_1101-2020_12_31_424971 47 27 cells cell NNS 10_1101-2020_12_31_424971 47 28 . . . 10_1101-2020_12_31_424971 48 1 As as IN 10_1101-2020_12_31_424971 48 2 a a DT 10_1101-2020_12_31_424971 48 3 control control NN 10_1101-2020_12_31_424971 48 4 , , , 10_1101-2020_12_31_424971 48 5 no no DT 10_1101-2020_12_31_424971 48 6 inhibition inhibition NN 10_1101-2020_12_31_424971 48 7 of of IN 10_1101-2020_12_31_424971 48 8 extension extension NN 10_1101-2020_12_31_424971 48 9 at at IN 10_1101-2020_12_31_424971 48 10 G37 G37 NNP 10_1101-2020_12_31_424971 48 11 or or CC 10_1101-2020_12_31_424971 48 12 G37a g37a NN 10_1101-2020_12_31_424971 48 13 was be VBD 10_1101-2020_12_31_424971 48 14 142 142 CD 10_1101-2020_12_31_424971 48 15 observed observe VBN 10_1101-2020_12_31_424971 48 16 for for IN 10_1101-2020_12_31_424971 48 17 G37-state G37-state NNP 10_1101-2020_12_31_424971 48 18 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 48 19 . . . 10_1101-2020_12_31_424971 49 1 Complementary complementary JJ 10_1101-2020_12_31_424971 49 2 kinetics kinetics NN 10_1101-2020_12_31_424971 49 3 experiments experiment NNS 10_1101-2020_12_31_424971 49 4 showed show VBD 10_1101-2020_12_31_424971 49 5 that that IN 10_1101-2020_12_31_424971 49 6 the the DT 10_1101-2020_12_31_424971 49 7 yield yield NN 10_1101-2020_12_31_424971 49 8 and and CC 10_1101-2020_12_31_424971 49 9 143 143 CD 10_1101-2020_12_31_424971 49 10 rate rate NN 10_1101-2020_12_31_424971 49 11 of of IN 10_1101-2020_12_31_424971 49 12 N1-methylation n1-methylation DT 10_1101-2020_12_31_424971 49 13 of of IN 10_1101-2020_12_31_424971 49 14 G37-state G37-state NNP 10_1101-2020_12_31_424971 49 15 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 49 16 were be VBD 10_1101-2020_12_31_424971 49 17 similar similar JJ 10_1101-2020_12_31_424971 49 18 to to IN 10_1101-2020_12_31_424971 49 19 those those DT 10_1101-2020_12_31_424971 49 20 of of IN 10_1101-2020_12_31_424971 49 21 G37-state G37-state NNP 10_1101-2020_12_31_424971 49 22 ProL ProL NNP 10_1101-2020_12_31_424971 49 23 ( ( -LRB- 10_1101-2020_12_31_424971 49 24 Figure Figure NNP 10_1101-2020_12_31_424971 49 25 1d 1d CD 10_1101-2020_12_31_424971 49 26 ) ) -RRB- 10_1101-2020_12_31_424971 49 27 . . . 10_1101-2020_12_31_424971 50 1 144 144 CD 10_1101-2020_12_31_424971 50 2 Likewise likewise RB 10_1101-2020_12_31_424971 50 3 , , , 10_1101-2020_12_31_424971 50 4 kinetics kinetic NNS 10_1101-2020_12_31_424971 50 5 experiments experiment NNS 10_1101-2020_12_31_424971 50 6 revealed reveal VBD 10_1101-2020_12_31_424971 50 7 that that IN 10_1101-2020_12_31_424971 50 8 the the DT 10_1101-2020_12_31_424971 50 9 yield yield NN 10_1101-2020_12_31_424971 50 10 and and CC 10_1101-2020_12_31_424971 50 11 rate rate NN 10_1101-2020_12_31_424971 50 12 of of IN 10_1101-2020_12_31_424971 50 13 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 50 14 of of IN 10_1101-2020_12_31_424971 50 15 native native JJ 10_1101-2020_12_31_424971 50 16 - - HYPH 10_1101-2020_12_31_424971 50 17 state state NN 10_1101-2020_12_31_424971 50 18 145 145 CD 10_1101-2020_12_31_424971 50 19 SufB2 sufb2 NN 10_1101-2020_12_31_424971 50 20 with with IN 10_1101-2020_12_31_424971 50 21 Pro Pro NNP 10_1101-2020_12_31_424971 50 22 were be VBD 10_1101-2020_12_31_424971 50 23 similar similar JJ 10_1101-2020_12_31_424971 50 24 to to IN 10_1101-2020_12_31_424971 50 25 those those DT 10_1101-2020_12_31_424971 50 26 of of IN 10_1101-2020_12_31_424971 50 27 native native JJ 10_1101-2020_12_31_424971 50 28 - - HYPH 10_1101-2020_12_31_424971 50 29 state state NN 10_1101-2020_12_31_424971 50 30 ProL ProL NNP 10_1101-2020_12_31_424971 50 31 ( ( -LRB- 10_1101-2020_12_31_424971 50 32 Figure Figure NNP 10_1101-2020_12_31_424971 50 33 1e 1e NNP 10_1101-2020_12_31_424971 50 34 ) ) -RRB- 10_1101-2020_12_31_424971 50 35 . . . 10_1101-2020_12_31_424971 51 1 In in IN 10_1101-2020_12_31_424971 51 2 contrast contrast NN 10_1101-2020_12_31_424971 51 3 , , , 10_1101-2020_12_31_424971 51 4 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 51 5 146 146 CD 10_1101-2020_12_31_424971 51 6 of of IN 10_1101-2020_12_31_424971 51 7 G37-state G37-state NNP 10_1101-2020_12_31_424971 51 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 51 9 was be VBD 10_1101-2020_12_31_424971 51 10 inhibited inhibit VBN 10_1101-2020_12_31_424971 51 11 ( ( -LRB- 10_1101-2020_12_31_424971 51 12 Figure Figure NNP 10_1101-2020_12_31_424971 51 13 1f 1f NNP 10_1101-2020_12_31_424971 51 14 ) ) -RRB- 10_1101-2020_12_31_424971 51 15 . . . 10_1101-2020_12_31_424971 52 1 These these DT 10_1101-2020_12_31_424971 52 2 results result NNS 10_1101-2020_12_31_424971 52 3 demonstrate demonstrate VBP 10_1101-2020_12_31_424971 52 4 that that IN 10_1101-2020_12_31_424971 52 5 the the DT 10_1101-2020_12_31_424971 52 6 native native JJ 10_1101-2020_12_31_424971 52 7 - - HYPH 10_1101-2020_12_31_424971 52 8 state state NN 10_1101-2020_12_31_424971 52 9 147 147 CD 10_1101-2020_12_31_424971 52 10 SufB2 sufb2 NN 10_1101-2020_12_31_424971 52 11 synthesized synthesize VBN 10_1101-2020_12_31_424971 52 12 in in IN 10_1101-2020_12_31_424971 52 13 cells cell NNS 10_1101-2020_12_31_424971 52 14 is be VBZ 10_1101-2020_12_31_424971 52 15 quantitatively quantitatively RB 10_1101-2020_12_31_424971 52 16 N1-methylated n1-methylate VBN 10_1101-2020_12_31_424971 52 17 to to TO 10_1101-2020_12_31_424971 52 18 generate generate VB 10_1101-2020_12_31_424971 52 19 m1G37 m1G37 NNP 10_1101-2020_12_31_424971 52 20 and and CC 10_1101-2020_12_31_424971 52 21 is be VBZ 10_1101-2020_12_31_424971 52 22 readily readily RB 10_1101-2020_12_31_424971 52 23 148 148 CD 10_1101-2020_12_31_424971 52 24 aminoacylated aminoacylate VBN 10_1101-2020_12_31_424971 52 25 with with IN 10_1101-2020_12_31_424971 52 26 Pro Pro NNP 10_1101-2020_12_31_424971 52 27 . . . 10_1101-2020_12_31_424971 53 1 149 149 CD 10_1101-2020_12_31_424971 53 2 150 150 CD 10_1101-2020_12_31_424971 53 3 SufB2 sufb2 NN 10_1101-2020_12_31_424971 53 4 promotes promote NNS 10_1101-2020_12_31_424971 53 5 +1 +1 NNS 10_1101-2020_12_31_424971 53 6 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 53 7 using use VBG 10_1101-2020_12_31_424971 53 8 triplet triplet NN 10_1101-2020_12_31_424971 53 9 - - HYPH 10_1101-2020_12_31_424971 53 10 slippage slippage NN 10_1101-2020_12_31_424971 53 11 and and CC 10_1101-2020_12_31_424971 53 12 possibly possibly RB 10_1101-2020_12_31_424971 53 13 other other JJ 10_1101-2020_12_31_424971 53 14 mechanisms mechanism NNS 10_1101-2020_12_31_424971 53 15 151 151 CD 10_1101-2020_12_31_424971 53 16 .CC .CC , 10_1101-2020_12_31_424971 53 17 - - : 10_1101-2020_12_31_424971 53 18 BY by IN 10_1101-2020_12_31_424971 53 19 - - HYPH 10_1101-2020_12_31_424971 53 20 NC NC NNP 10_1101-2020_12_31_424971 53 21 - - HYPH 10_1101-2020_12_31_424971 53 22 ND ND NNP 10_1101-2020_12_31_424971 53 23 4.0 4.0 CD 10_1101-2020_12_31_424971 53 24 International International NNP 10_1101-2020_12_31_424971 53 25 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 53 26 under under IN 10_1101-2020_12_31_424971 53 27 a a DT 10_1101-2020_12_31_424971 53 28 ( ( -LRB- 10_1101-2020_12_31_424971 53 29 which which WDT 10_1101-2020_12_31_424971 53 30 was be VBD 10_1101-2020_12_31_424971 53 31 not not RB 10_1101-2020_12_31_424971 53 32 certified certify VBN 10_1101-2020_12_31_424971 53 33 by by IN 10_1101-2020_12_31_424971 53 34 peer peer NN 10_1101-2020_12_31_424971 53 35 review review NN 10_1101-2020_12_31_424971 53 36 ) ) -RRB- 10_1101-2020_12_31_424971 53 37 is be VBZ 10_1101-2020_12_31_424971 53 38 the the DT 10_1101-2020_12_31_424971 53 39 author author NN 10_1101-2020_12_31_424971 53 40 / / SYM 10_1101-2020_12_31_424971 53 41 funder funder NN 10_1101-2020_12_31_424971 53 42 , , , 10_1101-2020_12_31_424971 53 43 who who WP 10_1101-2020_12_31_424971 53 44 has have VBZ 10_1101-2020_12_31_424971 53 45 granted grant VBN 10_1101-2020_12_31_424971 53 46 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 53 47 a a DT 10_1101-2020_12_31_424971 53 48 license license NN 10_1101-2020_12_31_424971 53 49 to to TO 10_1101-2020_12_31_424971 53 50 display display VB 10_1101-2020_12_31_424971 53 51 the the DT 10_1101-2020_12_31_424971 53 52 preprint preprint NN 10_1101-2020_12_31_424971 53 53 in in IN 10_1101-2020_12_31_424971 53 54 perpetuity perpetuity NN 10_1101-2020_12_31_424971 53 55 . . . 10_1101-2020_12_31_424971 54 1 It -PRON- PRP 10_1101-2020_12_31_424971 54 2 is be VBZ 10_1101-2020_12_31_424971 54 3 made make VBN 10_1101-2020_12_31_424971 54 4 The the DT 10_1101-2020_12_31_424971 54 5 copyright copyright NN 10_1101-2020_12_31_424971 54 6 holder holder NN 10_1101-2020_12_31_424971 54 7 for for IN 10_1101-2020_12_31_424971 54 8 this this DT 10_1101-2020_12_31_424971 54 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 54 10 version version NN 10_1101-2020_12_31_424971 54 11 posted post VBD 10_1101-2020_12_31_424971 54 12 January January NNP 10_1101-2020_12_31_424971 54 13 2 2 CD 10_1101-2020_12_31_424971 54 14 , , , 10_1101-2020_12_31_424971 54 15 2021 2021 CD 10_1101-2020_12_31_424971 54 16 . . . 10_1101-2020_12_31_424971 54 17 ; ; : 10_1101-2020_12_31_424971 54 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 54 19 : : : 10_1101-2020_12_31_424971 54 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 54 21 preprint preprint NN 10_1101-2020_12_31_424971 54 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 54 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 54 24 7 7 CD 10_1101-2020_12_31_424971 54 25 We -PRON- PRP 10_1101-2020_12_31_424971 54 26 next next RB 10_1101-2020_12_31_424971 54 27 determined determine VBD 10_1101-2020_12_31_424971 54 28 the the DT 10_1101-2020_12_31_424971 54 29 mechanism(s mechanism(s NNP 10_1101-2020_12_31_424971 54 30 ) ) -RRB- 10_1101-2020_12_31_424971 54 31 through through IN 10_1101-2020_12_31_424971 54 32 which which WDT 10_1101-2020_12_31_424971 54 33 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 54 34 promotes promote VBZ 10_1101-2020_12_31_424971 54 35 +1 +1 NNS 10_1101-2020_12_31_424971 54 36 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 54 37 in in IN 10_1101-2020_12_31_424971 54 38 a a DT 10_1101-2020_12_31_424971 54 39 152 152 CD 10_1101-2020_12_31_424971 54 40 cellular cellular JJ 10_1101-2020_12_31_424971 54 41 context context NN 10_1101-2020_12_31_424971 54 42 . . . 10_1101-2020_12_31_424971 55 1 We -PRON- PRP 10_1101-2020_12_31_424971 55 2 created create VBD 10_1101-2020_12_31_424971 55 3 a a DT 10_1101-2020_12_31_424971 55 4 pair pair NN 10_1101-2020_12_31_424971 55 5 of of IN 10_1101-2020_12_31_424971 55 6 isogenic isogenic JJ 10_1101-2020_12_31_424971 55 7 E. E. NNP 10_1101-2020_12_31_424971 55 8 coli coli NNS 10_1101-2020_12_31_424971 55 9 strains strain NNS 10_1101-2020_12_31_424971 55 10 expressing express VBG 10_1101-2020_12_31_424971 55 11 SufB2 sufb2 NN 10_1101-2020_12_31_424971 55 12 or or CC 10_1101-2020_12_31_424971 55 13 ProL ProL NNP 10_1101-2020_12_31_424971 55 14 from from IN 10_1101-2020_12_31_424971 55 15 the the DT 10_1101-2020_12_31_424971 55 16 153 153 CD 10_1101-2020_12_31_424971 55 17 chromosome chromosome NN 10_1101-2020_12_31_424971 55 18 in in IN 10_1101-2020_12_31_424971 55 19 a a DT 10_1101-2020_12_31_424971 55 20 trmD trmd CD 10_1101-2020_12_31_424971 55 21 - - HYPH 10_1101-2020_12_31_424971 55 22 knockdown knockdown VBN 10_1101-2020_12_31_424971 55 23 ( ( -LRB- 10_1101-2020_12_31_424971 55 24 trmD trmD NNP 10_1101-2020_12_31_424971 55 25 - - HYPH 10_1101-2020_12_31_424971 55 26 KD KD NNP 10_1101-2020_12_31_424971 55 27 ) ) -RRB- 10_1101-2020_12_31_424971 55 28 background30 background30 VBZ 10_1101-2020_12_31_424971 55 29 . . . 10_1101-2020_12_31_424971 56 1 This this DT 10_1101-2020_12_31_424971 56 2 background background NN 10_1101-2020_12_31_424971 56 3 strain strain NN 10_1101-2020_12_31_424971 56 4 was be VBD 10_1101-2020_12_31_424971 56 5 154 154 CD 10_1101-2020_12_31_424971 56 6 designed design VBN 10_1101-2020_12_31_424971 56 7 to to TO 10_1101-2020_12_31_424971 56 8 evaluate evaluate VB 10_1101-2020_12_31_424971 56 9 the the DT 10_1101-2020_12_31_424971 56 10 effect effect NN 10_1101-2020_12_31_424971 56 11 of of IN 10_1101-2020_12_31_424971 56 12 m1G37 m1g37 NN 10_1101-2020_12_31_424971 56 13 on on IN 10_1101-2020_12_31_424971 56 14 +1 +1 NNS 10_1101-2020_12_31_424971 56 15 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 56 16 and and CC 10_1101-2020_12_31_424971 56 17 it -PRON- PRP 10_1101-2020_12_31_424971 56 18 was be VBD 10_1101-2020_12_31_424971 56 19 generated generate VBN 10_1101-2020_12_31_424971 56 20 by by IN 10_1101-2020_12_31_424971 56 21 deleting delete VBG 10_1101-2020_12_31_424971 56 22 155 155 CD 10_1101-2020_12_31_424971 56 23 chromosomal chromosomal JJ 10_1101-2020_12_31_424971 56 24 trmD trmD NNS 10_1101-2020_12_31_424971 56 25 and and CC 10_1101-2020_12_31_424971 56 26 controlling control VBG 10_1101-2020_12_31_424971 56 27 cellular cellular JJ 10_1101-2020_12_31_424971 56 28 levels level NNS 10_1101-2020_12_31_424971 56 29 of of IN 10_1101-2020_12_31_424971 56 30 m1G37 m1g37 NN 10_1101-2020_12_31_424971 56 31 using use VBG 10_1101-2020_12_31_424971 56 32 arabinose arabinose RB 10_1101-2020_12_31_424971 56 33 - - HYPH 10_1101-2020_12_31_424971 56 34 induced induce VBN 10_1101-2020_12_31_424971 56 35 expression expression NN 10_1101-2020_12_31_424971 56 36 156 156 CD 10_1101-2020_12_31_424971 56 37 of of IN 10_1101-2020_12_31_424971 56 38 the the DT 10_1101-2020_12_31_424971 56 39 human human JJ 10_1101-2020_12_31_424971 56 40 counterpart counterpart NN 10_1101-2020_12_31_424971 56 41 trm5 trm5 NN 10_1101-2020_12_31_424971 56 42 , , , 10_1101-2020_12_31_424971 56 43 which which WDT 10_1101-2020_12_31_424971 56 44 is be VBZ 10_1101-2020_12_31_424971 56 45 competent competent JJ 10_1101-2020_12_31_424971 56 46 to to TO 10_1101-2020_12_31_424971 56 47 stoichiometrically stoichiometrically RB 10_1101-2020_12_31_424971 56 48 N1-methylate n1-methylate VB 10_1101-2020_12_31_424971 56 49 intracellular intracellular JJ 10_1101-2020_12_31_424971 56 50 157 157 CD 10_1101-2020_12_31_424971 56 51 tRNA trna NN 10_1101-2020_12_31_424971 56 52 substrates30 substrates30 NNS 10_1101-2020_12_31_424971 56 53 . . . 10_1101-2020_12_31_424971 57 1 The the DT 10_1101-2020_12_31_424971 57 2 isogenic isogenic JJ 10_1101-2020_12_31_424971 57 3 pair pair NN 10_1101-2020_12_31_424971 57 4 of of IN 10_1101-2020_12_31_424971 57 5 the the DT 10_1101-2020_12_31_424971 57 6 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 57 7 and and CC 10_1101-2020_12_31_424971 57 8 ProL ProL NNP 10_1101-2020_12_31_424971 57 9 strains strain NNS 10_1101-2020_12_31_424971 57 10 were be VBD 10_1101-2020_12_31_424971 57 11 measured measure VBN 10_1101-2020_12_31_424971 57 12 for for IN 10_1101-2020_12_31_424971 57 13 +1 +1 NNS 10_1101-2020_12_31_424971 57 14 158 158 CD 10_1101-2020_12_31_424971 57 15 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 57 16 in in IN 10_1101-2020_12_31_424971 57 17 a a DT 10_1101-2020_12_31_424971 57 18 cell cell NN 10_1101-2020_12_31_424971 57 19 - - HYPH 10_1101-2020_12_31_424971 57 20 based base VBN 10_1101-2020_12_31_424971 57 21 lacZ lacz JJ 10_1101-2020_12_31_424971 57 22 reporter reporter NN 10_1101-2020_12_31_424971 57 23 assay assay NN 10_1101-2020_12_31_424971 57 24 in in IN 10_1101-2020_12_31_424971 57 25 which which WDT 10_1101-2020_12_31_424971 57 26 a a DT 10_1101-2020_12_31_424971 57 27 CCC CCC NNP 10_1101-2020_12_31_424971 57 28 - - HYPH 10_1101-2020_12_31_424971 57 29 C C NNP 10_1101-2020_12_31_424971 57 30 motif motif NN 10_1101-2020_12_31_424971 57 31 was be VBD 10_1101-2020_12_31_424971 57 32 inserted insert VBN 10_1101-2020_12_31_424971 57 33 into into IN 10_1101-2020_12_31_424971 57 34 the the DT 10_1101-2020_12_31_424971 57 35 2nd 2nd JJ 10_1101-2020_12_31_424971 57 36 159 159 CD 10_1101-2020_12_31_424971 57 37 codon codon NN 10_1101-2020_12_31_424971 57 38 position position NN 10_1101-2020_12_31_424971 57 39 of of IN 10_1101-2020_12_31_424971 57 40 lacZ lacZ , 10_1101-2020_12_31_424971 57 41 such such JJ 10_1101-2020_12_31_424971 57 42 that that IN 10_1101-2020_12_31_424971 57 43 a a DT 10_1101-2020_12_31_424971 57 44 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 57 45 event event NN 10_1101-2020_12_31_424971 57 46 at at IN 10_1101-2020_12_31_424971 57 47 the the DT 10_1101-2020_12_31_424971 57 48 motif motif NN 10_1101-2020_12_31_424971 57 49 was be VBD 10_1101-2020_12_31_424971 57 50 necessary necessary JJ 10_1101-2020_12_31_424971 57 51 to to TO 10_1101-2020_12_31_424971 57 52 synthesize synthesize VB 10_1101-2020_12_31_424971 57 53 160 160 CD 10_1101-2020_12_31_424971 57 54 full full JJ 10_1101-2020_12_31_424971 57 55 - - HYPH 10_1101-2020_12_31_424971 57 56 length length NN 10_1101-2020_12_31_424971 57 57 b b NN 10_1101-2020_12_31_424971 57 58 - - HYPH 10_1101-2020_12_31_424971 57 59 galactosidase galactosidase NN 10_1101-2020_12_31_424971 57 60 ( ( -LRB- 10_1101-2020_12_31_424971 57 61 b b NNP 10_1101-2020_12_31_424971 57 62 - - HYPH 10_1101-2020_12_31_424971 57 63 Gal)29 Gal)29 NNP 10_1101-2020_12_31_424971 57 64 . . . 10_1101-2020_12_31_424971 58 1 The the DT 10_1101-2020_12_31_424971 58 2 efficiency efficiency NN 10_1101-2020_12_31_424971 58 3 of of IN 10_1101-2020_12_31_424971 58 4 +1 +1 NNS 10_1101-2020_12_31_424971 58 5 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 58 6 was be VBD 10_1101-2020_12_31_424971 58 7 calculated calculate VBN 10_1101-2020_12_31_424971 58 8 as as IN 10_1101-2020_12_31_424971 58 9 the the DT 10_1101-2020_12_31_424971 58 10 ratio ratio NN 10_1101-2020_12_31_424971 58 11 161 161 CD 10_1101-2020_12_31_424971 58 12 of of IN 10_1101-2020_12_31_424971 58 13 b b NN 10_1101-2020_12_31_424971 58 14 - - : 10_1101-2020_12_31_424971 58 15 Gal Gal NNP 10_1101-2020_12_31_424971 58 16 expressed express VBD 10_1101-2020_12_31_424971 58 17 in in IN 10_1101-2020_12_31_424971 58 18 cells cell NNS 10_1101-2020_12_31_424971 58 19 containing contain VBG 10_1101-2020_12_31_424971 58 20 the the DT 10_1101-2020_12_31_424971 58 21 CCC CCC NNP 10_1101-2020_12_31_424971 58 22 - - HYPH 10_1101-2020_12_31_424971 58 23 C C NNP 10_1101-2020_12_31_424971 58 24 insertion insertion NN 10_1101-2020_12_31_424971 58 25 relative relative JJ 10_1101-2020_12_31_424971 58 26 to to IN 10_1101-2020_12_31_424971 58 27 cells cell NNS 10_1101-2020_12_31_424971 58 28 containing contain VBG 10_1101-2020_12_31_424971 58 29 an an DT 10_1101-2020_12_31_424971 58 30 in in IN 10_1101-2020_12_31_424971 58 31 - - HYPH 10_1101-2020_12_31_424971 58 32 frame frame NN 10_1101-2020_12_31_424971 58 33 162 162 CD 10_1101-2020_12_31_424971 58 34 CCC CCC NNP 10_1101-2020_12_31_424971 58 35 insertion insertion NN 10_1101-2020_12_31_424971 58 36 . . . 10_1101-2020_12_31_424971 59 1 163 163 CD 10_1101-2020_12_31_424971 59 2 In in IN 10_1101-2020_12_31_424971 59 3 the the DT 10_1101-2020_12_31_424971 59 4 m1G37-abundant m1G37-abundant '' 10_1101-2020_12_31_424971 59 5 ( ( -LRB- 10_1101-2020_12_31_424971 59 6 m1G37 m1G37 NNP 10_1101-2020_12_31_424971 59 7 + + SYM 10_1101-2020_12_31_424971 59 8 ) ) -RRB- 10_1101-2020_12_31_424971 59 9 condition condition NN 10_1101-2020_12_31_424971 59 10 , , , 10_1101-2020_12_31_424971 59 11 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 59 12 displayed display VBD 10_1101-2020_12_31_424971 59 13 a a DT 10_1101-2020_12_31_424971 59 14 high high JJ 10_1101-2020_12_31_424971 59 15 +1-frameshifting +1-frameshifting PRP$ 10_1101-2020_12_31_424971 59 16 164 164 CD 10_1101-2020_12_31_424971 59 17 efficiency efficiency NN 10_1101-2020_12_31_424971 59 18 ( ( -LRB- 10_1101-2020_12_31_424971 59 19 8.2 8.2 CD 10_1101-2020_12_31_424971 59 20 % % NN 10_1101-2020_12_31_424971 59 21 , , , 10_1101-2020_12_31_424971 59 22 Figure Figure NNP 10_1101-2020_12_31_424971 59 23 2a 2a CD 10_1101-2020_12_31_424971 59 24 ) ) -RRB- 10_1101-2020_12_31_424971 59 25 relative relative JJ 10_1101-2020_12_31_424971 59 26 to to IN 10_1101-2020_12_31_424971 59 27 ProL ProL NNP 10_1101-2020_12_31_424971 59 28 ( ( -LRB- 10_1101-2020_12_31_424971 59 29 1.4 1.4 CD 10_1101-2020_12_31_424971 59 30 % % NN 10_1101-2020_12_31_424971 59 31 ) ) -RRB- 10_1101-2020_12_31_424971 59 32 . . . 10_1101-2020_12_31_424971 60 1 In in IN 10_1101-2020_12_31_424971 60 2 the the DT 10_1101-2020_12_31_424971 60 3 m1G37-deficient m1G37-deficient , 10_1101-2020_12_31_424971 60 4 ( ( -LRB- 10_1101-2020_12_31_424971 60 5 m1G37 m1G37 NNP 10_1101-2020_12_31_424971 60 6 – – : 10_1101-2020_12_31_424971 60 7 ) ) -RRB- 10_1101-2020_12_31_424971 60 8 condition condition NN 10_1101-2020_12_31_424971 60 9 , , , 10_1101-2020_12_31_424971 60 10 165 165 CD 10_1101-2020_12_31_424971 60 11 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 60 12 exhibited exhibit VBD 10_1101-2020_12_31_424971 60 13 an an DT 10_1101-2020_12_31_424971 60 14 even even RB 10_1101-2020_12_31_424971 60 15 higher high JJR 10_1101-2020_12_31_424971 60 16 efficiency efficiency NN 10_1101-2020_12_31_424971 60 17 ( ( -LRB- 10_1101-2020_12_31_424971 60 18 20.8 20.8 CD 10_1101-2020_12_31_424971 60 19 % % NN 10_1101-2020_12_31_424971 60 20 ) ) -RRB- 10_1101-2020_12_31_424971 60 21 and and CC 10_1101-2020_12_31_424971 60 22 , , , 10_1101-2020_12_31_424971 60 23 consistent consistent JJ 10_1101-2020_12_31_424971 60 24 with with IN 10_1101-2020_12_31_424971 60 25 our -PRON- PRP$ 10_1101-2020_12_31_424971 60 26 previous previous JJ 10_1101-2020_12_31_424971 60 27 work29 work29 NNS 10_1101-2020_12_31_424971 60 28 , , , 10_1101-2020_12_31_424971 60 29 ProL ProL NNP 10_1101-2020_12_31_424971 60 30 166 166 CD 10_1101-2020_12_31_424971 60 31 also also RB 10_1101-2020_12_31_424971 60 32 displayed display VBD 10_1101-2020_12_31_424971 60 33 an an DT 10_1101-2020_12_31_424971 60 34 increased increase VBN 10_1101-2020_12_31_424971 60 35 efficiency efficiency NN 10_1101-2020_12_31_424971 60 36 ( ( -LRB- 10_1101-2020_12_31_424971 60 37 7.0 7.0 CD 10_1101-2020_12_31_424971 60 38 % % NN 10_1101-2020_12_31_424971 60 39 ) ) -RRB- 10_1101-2020_12_31_424971 60 40 relative relative JJ 10_1101-2020_12_31_424971 60 41 to to IN 10_1101-2020_12_31_424971 60 42 background background NN 10_1101-2020_12_31_424971 60 43 ( ( -LRB- 10_1101-2020_12_31_424971 60 44 1.4 1.4 CD 10_1101-2020_12_31_424971 60 45 % % NN 10_1101-2020_12_31_424971 60 46 ) ) -RRB- 10_1101-2020_12_31_424971 60 47 . . . 10_1101-2020_12_31_424971 61 1 Because because IN 10_1101-2020_12_31_424971 61 2 N1 N1 NNP 10_1101-2020_12_31_424971 61 3 - - HYPH 10_1101-2020_12_31_424971 61 4 167 167 CD 10_1101-2020_12_31_424971 61 5 methylation methylation NN 10_1101-2020_12_31_424971 61 6 in in IN 10_1101-2020_12_31_424971 61 7 the the DT 10_1101-2020_12_31_424971 61 8 m1G37 m1g37 NN 10_1101-2020_12_31_424971 61 9 + + SYM 10_1101-2020_12_31_424971 61 10 condition condition NN 10_1101-2020_12_31_424971 61 11 was be VBD 10_1101-2020_12_31_424971 61 12 stoichiometric stoichiometric JJ 10_1101-2020_12_31_424971 61 13 ( ( -LRB- 10_1101-2020_12_31_424971 61 14 Figure Figure NNP 10_1101-2020_12_31_424971 61 15 1c 1c NNP 10_1101-2020_12_31_424971 61 16 ) ) -RRB- 10_1101-2020_12_31_424971 61 17 , , , 10_1101-2020_12_31_424971 61 18 thereby thereby RB 10_1101-2020_12_31_424971 61 19 preventing prevent VBG 10_1101-2020_12_31_424971 61 20 168 168 CD 10_1101-2020_12_31_424971 61 21 quadruplet quadruplet NN 10_1101-2020_12_31_424971 61 22 - - HYPH 10_1101-2020_12_31_424971 61 23 pairing pair VBG 10_1101-2020_12_31_424971 61 24 , , , 10_1101-2020_12_31_424971 61 25 we -PRON- PRP 10_1101-2020_12_31_424971 61 26 attribute attribute VBP 10_1101-2020_12_31_424971 61 27 the the DT 10_1101-2020_12_31_424971 61 28 8.2 8.2 CD 10_1101-2020_12_31_424971 61 29 % % NN 10_1101-2020_12_31_424971 61 30 efficiency efficiency NN 10_1101-2020_12_31_424971 61 31 of of IN 10_1101-2020_12_31_424971 61 32 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 61 33 in in IN 10_1101-2020_12_31_424971 61 34 this this DT 10_1101-2020_12_31_424971 61 35 condition condition NN 10_1101-2020_12_31_424971 61 36 as as IN 10_1101-2020_12_31_424971 61 37 arising arise VBG 10_1101-2020_12_31_424971 61 38 exclusively exclusively RB 10_1101-2020_12_31_424971 61 39 169 169 CD 10_1101-2020_12_31_424971 61 40 from from IN 10_1101-2020_12_31_424971 61 41 triplet triplet NN 10_1101-2020_12_31_424971 61 42 - - HYPH 10_1101-2020_12_31_424971 61 43 slippage slippage NN 10_1101-2020_12_31_424971 61 44 . . . 10_1101-2020_12_31_424971 62 1 In in IN 10_1101-2020_12_31_424971 62 2 the the DT 10_1101-2020_12_31_424971 62 3 m1G37 m1g37 NN 10_1101-2020_12_31_424971 62 4 – – : 10_1101-2020_12_31_424971 62 5 condition condition NN 10_1101-2020_12_31_424971 62 6 , , , 10_1101-2020_12_31_424971 62 7 we -PRON- PRP 10_1101-2020_12_31_424971 62 8 observed observe VBD 10_1101-2020_12_31_424971 62 9 an an DT 10_1101-2020_12_31_424971 62 10 increase increase NN 10_1101-2020_12_31_424971 62 11 in in IN 10_1101-2020_12_31_424971 62 12 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 62 13 170 170 CD 10_1101-2020_12_31_424971 62 14 efficiency efficiency NN 10_1101-2020_12_31_424971 62 15 of of IN 10_1101-2020_12_31_424971 62 16 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 62 17 to to IN 10_1101-2020_12_31_424971 62 18 20.8 20.8 CD 10_1101-2020_12_31_424971 62 19 % % NN 10_1101-2020_12_31_424971 62 20 . . . 10_1101-2020_12_31_424971 63 1 While while IN 10_1101-2020_12_31_424971 63 2 multiple multiple JJ 10_1101-2020_12_31_424971 63 3 mechanisms mechanism NNS 10_1101-2020_12_31_424971 63 4 may may MD 10_1101-2020_12_31_424971 63 5 exist exist VB 10_1101-2020_12_31_424971 63 6 for for IN 10_1101-2020_12_31_424971 63 7 the the DT 10_1101-2020_12_31_424971 63 8 increased increase VBN 10_1101-2020_12_31_424971 63 9 +1 +1 NNP 10_1101-2020_12_31_424971 63 10 171 171 CD 10_1101-2020_12_31_424971 63 11 frameshifting frameshifting NN 10_1101-2020_12_31_424971 63 12 , , , 10_1101-2020_12_31_424971 63 13 the the DT 10_1101-2020_12_31_424971 63 14 exploration exploration NN 10_1101-2020_12_31_424971 63 15 of of IN 10_1101-2020_12_31_424971 63 16 both both CC 10_1101-2020_12_31_424971 63 17 triplet triplet NN 10_1101-2020_12_31_424971 63 18 - - HYPH 10_1101-2020_12_31_424971 63 19 slippage slippage NN 10_1101-2020_12_31_424971 63 20 and and CC 10_1101-2020_12_31_424971 63 21 quadruplet quadruplet NN 10_1101-2020_12_31_424971 63 22 - - HYPH 10_1101-2020_12_31_424971 63 23 pairing pairing NN 10_1101-2020_12_31_424971 63 24 is be VBZ 10_1101-2020_12_31_424971 63 25 one one CD 10_1101-2020_12_31_424971 63 26 possibility possibility NN 10_1101-2020_12_31_424971 63 27 . . . 10_1101-2020_12_31_424971 64 1 172 172 CD 10_1101-2020_12_31_424971 64 2 To to TO 10_1101-2020_12_31_424971 64 3 confirm confirm VB 10_1101-2020_12_31_424971 64 4 our -PRON- PRP$ 10_1101-2020_12_31_424971 64 5 results result NNS 10_1101-2020_12_31_424971 64 6 , , , 10_1101-2020_12_31_424971 64 7 we -PRON- PRP 10_1101-2020_12_31_424971 64 8 performed perform VBD 10_1101-2020_12_31_424971 64 9 similar similar JJ 10_1101-2020_12_31_424971 64 10 studies study NNS 10_1101-2020_12_31_424971 64 11 with with IN 10_1101-2020_12_31_424971 64 12 the the DT 10_1101-2020_12_31_424971 64 13 isogenic isogenic JJ 10_1101-2020_12_31_424971 64 14 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 64 15 and and CC 10_1101-2020_12_31_424971 64 16 ProL ProL NNP 10_1101-2020_12_31_424971 64 17 strains strain VBZ 10_1101-2020_12_31_424971 64 18 173 173 CD 10_1101-2020_12_31_424971 64 19 on on IN 10_1101-2020_12_31_424971 64 20 the the DT 10_1101-2020_12_31_424971 64 21 endogenous endogenous JJ 10_1101-2020_12_31_424971 64 22 E. E. NNP 10_1101-2020_12_31_424971 64 23 coli coli NNS 10_1101-2020_12_31_424971 64 24 lolB lolb NN 10_1101-2020_12_31_424971 64 25 gene gene NN 10_1101-2020_12_31_424971 64 26 , , , 10_1101-2020_12_31_424971 64 27 encoding encode VBG 10_1101-2020_12_31_424971 64 28 the the DT 10_1101-2020_12_31_424971 64 29 outer outer JJ 10_1101-2020_12_31_424971 64 30 membrane membrane NN 10_1101-2020_12_31_424971 64 31 lipoprotein lipoprotein NN 10_1101-2020_12_31_424971 64 32 . . . 10_1101-2020_12_31_424971 65 1 The the DT 10_1101-2020_12_31_424971 65 2 lolB lolB NNP 10_1101-2020_12_31_424971 65 3 gene gene NN 10_1101-2020_12_31_424971 65 4 174 174 CD 10_1101-2020_12_31_424971 65 5 naturally naturally RB 10_1101-2020_12_31_424971 65 6 contains contain VBZ 10_1101-2020_12_31_424971 65 7 a a DT 10_1101-2020_12_31_424971 65 8 CCC CCC NNP 10_1101-2020_12_31_424971 65 9 - - HYPH 10_1101-2020_12_31_424971 65 10 C C NNP 10_1101-2020_12_31_424971 65 11 motif motif NN 10_1101-2020_12_31_424971 65 12 at at IN 10_1101-2020_12_31_424971 65 13 the the DT 10_1101-2020_12_31_424971 65 14 2nd 2nd JJ 10_1101-2020_12_31_424971 65 15 codon codon NN 10_1101-2020_12_31_424971 65 16 position position NN 10_1101-2020_12_31_424971 65 17 such such JJ 10_1101-2020_12_31_424971 65 18 that that IN 10_1101-2020_12_31_424971 65 19 +1 +1 NNS 10_1101-2020_12_31_424971 65 20 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 65 21 at at IN 10_1101-2020_12_31_424971 65 22 this this DT 10_1101-2020_12_31_424971 65 23 motif motif NN 10_1101-2020_12_31_424971 65 24 175 175 CD 10_1101-2020_12_31_424971 65 25 would would MD 10_1101-2020_12_31_424971 65 26 decrease decrease VB 10_1101-2020_12_31_424971 65 27 protein protein NN 10_1101-2020_12_31_424971 65 28 synthesis synthesis NN 10_1101-2020_12_31_424971 65 29 due due JJ 10_1101-2020_12_31_424971 65 30 to to IN 10_1101-2020_12_31_424971 65 31 premature premature JJ 10_1101-2020_12_31_424971 65 32 termination termination NN 10_1101-2020_12_31_424971 65 33 . . . 10_1101-2020_12_31_424971 66 1 As as IN 10_1101-2020_12_31_424971 66 2 a a DT 10_1101-2020_12_31_424971 66 3 reference reference NN 10_1101-2020_12_31_424971 66 4 , , , 10_1101-2020_12_31_424971 66 5 we -PRON- PRP 10_1101-2020_12_31_424971 66 6 used use VBD 10_1101-2020_12_31_424971 66 7 E. E. NNP 10_1101-2020_12_31_424971 66 8 coli coli VBZ 10_1101-2020_12_31_424971 66 9 176 176 CD 10_1101-2020_12_31_424971 66 10 .CC .CC : 10_1101-2020_12_31_424971 66 11 - - HYPH 10_1101-2020_12_31_424971 66 12 BY by IN 10_1101-2020_12_31_424971 66 13 - - HYPH 10_1101-2020_12_31_424971 66 14 NC NC NNP 10_1101-2020_12_31_424971 66 15 - - HYPH 10_1101-2020_12_31_424971 66 16 ND ND NNP 10_1101-2020_12_31_424971 66 17 4.0 4.0 CD 10_1101-2020_12_31_424971 66 18 International International NNP 10_1101-2020_12_31_424971 66 19 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 66 20 under under IN 10_1101-2020_12_31_424971 66 21 a a DT 10_1101-2020_12_31_424971 66 22 ( ( -LRB- 10_1101-2020_12_31_424971 66 23 which which WDT 10_1101-2020_12_31_424971 66 24 was be VBD 10_1101-2020_12_31_424971 66 25 not not RB 10_1101-2020_12_31_424971 66 26 certified certify VBN 10_1101-2020_12_31_424971 66 27 by by IN 10_1101-2020_12_31_424971 66 28 peer peer NN 10_1101-2020_12_31_424971 66 29 review review NN 10_1101-2020_12_31_424971 66 30 ) ) -RRB- 10_1101-2020_12_31_424971 66 31 is be VBZ 10_1101-2020_12_31_424971 66 32 the the DT 10_1101-2020_12_31_424971 66 33 author author NN 10_1101-2020_12_31_424971 66 34 / / SYM 10_1101-2020_12_31_424971 66 35 funder funder NN 10_1101-2020_12_31_424971 66 36 , , , 10_1101-2020_12_31_424971 66 37 who who WP 10_1101-2020_12_31_424971 66 38 has have VBZ 10_1101-2020_12_31_424971 66 39 granted grant VBN 10_1101-2020_12_31_424971 66 40 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 66 41 a a DT 10_1101-2020_12_31_424971 66 42 license license NN 10_1101-2020_12_31_424971 66 43 to to TO 10_1101-2020_12_31_424971 66 44 display display VB 10_1101-2020_12_31_424971 66 45 the the DT 10_1101-2020_12_31_424971 66 46 preprint preprint NN 10_1101-2020_12_31_424971 66 47 in in IN 10_1101-2020_12_31_424971 66 48 perpetuity perpetuity NN 10_1101-2020_12_31_424971 66 49 . . . 10_1101-2020_12_31_424971 67 1 It -PRON- PRP 10_1101-2020_12_31_424971 67 2 is be VBZ 10_1101-2020_12_31_424971 67 3 made make VBN 10_1101-2020_12_31_424971 67 4 The the DT 10_1101-2020_12_31_424971 67 5 copyright copyright NN 10_1101-2020_12_31_424971 67 6 holder holder NN 10_1101-2020_12_31_424971 67 7 for for IN 10_1101-2020_12_31_424971 67 8 this this DT 10_1101-2020_12_31_424971 67 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 67 10 version version NN 10_1101-2020_12_31_424971 67 11 posted post VBD 10_1101-2020_12_31_424971 67 12 January January NNP 10_1101-2020_12_31_424971 67 13 2 2 CD 10_1101-2020_12_31_424971 67 14 , , , 10_1101-2020_12_31_424971 67 15 2021 2021 CD 10_1101-2020_12_31_424971 67 16 . . . 10_1101-2020_12_31_424971 67 17 ; ; : 10_1101-2020_12_31_424971 67 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 67 19 : : : 10_1101-2020_12_31_424971 67 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 67 21 preprint preprint NN 10_1101-2020_12_31_424971 67 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 67 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 67 24 8 8 CD 10_1101-2020_12_31_424971 67 25 cysS cys NNS 10_1101-2020_12_31_424971 67 26 , , , 10_1101-2020_12_31_424971 67 27 encoding encode VBG 10_1101-2020_12_31_424971 67 28 cysteinyl cysteinyl NN 10_1101-2020_12_31_424971 67 29 - - HYPH 10_1101-2020_12_31_424971 67 30 tRNA trna NN 10_1101-2020_12_31_424971 67 31 synthetase synthetase NN 10_1101-2020_12_31_424971 67 32 ( ( -LRB- 10_1101-2020_12_31_424971 67 33 CysRS)30 CysRS)30 NNS 10_1101-2020_12_31_424971 67 34 , , , 10_1101-2020_12_31_424971 67 35 which which WDT 10_1101-2020_12_31_424971 67 36 has have VBZ 10_1101-2020_12_31_424971 67 37 no no DT 10_1101-2020_12_31_424971 67 38 CCC CCC NNP 10_1101-2020_12_31_424971 67 39 - - HYPH 10_1101-2020_12_31_424971 67 40 C C NNP 10_1101-2020_12_31_424971 67 41 motif motif NN 10_1101-2020_12_31_424971 67 42 in in IN 10_1101-2020_12_31_424971 67 43 the the DT 10_1101-2020_12_31_424971 67 44 first first JJ 10_1101-2020_12_31_424971 67 45 16 16 CD 10_1101-2020_12_31_424971 67 46 177 177 CD 10_1101-2020_12_31_424971 67 47 codons codon NNS 10_1101-2020_12_31_424971 67 48 and and CC 10_1101-2020_12_31_424971 67 49 would would MD 10_1101-2020_12_31_424971 67 50 be be VB 10_1101-2020_12_31_424971 67 51 less less RBR 10_1101-2020_12_31_424971 67 52 sensitive sensitive JJ 10_1101-2020_12_31_424971 67 53 to to IN 10_1101-2020_12_31_424971 67 54 +1 +1 NNS 10_1101-2020_12_31_424971 67 55 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 67 56 at at IN 10_1101-2020_12_31_424971 67 57 CCC CCC NNP 10_1101-2020_12_31_424971 67 58 - - HYPH 10_1101-2020_12_31_424971 67 59 C C NNP 10_1101-2020_12_31_424971 67 60 motifs motif NNS 10_1101-2020_12_31_424971 67 61 during during IN 10_1101-2020_12_31_424971 67 62 protein protein NN 10_1101-2020_12_31_424971 67 63 synthesis synthesis NN 10_1101-2020_12_31_424971 67 64 . . . 10_1101-2020_12_31_424971 68 1 178 178 CD 10_1101-2020_12_31_424971 68 2 The the DT 10_1101-2020_12_31_424971 68 3 ratio ratio NN 10_1101-2020_12_31_424971 68 4 of of IN 10_1101-2020_12_31_424971 68 5 protein protein NN 10_1101-2020_12_31_424971 68 6 synthesis synthesis NN 10_1101-2020_12_31_424971 68 7 of of IN 10_1101-2020_12_31_424971 68 8 lolB lolB NNP 10_1101-2020_12_31_424971 68 9 to to IN 10_1101-2020_12_31_424971 68 10 cysS cys NNS 10_1101-2020_12_31_424971 68 11 for for IN 10_1101-2020_12_31_424971 68 12 the the DT 10_1101-2020_12_31_424971 68 13 control control NN 10_1101-2020_12_31_424971 68 14 sample sample NN 10_1101-2020_12_31_424971 68 15 ProL ProL NNP 10_1101-2020_12_31_424971 68 16 in in IN 10_1101-2020_12_31_424971 68 17 the the DT 10_1101-2020_12_31_424971 68 18 m1G37 m1g37 JJ 10_1101-2020_12_31_424971 68 19 condition condition NN 10_1101-2020_12_31_424971 68 20 , , , 10_1101-2020_12_31_424971 68 21 179 179 CD 10_1101-2020_12_31_424971 68 22 measured measure VBN 10_1101-2020_12_31_424971 68 23 from from IN 10_1101-2020_12_31_424971 68 24 Western western JJ 10_1101-2020_12_31_424971 68 25 blots blot NNS 10_1101-2020_12_31_424971 68 26 ( ( -LRB- 10_1101-2020_12_31_424971 68 27 Methods Methods NNP 10_1101-2020_12_31_424971 68 28 ) ) -RRB- 10_1101-2020_12_31_424971 68 29 , , , 10_1101-2020_12_31_424971 68 30 was be VBD 10_1101-2020_12_31_424971 68 31 normalized normalize VBN 10_1101-2020_12_31_424971 68 32 to to IN 10_1101-2020_12_31_424971 68 33 1.00 1.00 CD 10_1101-2020_12_31_424971 68 34 , , , 10_1101-2020_12_31_424971 68 35 denoting denote VBG 10_1101-2020_12_31_424971 68 36 that that IN 10_1101-2020_12_31_424971 68 37 lolB lolb NN 10_1101-2020_12_31_424971 68 38 and and CC 10_1101-2020_12_31_424971 68 39 cysS cyss IN 10_1101-2020_12_31_424971 68 40 180 180 CD 10_1101-2020_12_31_424971 68 41 were be VBD 10_1101-2020_12_31_424971 68 42 maximally maximally RB 10_1101-2020_12_31_424971 68 43 translated translate VBN 10_1101-2020_12_31_424971 68 44 in in IN 10_1101-2020_12_31_424971 68 45 the the DT 10_1101-2020_12_31_424971 68 46 0-frame 0-frame CD 10_1101-2020_12_31_424971 68 47 without without IN 10_1101-2020_12_31_424971 68 48 +1 +1 NNS 10_1101-2020_12_31_424971 68 49 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 68 50 ( ( -LRB- 10_1101-2020_12_31_424971 68 51 i.e. i.e. FW 10_1101-2020_12_31_424971 68 52 , , , 10_1101-2020_12_31_424971 68 53 a a DT 10_1101-2020_12_31_424971 68 54 relative relative JJ 10_1101-2020_12_31_424971 68 55 +1 +1 NNS 10_1101-2020_12_31_424971 68 56 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 68 57 181 181 CD 10_1101-2020_12_31_424971 68 58 efficiency efficiency NN 10_1101-2020_12_31_424971 68 59 of of IN 10_1101-2020_12_31_424971 68 60 0.00 0.00 CD 10_1101-2020_12_31_424971 68 61 ) ) -RRB- 10_1101-2020_12_31_424971 68 62 ( ( -LRB- 10_1101-2020_12_31_424971 68 63 Figures figure NNS 10_1101-2020_12_31_424971 68 64 2b 2b NNP 10_1101-2020_12_31_424971 68 65 , , , 10_1101-2020_12_31_424971 68 66 2c 2c NN 10_1101-2020_12_31_424971 68 67 ) ) -RRB- 10_1101-2020_12_31_424971 68 68 . . . 10_1101-2020_12_31_424971 69 1 In in IN 10_1101-2020_12_31_424971 69 2 the the DT 10_1101-2020_12_31_424971 69 3 m1G37 m1g37 NN 10_1101-2020_12_31_424971 69 4 + + CD 10_1101-2020_12_31_424971 69 5 condition condition NN 10_1101-2020_12_31_424971 69 6 , , , 10_1101-2020_12_31_424971 69 7 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 69 8 displayed display VBD 10_1101-2020_12_31_424971 69 9 a a DT 10_1101-2020_12_31_424971 69 10 ratio ratio NN 10_1101-2020_12_31_424971 69 11 of of IN 10_1101-2020_12_31_424971 69 12 LolB LolB NNP 10_1101-2020_12_31_424971 69 13 to to IN 10_1101-2020_12_31_424971 69 14 182 182 CD 10_1101-2020_12_31_424971 69 15 CysRS CysRS NNP 10_1101-2020_12_31_424971 69 16 of of IN 10_1101-2020_12_31_424971 69 17 0.62 0.62 CD 10_1101-2020_12_31_424971 69 18 , , , 10_1101-2020_12_31_424971 69 19 indicating indicate VBG 10_1101-2020_12_31_424971 69 20 an an DT 10_1101-2020_12_31_424971 69 21 increase increase NN 10_1101-2020_12_31_424971 69 22 in in IN 10_1101-2020_12_31_424971 69 23 the the DT 10_1101-2020_12_31_424971 69 24 relative relative JJ 10_1101-2020_12_31_424971 69 25 +1 +1 : 10_1101-2020_12_31_424971 69 26 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 69 27 efficiency efficiency NN 10_1101-2020_12_31_424971 69 28 to to IN 10_1101-2020_12_31_424971 69 29 0.38 0.38 CD 10_1101-2020_12_31_424971 69 30 , , , 10_1101-2020_12_31_424971 69 31 and and CC 10_1101-2020_12_31_424971 69 32 in in IN 10_1101-2020_12_31_424971 69 33 the the DT 10_1101-2020_12_31_424971 69 34 183 183 CD 10_1101-2020_12_31_424971 69 35 m1G37 m1g37 NN 10_1101-2020_12_31_424971 69 36 – – : 10_1101-2020_12_31_424971 69 37 condition condition NN 10_1101-2020_12_31_424971 69 38 , , , 10_1101-2020_12_31_424971 69 39 it -PRON- PRP 10_1101-2020_12_31_424971 69 40 displayed display VBD 10_1101-2020_12_31_424971 69 41 a a DT 10_1101-2020_12_31_424971 69 42 ratio ratio NN 10_1101-2020_12_31_424971 69 43 of of IN 10_1101-2020_12_31_424971 69 44 0.17 0.17 CD 10_1101-2020_12_31_424971 69 45 , , , 10_1101-2020_12_31_424971 69 46 indicating indicate VBG 10_1101-2020_12_31_424971 69 47 an an DT 10_1101-2020_12_31_424971 69 48 increase increase NN 10_1101-2020_12_31_424971 69 49 in in IN 10_1101-2020_12_31_424971 69 50 the the DT 10_1101-2020_12_31_424971 69 51 relative relative JJ 10_1101-2020_12_31_424971 69 52 +1 +1 : 10_1101-2020_12_31_424971 69 53 184 184 CD 10_1101-2020_12_31_424971 69 54 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 69 55 efficiency efficiency NN 10_1101-2020_12_31_424971 69 56 to to IN 10_1101-2020_12_31_424971 69 57 0.83 0.83 CD 10_1101-2020_12_31_424971 69 58 ( ( -LRB- 10_1101-2020_12_31_424971 69 59 Figures figure NNS 10_1101-2020_12_31_424971 69 60 2b 2b NNP 10_1101-2020_12_31_424971 69 61 , , , 10_1101-2020_12_31_424971 69 62 2c 2c NN 10_1101-2020_12_31_424971 69 63 ) ) -RRB- 10_1101-2020_12_31_424971 69 64 . . . 10_1101-2020_12_31_424971 70 1 Similarly similarly RB 10_1101-2020_12_31_424971 70 2 , , , 10_1101-2020_12_31_424971 70 3 ProL ProL NNP 10_1101-2020_12_31_424971 70 4 in in IN 10_1101-2020_12_31_424971 70 5 the the DT 10_1101-2020_12_31_424971 70 6 m1G37 m1g37 NN 10_1101-2020_12_31_424971 70 7 – – : 10_1101-2020_12_31_424971 70 8 condition condition NN 10_1101-2020_12_31_424971 70 9 displayed display VBD 10_1101-2020_12_31_424971 70 10 185 185 CD 10_1101-2020_12_31_424971 70 11 a a DT 10_1101-2020_12_31_424971 70 12 ratio ratio NN 10_1101-2020_12_31_424971 70 13 of of IN 10_1101-2020_12_31_424971 70 14 LolB LolB NNP 10_1101-2020_12_31_424971 70 15 to to IN 10_1101-2020_12_31_424971 70 16 CysRS CysRS NNP 10_1101-2020_12_31_424971 70 17 of of IN 10_1101-2020_12_31_424971 70 18 0.47 0.47 CD 10_1101-2020_12_31_424971 70 19 , , , 10_1101-2020_12_31_424971 70 20 indicating indicate VBG 10_1101-2020_12_31_424971 70 21 an an DT 10_1101-2020_12_31_424971 70 22 increase increase NN 10_1101-2020_12_31_424971 70 23 in in IN 10_1101-2020_12_31_424971 70 24 the the DT 10_1101-2020_12_31_424971 70 25 +1-frameshifting +1-frameshifting JJ 10_1101-2020_12_31_424971 70 26 efficiency efficiency NN 10_1101-2020_12_31_424971 70 27 to to TO 10_1101-2020_12_31_424971 70 28 0.53 0.53 CD 10_1101-2020_12_31_424971 70 29 . . . 10_1101-2020_12_31_424971 71 1 186 186 CD 10_1101-2020_12_31_424971 71 2 187 187 CD 10_1101-2020_12_31_424971 71 3 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 71 4 can can MD 10_1101-2020_12_31_424971 71 5 insert insert VB 10_1101-2020_12_31_424971 71 6 non non JJ 10_1101-2020_12_31_424971 71 7 - - JJ 10_1101-2020_12_31_424971 71 8 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 71 9 amino amino JJ 10_1101-2020_12_31_424971 71 10 acids acid NNS 10_1101-2020_12_31_424971 71 11 at at IN 10_1101-2020_12_31_424971 71 12 CCC CCC NNP 10_1101-2020_12_31_424971 71 13 - - HYPH 10_1101-2020_12_31_424971 71 14 C C NNP 10_1101-2020_12_31_424971 71 15 motifs motif VBZ 10_1101-2020_12_31_424971 71 16 188 188 CD 10_1101-2020_12_31_424971 71 17 We -PRON- PRP 10_1101-2020_12_31_424971 71 18 next next RB 10_1101-2020_12_31_424971 71 19 asked ask VBD 10_1101-2020_12_31_424971 71 20 whether whether IN 10_1101-2020_12_31_424971 71 21 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 71 22 can can MD 10_1101-2020_12_31_424971 71 23 deliver deliver VB 10_1101-2020_12_31_424971 71 24 non non JJ 10_1101-2020_12_31_424971 71 25 - - JJ 10_1101-2020_12_31_424971 71 26 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 71 27 amino amino JJ 10_1101-2020_12_31_424971 71 28 acids acid NNS 10_1101-2020_12_31_424971 71 29 to to IN 10_1101-2020_12_31_424971 71 30 the the DT 10_1101-2020_12_31_424971 71 31 ribosome ribosome NN 10_1101-2020_12_31_424971 71 32 by by IN 10_1101-2020_12_31_424971 71 33 189 189 CD 10_1101-2020_12_31_424971 71 34 inducing induce VBG 10_1101-2020_12_31_424971 71 35 +1 +1 NNS 10_1101-2020_12_31_424971 71 36 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 71 37 at at IN 10_1101-2020_12_31_424971 71 38 a a DT 10_1101-2020_12_31_424971 71 39 CCC CCC NNP 10_1101-2020_12_31_424971 71 40 - - HYPH 10_1101-2020_12_31_424971 71 41 C C NNP 10_1101-2020_12_31_424971 71 42 motif motif NN 10_1101-2020_12_31_424971 71 43 ( ( -LRB- 10_1101-2020_12_31_424971 71 44 Figure Figure NNP 10_1101-2020_12_31_424971 71 45 2d 2d NNP 10_1101-2020_12_31_424971 71 46 ) ) -RRB- 10_1101-2020_12_31_424971 71 47 . . . 10_1101-2020_12_31_424971 72 1 We -PRON- PRP 10_1101-2020_12_31_424971 72 2 inserted insert VBD 10_1101-2020_12_31_424971 72 3 a a DT 10_1101-2020_12_31_424971 72 4 CCC CCC NNP 10_1101-2020_12_31_424971 72 5 - - HYPH 10_1101-2020_12_31_424971 72 6 C C NNP 10_1101-2020_12_31_424971 72 7 motif motif NN 10_1101-2020_12_31_424971 72 8 at at IN 10_1101-2020_12_31_424971 72 9 the the DT 10_1101-2020_12_31_424971 72 10 5th 5th JJ 10_1101-2020_12_31_424971 72 11 190 190 CD 10_1101-2020_12_31_424971 72 12 codon codon NN 10_1101-2020_12_31_424971 72 13 position position NN 10_1101-2020_12_31_424971 72 14 of of IN 10_1101-2020_12_31_424971 72 15 the the DT 10_1101-2020_12_31_424971 72 16 E. E. NNP 10_1101-2020_12_31_424971 72 17 coli coli NNS 10_1101-2020_12_31_424971 72 18 folA fola JJ 10_1101-2020_12_31_424971 72 19 gene gene NN 10_1101-2020_12_31_424971 72 20 , , , 10_1101-2020_12_31_424971 72 21 encoding encode VBG 10_1101-2020_12_31_424971 72 22 dihydrofolate dihydrofolate JJ 10_1101-2020_12_31_424971 72 23 reductase reductase NN 10_1101-2020_12_31_424971 72 24 ( ( -LRB- 10_1101-2020_12_31_424971 72 25 DHFR DHFR NNP 10_1101-2020_12_31_424971 72 26 ) ) -RRB- 10_1101-2020_12_31_424971 72 27 . . . 10_1101-2020_12_31_424971 73 1 A a DT 10_1101-2020_12_31_424971 73 2 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 73 3 - - SYM 10_1101-2020_12_31_424971 73 4 191 191 CD 10_1101-2020_12_31_424971 73 5 induced induce VBN 10_1101-2020_12_31_424971 73 6 +1 +1 NNS 10_1101-2020_12_31_424971 73 7 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 73 8 event event NN 10_1101-2020_12_31_424971 73 9 at at IN 10_1101-2020_12_31_424971 73 10 the the DT 10_1101-2020_12_31_424971 73 11 insertion insertion NN 10_1101-2020_12_31_424971 73 12 would would MD 10_1101-2020_12_31_424971 73 13 result result VB 10_1101-2020_12_31_424971 73 14 in in IN 10_1101-2020_12_31_424971 73 15 full full JJ 10_1101-2020_12_31_424971 73 16 - - HYPH 10_1101-2020_12_31_424971 73 17 length length NN 10_1101-2020_12_31_424971 73 18 DHFR DHFR NNP 10_1101-2020_12_31_424971 73 19 , , , 10_1101-2020_12_31_424971 73 20 whereas whereas IN 10_1101-2020_12_31_424971 73 21 the the DT 10_1101-2020_12_31_424971 73 22 192 192 CD 10_1101-2020_12_31_424971 73 23 absence absence NN 10_1101-2020_12_31_424971 73 24 of of IN 10_1101-2020_12_31_424971 73 25 +1 +1 NNS 10_1101-2020_12_31_424971 73 26 frameshifting frameshifting NN 10_1101-2020_12_31_424971 73 27 would would MD 10_1101-2020_12_31_424971 73 28 result result VB 10_1101-2020_12_31_424971 73 29 in in IN 10_1101-2020_12_31_424971 73 30 a a DT 10_1101-2020_12_31_424971 73 31 C c NN 10_1101-2020_12_31_424971 73 32 - - HYPH 10_1101-2020_12_31_424971 73 33 terminal terminal JJ 10_1101-2020_12_31_424971 73 34 truncated truncate VBN 10_1101-2020_12_31_424971 73 35 DHFR DHFR NNP 10_1101-2020_12_31_424971 73 36 fragment fragment NN 10_1101-2020_12_31_424971 73 37 ( ( -LRB- 10_1101-2020_12_31_424971 73 38 DC DC NNP 10_1101-2020_12_31_424971 73 39 ) ) -RRB- 10_1101-2020_12_31_424971 73 40 . . . 10_1101-2020_12_31_424971 74 1 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 74 2 193 193 CD 10_1101-2020_12_31_424971 74 3 was be VBD 10_1101-2020_12_31_424971 74 4 aminoacylated aminoacylate VBN 10_1101-2020_12_31_424971 74 5 with with IN 10_1101-2020_12_31_424971 74 6 non non JJ 10_1101-2020_12_31_424971 74 7 - - JJ 10_1101-2020_12_31_424971 74 8 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 74 9 amino amino JJ 10_1101-2020_12_31_424971 74 10 acids acid NNS 10_1101-2020_12_31_424971 74 11 using use VBG 10_1101-2020_12_31_424971 74 12 a a DT 10_1101-2020_12_31_424971 74 13 Flexizyme32 Flexizyme32 NNP 10_1101-2020_12_31_424971 74 14 and and CC 10_1101-2020_12_31_424971 74 15 subsequently subsequently RB 10_1101-2020_12_31_424971 74 16 194 194 CD 10_1101-2020_12_31_424971 74 17 tested test VBN 10_1101-2020_12_31_424971 74 18 in in IN 10_1101-2020_12_31_424971 74 19 [ [ -LRB- 10_1101-2020_12_31_424971 74 20 35S]-Met 35s]-met CD 10_1101-2020_12_31_424971 74 21 - - HYPH 10_1101-2020_12_31_424971 74 22 dependent dependent JJ 10_1101-2020_12_31_424971 74 23 in in FW 10_1101-2020_12_31_424971 74 24 vitro vitro FW 10_1101-2020_12_31_424971 74 25 translation translation NN 10_1101-2020_12_31_424971 74 26 reactions reaction NNS 10_1101-2020_12_31_424971 74 27 using use VBG 10_1101-2020_12_31_424971 74 28 the the DT 10_1101-2020_12_31_424971 74 29 E. E. NNP 10_1101-2020_12_31_424971 74 30 coli coli NNS 10_1101-2020_12_31_424971 74 31 PURExpress purexpress NN 10_1101-2020_12_31_424971 74 32 system system NN 10_1101-2020_12_31_424971 74 33 . . . 10_1101-2020_12_31_424971 75 1 195 195 CD 10_1101-2020_12_31_424971 75 2 The the DT 10_1101-2020_12_31_424971 75 3 resulting result VBG 10_1101-2020_12_31_424971 75 4 protein protein NN 10_1101-2020_12_31_424971 75 5 products product NNS 10_1101-2020_12_31_424971 75 6 were be VBD 10_1101-2020_12_31_424971 75 7 separated separate VBN 10_1101-2020_12_31_424971 75 8 by by IN 10_1101-2020_12_31_424971 75 9 sodium sodium VBG 10_1101-2020_12_31_424971 75 10 dodecyl dodecyl JJ 10_1101-2020_12_31_424971 75 11 sulfate sulfate NN 10_1101-2020_12_31_424971 75 12 ( ( -LRB- 10_1101-2020_12_31_424971 75 13 SDS)-polyacrylamide sds)-polyacrylamide NN 10_1101-2020_12_31_424971 75 14 196 196 CD 10_1101-2020_12_31_424971 75 15 gel gel NN 10_1101-2020_12_31_424971 75 16 electrophoresis electrophoresis NN 10_1101-2020_12_31_424971 75 17 and and CC 10_1101-2020_12_31_424971 75 18 quantified quantify VBN 10_1101-2020_12_31_424971 75 19 by by IN 10_1101-2020_12_31_424971 75 20 phosphorimaging phosphorimage VBG 10_1101-2020_12_31_424971 75 21 . . . 10_1101-2020_12_31_424971 76 1 Control control NN 10_1101-2020_12_31_424971 76 2 experiments experiment NNS 10_1101-2020_12_31_424971 76 3 with with IN 10_1101-2020_12_31_424971 76 4 no no DT 10_1101-2020_12_31_424971 76 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 76 6 or or CC 10_1101-2020_12_31_424971 76 7 197 197 CD 10_1101-2020_12_31_424971 76 8 with with IN 10_1101-2020_12_31_424971 76 9 a a DT 10_1101-2020_12_31_424971 76 10 non non JJ 10_1101-2020_12_31_424971 76 11 - - HYPH 10_1101-2020_12_31_424971 76 12 acylated acylated JJ 10_1101-2020_12_31_424971 76 13 SufB2 sufb2 NN 10_1101-2020_12_31_424971 76 14 showed show VBD 10_1101-2020_12_31_424971 76 15 no no DT 10_1101-2020_12_31_424971 76 16 full full JJ 10_1101-2020_12_31_424971 76 17 - - HYPH 10_1101-2020_12_31_424971 76 18 length length NN 10_1101-2020_12_31_424971 76 19 DHFR DHFR NNP 10_1101-2020_12_31_424971 76 20 , , , 10_1101-2020_12_31_424971 76 21 demonstrating demonstrate VBG 10_1101-2020_12_31_424971 76 22 that that IN 10_1101-2020_12_31_424971 76 23 synthesis synthesis NN 10_1101-2020_12_31_424971 76 24 of of IN 10_1101-2020_12_31_424971 76 25 full-198 full-198 NN 10_1101-2020_12_31_424971 76 26 length length NN 10_1101-2020_12_31_424971 76 27 DHFR DHFR NNP 10_1101-2020_12_31_424971 76 28 depended depend VBD 10_1101-2020_12_31_424971 76 29 upon upon IN 10_1101-2020_12_31_424971 76 30 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 76 31 delivery delivery NN 10_1101-2020_12_31_424971 76 32 of of IN 10_1101-2020_12_31_424971 76 33 an an DT 10_1101-2020_12_31_424971 76 34 amino amino JJ 10_1101-2020_12_31_424971 76 35 acid acid NN 10_1101-2020_12_31_424971 76 36 as as IN 10_1101-2020_12_31_424971 76 37 a a DT 10_1101-2020_12_31_424971 76 38 result result NN 10_1101-2020_12_31_424971 76 39 of of IN 10_1101-2020_12_31_424971 76 40 +1 +1 NNS 10_1101-2020_12_31_424971 76 41 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 76 42 at at IN 10_1101-2020_12_31_424971 76 43 199 199 CD 10_1101-2020_12_31_424971 76 44 the the DT 10_1101-2020_12_31_424971 76 45 CCC CCC NNP 10_1101-2020_12_31_424971 76 46 - - HYPH 10_1101-2020_12_31_424971 76 47 C C NNP 10_1101-2020_12_31_424971 76 48 motif motif NN 10_1101-2020_12_31_424971 76 49 . . . 10_1101-2020_12_31_424971 77 1 We -PRON- PRP 10_1101-2020_12_31_424971 77 2 showed show VBD 10_1101-2020_12_31_424971 77 3 that that IN 10_1101-2020_12_31_424971 77 4 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 77 5 was be VBD 10_1101-2020_12_31_424971 77 6 able able JJ 10_1101-2020_12_31_424971 77 7 to to TO 10_1101-2020_12_31_424971 77 8 deliver deliver VB 10_1101-2020_12_31_424971 77 9 Pro Pro NNP 10_1101-2020_12_31_424971 77 10 , , , 10_1101-2020_12_31_424971 77 11 Arg Arg NNP 10_1101-2020_12_31_424971 77 12 , , , 10_1101-2020_12_31_424971 77 13 Val Val NNP 10_1101-2020_12_31_424971 77 14 , , , 10_1101-2020_12_31_424971 77 15 and and CC 10_1101-2020_12_31_424971 77 16 the the DT 10_1101-2020_12_31_424971 77 17 Pro Pro NNP 10_1101-2020_12_31_424971 77 18 analogs analog VBZ 10_1101-2020_12_31_424971 77 19 200 200 CD 10_1101-2020_12_31_424971 77 20 cis cis NN 10_1101-2020_12_31_424971 77 21 - - HYPH 10_1101-2020_12_31_424971 77 22 hydroxypro hydroxypro JJ 10_1101-2020_12_31_424971 77 23 , , , 10_1101-2020_12_31_424971 77 24 trans trans NNP 10_1101-2020_12_31_424971 77 25 - - NNP 10_1101-2020_12_31_424971 77 26 hydroxypro hydroxypro JJ 10_1101-2020_12_31_424971 77 27 , , , 10_1101-2020_12_31_424971 77 28 azetidine azetidine NN 10_1101-2020_12_31_424971 77 29 , , , 10_1101-2020_12_31_424971 77 30 and and CC 10_1101-2020_12_31_424971 77 31 thiapro thiapro NN 10_1101-2020_12_31_424971 77 32 ( ( -LRB- 10_1101-2020_12_31_424971 77 33 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 77 34 Figure Figure NNP 10_1101-2020_12_31_424971 77 35 1 1 CD 10_1101-2020_12_31_424971 77 36 ) ) -RRB- 10_1101-2020_12_31_424971 77 37 to to IN 10_1101-2020_12_31_424971 77 38 the the DT 10_1101-2020_12_31_424971 77 39 201 201 CD 10_1101-2020_12_31_424971 77 40 ribosome ribosome NN 10_1101-2020_12_31_424971 77 41 in in IN 10_1101-2020_12_31_424971 77 42 response response NN 10_1101-2020_12_31_424971 77 43 to to IN 10_1101-2020_12_31_424971 77 44 the the DT 10_1101-2020_12_31_424971 77 45 CCC CCC NNP 10_1101-2020_12_31_424971 77 46 - - HYPH 10_1101-2020_12_31_424971 77 47 C C NNP 10_1101-2020_12_31_424971 77 48 motif motif NN 10_1101-2020_12_31_424971 77 49 , , , 10_1101-2020_12_31_424971 77 50 and and CC 10_1101-2020_12_31_424971 77 51 that that IN 10_1101-2020_12_31_424971 77 52 the the DT 10_1101-2020_12_31_424971 77 53 efficiency efficiency NN 10_1101-2020_12_31_424971 77 54 of of IN 10_1101-2020_12_31_424971 77 55 delivery delivery NN 10_1101-2020_12_31_424971 77 56 by by IN 10_1101-2020_12_31_424971 77 57 G37-state G37-state NNP 10_1101-2020_12_31_424971 77 58 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 77 59 202 202 CD 10_1101-2020_12_31_424971 77 60 .CC .CC : 10_1101-2020_12_31_424971 77 61 - - HYPH 10_1101-2020_12_31_424971 77 62 BY by IN 10_1101-2020_12_31_424971 77 63 - - HYPH 10_1101-2020_12_31_424971 77 64 NC NC NNP 10_1101-2020_12_31_424971 77 65 - - HYPH 10_1101-2020_12_31_424971 77 66 ND ND NNP 10_1101-2020_12_31_424971 77 67 4.0 4.0 CD 10_1101-2020_12_31_424971 77 68 International International NNP 10_1101-2020_12_31_424971 77 69 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 77 70 under under IN 10_1101-2020_12_31_424971 77 71 a a DT 10_1101-2020_12_31_424971 77 72 ( ( -LRB- 10_1101-2020_12_31_424971 77 73 which which WDT 10_1101-2020_12_31_424971 77 74 was be VBD 10_1101-2020_12_31_424971 77 75 not not RB 10_1101-2020_12_31_424971 77 76 certified certify VBN 10_1101-2020_12_31_424971 77 77 by by IN 10_1101-2020_12_31_424971 77 78 peer peer NN 10_1101-2020_12_31_424971 77 79 review review NN 10_1101-2020_12_31_424971 77 80 ) ) -RRB- 10_1101-2020_12_31_424971 77 81 is be VBZ 10_1101-2020_12_31_424971 77 82 the the DT 10_1101-2020_12_31_424971 77 83 author author NN 10_1101-2020_12_31_424971 77 84 / / SYM 10_1101-2020_12_31_424971 77 85 funder funder NN 10_1101-2020_12_31_424971 77 86 , , , 10_1101-2020_12_31_424971 77 87 who who WP 10_1101-2020_12_31_424971 77 88 has have VBZ 10_1101-2020_12_31_424971 77 89 granted grant VBN 10_1101-2020_12_31_424971 77 90 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 77 91 a a DT 10_1101-2020_12_31_424971 77 92 license license NN 10_1101-2020_12_31_424971 77 93 to to TO 10_1101-2020_12_31_424971 77 94 display display VB 10_1101-2020_12_31_424971 77 95 the the DT 10_1101-2020_12_31_424971 77 96 preprint preprint NN 10_1101-2020_12_31_424971 77 97 in in IN 10_1101-2020_12_31_424971 77 98 perpetuity perpetuity NN 10_1101-2020_12_31_424971 77 99 . . . 10_1101-2020_12_31_424971 78 1 It -PRON- PRP 10_1101-2020_12_31_424971 78 2 is be VBZ 10_1101-2020_12_31_424971 78 3 made make VBN 10_1101-2020_12_31_424971 78 4 The the DT 10_1101-2020_12_31_424971 78 5 copyright copyright NN 10_1101-2020_12_31_424971 78 6 holder holder NN 10_1101-2020_12_31_424971 78 7 for for IN 10_1101-2020_12_31_424971 78 8 this this DT 10_1101-2020_12_31_424971 78 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 78 10 version version NN 10_1101-2020_12_31_424971 78 11 posted post VBD 10_1101-2020_12_31_424971 78 12 January January NNP 10_1101-2020_12_31_424971 78 13 2 2 CD 10_1101-2020_12_31_424971 78 14 , , , 10_1101-2020_12_31_424971 78 15 2021 2021 CD 10_1101-2020_12_31_424971 78 16 . . . 10_1101-2020_12_31_424971 78 17 ; ; : 10_1101-2020_12_31_424971 78 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 78 19 : : : 10_1101-2020_12_31_424971 78 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 78 21 preprint preprint NN 10_1101-2020_12_31_424971 78 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 78 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 78 24 9 9 CD 10_1101-2020_12_31_424971 78 25 was be VBD 10_1101-2020_12_31_424971 78 26 generally generally RB 10_1101-2020_12_31_424971 78 27 higher high JJR 10_1101-2020_12_31_424971 78 28 than than IN 10_1101-2020_12_31_424971 78 29 that that DT 10_1101-2020_12_31_424971 78 30 by by IN 10_1101-2020_12_31_424971 78 31 native native JJ 10_1101-2020_12_31_424971 78 32 - - HYPH 10_1101-2020_12_31_424971 78 33 state state NN 10_1101-2020_12_31_424971 78 34 SufB2 sufb2 NN 10_1101-2020_12_31_424971 78 35 . . . 10_1101-2020_12_31_424971 79 1 Notably notably RB 10_1101-2020_12_31_424971 79 2 , , , 10_1101-2020_12_31_424971 79 3 the the DT 10_1101-2020_12_31_424971 79 4 PURExpress purexpress NN 10_1101-2020_12_31_424971 79 5 system system NN 10_1101-2020_12_31_424971 79 6 contains contain VBZ 10_1101-2020_12_31_424971 79 7 203 203 CD 10_1101-2020_12_31_424971 79 8 all all DT 10_1101-2020_12_31_424971 79 9 canonical canonical JJ 10_1101-2020_12_31_424971 79 10 tRNAs trnas NN 10_1101-2020_12_31_424971 79 11 , , , 10_1101-2020_12_31_424971 79 12 including include VBG 10_1101-2020_12_31_424971 79 13 ProL ProL NNP 10_1101-2020_12_31_424971 79 14 and and CC 10_1101-2020_12_31_424971 79 15 ProM ProM NNP 10_1101-2020_12_31_424971 79 16 , , , 10_1101-2020_12_31_424971 79 17 indicating indicate VBG 10_1101-2020_12_31_424971 79 18 the the DT 10_1101-2020_12_31_424971 79 19 ability ability NN 10_1101-2020_12_31_424971 79 20 of of IN 10_1101-2020_12_31_424971 79 21 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 79 22 to to IN 10_1101-2020_12_31_424971 79 23 successfully successfully RB 10_1101-2020_12_31_424971 79 24 204 204 CD 10_1101-2020_12_31_424971 79 25 compete compete VB 10_1101-2020_12_31_424971 79 26 with with IN 10_1101-2020_12_31_424971 79 27 these these DT 10_1101-2020_12_31_424971 79 28 tRNAs trnas CD 10_1101-2020_12_31_424971 79 29 . . . 10_1101-2020_12_31_424971 80 1 205 205 CD 10_1101-2020_12_31_424971 80 2 206 206 CD 10_1101-2020_12_31_424971 80 3 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 80 4 uses use VBZ 10_1101-2020_12_31_424971 80 5 triplet triplet NN 10_1101-2020_12_31_424971 80 6 pairing pair VBG 10_1101-2020_12_31_424971 80 7 in in IN 10_1101-2020_12_31_424971 80 8 the the DT 10_1101-2020_12_31_424971 80 9 0-frame 0-frame CD 10_1101-2020_12_31_424971 80 10 at at IN 10_1101-2020_12_31_424971 80 11 the the DT 10_1101-2020_12_31_424971 80 12 A A NNP 10_1101-2020_12_31_424971 80 13 site site NN 10_1101-2020_12_31_424971 80 14 207 207 CD 10_1101-2020_12_31_424971 80 15 To to TO 10_1101-2020_12_31_424971 80 16 determine determine VB 10_1101-2020_12_31_424971 80 17 at at IN 10_1101-2020_12_31_424971 80 18 which which WDT 10_1101-2020_12_31_424971 80 19 step step NN 10_1101-2020_12_31_424971 80 20 in in IN 10_1101-2020_12_31_424971 80 21 the the DT 10_1101-2020_12_31_424971 80 22 elongation elongation NN 10_1101-2020_12_31_424971 80 23 cycle cycle NN 10_1101-2020_12_31_424971 80 24 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 80 25 undergoes undergoe NNS 10_1101-2020_12_31_424971 80 26 +1 +1 VBZ 10_1101-2020_12_31_424971 80 27 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 80 28 in in IN 10_1101-2020_12_31_424971 80 29 208 208 CD 10_1101-2020_12_31_424971 80 30 response response NN 10_1101-2020_12_31_424971 80 31 to to IN 10_1101-2020_12_31_424971 80 32 a a DT 10_1101-2020_12_31_424971 80 33 CCC CCC NNP 10_1101-2020_12_31_424971 80 34 - - HYPH 10_1101-2020_12_31_424971 80 35 C C NNP 10_1101-2020_12_31_424971 80 36 motif motif NN 10_1101-2020_12_31_424971 80 37 , , , 10_1101-2020_12_31_424971 80 38 we -PRON- PRP 10_1101-2020_12_31_424971 80 39 used use VBD 10_1101-2020_12_31_424971 80 40 an an DT 10_1101-2020_12_31_424971 80 41 E. E. NNP 10_1101-2020_12_31_424971 80 42 coli coli NNS 10_1101-2020_12_31_424971 80 43 in in IN 10_1101-2020_12_31_424971 80 44 vitro vitro FW 10_1101-2020_12_31_424971 80 45 translation translation NN 10_1101-2020_12_31_424971 80 46 system system NN 10_1101-2020_12_31_424971 80 47 composed compose VBN 10_1101-2020_12_31_424971 80 48 of of IN 10_1101-2020_12_31_424971 80 49 purified purify VBN 10_1101-2020_12_31_424971 80 50 209 209 CD 10_1101-2020_12_31_424971 80 51 components component NNS 10_1101-2020_12_31_424971 80 52 and and CC 10_1101-2020_12_31_424971 80 53 supplemented supplement VBN 10_1101-2020_12_31_424971 80 54 with with IN 10_1101-2020_12_31_424971 80 55 requisite requisite JJ 10_1101-2020_12_31_424971 80 56 tRNAs trnas NN 10_1101-2020_12_31_424971 80 57 and and CC 10_1101-2020_12_31_424971 80 58 translation translation NN 10_1101-2020_12_31_424971 80 59 factors factor NNS 10_1101-2020_12_31_424971 80 60 to to TO 10_1101-2020_12_31_424971 80 61 perform perform VB 10_1101-2020_12_31_424971 80 62 a a DT 10_1101-2020_12_31_424971 80 63 series series NN 10_1101-2020_12_31_424971 80 64 210 210 CD 10_1101-2020_12_31_424971 80 65 of of IN 10_1101-2020_12_31_424971 80 66 ensemble ensemble JJ 10_1101-2020_12_31_424971 80 67 rapid rapid JJ 10_1101-2020_12_31_424971 80 68 kinetic kinetic NNP 10_1101-2020_12_31_424971 80 69 studies study NNS 10_1101-2020_12_31_424971 80 70 . . . 10_1101-2020_12_31_424971 81 1 We -PRON- PRP 10_1101-2020_12_31_424971 81 2 began begin VBD 10_1101-2020_12_31_424971 81 3 with with IN 10_1101-2020_12_31_424971 81 4 a a DT 10_1101-2020_12_31_424971 81 5 GTPase GTPase NNP 10_1101-2020_12_31_424971 81 6 assay assay NN 10_1101-2020_12_31_424971 81 7 that that IN 10_1101-2020_12_31_424971 81 8 reports report VBZ 10_1101-2020_12_31_424971 81 9 on on IN 10_1101-2020_12_31_424971 81 10 the the DT 10_1101-2020_12_31_424971 81 11 yield yield NN 10_1101-2020_12_31_424971 81 12 and and CC 10_1101-2020_12_31_424971 81 13 211 211 CD 10_1101-2020_12_31_424971 81 14 rate rate NN 10_1101-2020_12_31_424971 81 15 with with IN 10_1101-2020_12_31_424971 81 16 which which WDT 10_1101-2020_12_31_424971 81 17 the the DT 10_1101-2020_12_31_424971 81 18 translational translational JJ 10_1101-2020_12_31_424971 81 19 GTPase GTPase NNP 10_1101-2020_12_31_424971 81 20 EF EF NNP 10_1101-2020_12_31_424971 81 21 - - HYPH 10_1101-2020_12_31_424971 81 22 Tu Tu NNP 10_1101-2020_12_31_424971 81 23 hydrolyzes hydrolyze VBZ 10_1101-2020_12_31_424971 81 24 GTP GTP NNP 10_1101-2020_12_31_424971 81 25 upon upon IN 10_1101-2020_12_31_424971 81 26 delivery delivery NN 10_1101-2020_12_31_424971 81 27 of of IN 10_1101-2020_12_31_424971 81 28 a a DT 10_1101-2020_12_31_424971 81 29 ternary ternary JJ 10_1101-2020_12_31_424971 81 30 212 212 CD 10_1101-2020_12_31_424971 81 31 complex complex NN 10_1101-2020_12_31_424971 81 32 ( ( -LRB- 10_1101-2020_12_31_424971 81 33 TC TC NNP 10_1101-2020_12_31_424971 81 34 ) ) -RRB- 10_1101-2020_12_31_424971 81 35 , , , 10_1101-2020_12_31_424971 81 36 composed compose VBN 10_1101-2020_12_31_424971 81 37 of of IN 10_1101-2020_12_31_424971 81 38 EF EF NNP 10_1101-2020_12_31_424971 81 39 - - HYPH 10_1101-2020_12_31_424971 81 40 Tu Tu NNP 10_1101-2020_12_31_424971 81 41 , , , 10_1101-2020_12_31_424971 81 42 [ [ -LRB- 10_1101-2020_12_31_424971 81 43 g-32P]-GTP g-32p]-gtp NN 10_1101-2020_12_31_424971 81 44 , , , 10_1101-2020_12_31_424971 81 45 and and CC 10_1101-2020_12_31_424971 81 46 prolyl prolyl NNP 10_1101-2020_12_31_424971 81 47 - - HYPH 10_1101-2020_12_31_424971 81 48 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 81 49 ( ( -LRB- 10_1101-2020_12_31_424971 81 50 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 81 51 ) ) -RRB- 10_1101-2020_12_31_424971 81 52 or or CC 10_1101-2020_12_31_424971 81 53 ProL ProL NNP 10_1101-2020_12_31_424971 81 54 ( ( -LRB- 10_1101-2020_12_31_424971 81 55 ProL ProL NNP 10_1101-2020_12_31_424971 81 56 - - HYPH 10_1101-2020_12_31_424971 81 57 TC TC NNP 10_1101-2020_12_31_424971 81 58 ) ) -RRB- 10_1101-2020_12_31_424971 81 59 , , , 10_1101-2020_12_31_424971 81 60 213 213 CD 10_1101-2020_12_31_424971 81 61 to to IN 10_1101-2020_12_31_424971 81 62 the the DT 10_1101-2020_12_31_424971 81 63 A A NNP 10_1101-2020_12_31_424971 81 64 site site NN 10_1101-2020_12_31_424971 81 65 of of IN 10_1101-2020_12_31_424971 81 66 a a DT 10_1101-2020_12_31_424971 81 67 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 81 68 70S 70S NNP 10_1101-2020_12_31_424971 81 69 initiation initiation NN 10_1101-2020_12_31_424971 81 70 complex complex NN 10_1101-2020_12_31_424971 81 71 ( ( -LRB- 10_1101-2020_12_31_424971 81 72 70S 70S NNP 10_1101-2020_12_31_424971 81 73 IC IC NNP 10_1101-2020_12_31_424971 81 74 ) ) -RRB- 10_1101-2020_12_31_424971 81 75 carrying carry VBG 10_1101-2020_12_31_424971 81 76 an an DT 10_1101-2020_12_31_424971 81 77 initiator initiator NN 10_1101-2020_12_31_424971 81 78 fMet fMet NNP 10_1101-2020_12_31_424971 81 79 - - HYPH 10_1101-2020_12_31_424971 81 80 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 81 81 in in IN 10_1101-2020_12_31_424971 81 82 214 214 CD 10_1101-2020_12_31_424971 81 83 the the DT 10_1101-2020_12_31_424971 81 84 P p NN 10_1101-2020_12_31_424971 81 85 site site NN 10_1101-2020_12_31_424971 81 86 and and CC 10_1101-2020_12_31_424971 81 87 a a DT 10_1101-2020_12_31_424971 81 88 programmed program VBN 10_1101-2020_12_31_424971 81 89 CCC CCC NNP 10_1101-2020_12_31_424971 81 90 - - HYPH 10_1101-2020_12_31_424971 81 91 C C NNP 10_1101-2020_12_31_424971 81 92 motif motif NN 10_1101-2020_12_31_424971 81 93 at at IN 10_1101-2020_12_31_424971 81 94 the the DT 10_1101-2020_12_31_424971 81 95 A A NNP 10_1101-2020_12_31_424971 81 96 site site NN 10_1101-2020_12_31_424971 81 97 . . . 10_1101-2020_12_31_424971 82 1 The the DT 10_1101-2020_12_31_424971 82 2 results result NNS 10_1101-2020_12_31_424971 82 3 of of IN 10_1101-2020_12_31_424971 82 4 these these DT 10_1101-2020_12_31_424971 82 5 experiments experiment NNS 10_1101-2020_12_31_424971 82 6 showed show VBD 10_1101-2020_12_31_424971 82 7 215 215 CD 10_1101-2020_12_31_424971 82 8 that that IN 10_1101-2020_12_31_424971 82 9 the the DT 10_1101-2020_12_31_424971 82 10 yield yield NN 10_1101-2020_12_31_424971 82 11 and and CC 10_1101-2020_12_31_424971 82 12 rate rate NN 10_1101-2020_12_31_424971 82 13 of of IN 10_1101-2020_12_31_424971 82 14 GTP GTP NNP 10_1101-2020_12_31_424971 82 15 hydrolysis hydrolysis NN 10_1101-2020_12_31_424971 82 16 ( ( -LRB- 10_1101-2020_12_31_424971 82 17 kGTP kGTP NNP 10_1101-2020_12_31_424971 82 18 , , , 10_1101-2020_12_31_424971 82 19 obs obs NNP 10_1101-2020_12_31_424971 82 20 ) ) -RRB- 10_1101-2020_12_31_424971 82 21 upon upon IN 10_1101-2020_12_31_424971 82 22 delivery delivery NN 10_1101-2020_12_31_424971 82 23 of of IN 10_1101-2020_12_31_424971 82 24 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 82 25 were be VBD 10_1101-2020_12_31_424971 82 26 quantitatively quantitatively RB 10_1101-2020_12_31_424971 82 27 216 216 CD 10_1101-2020_12_31_424971 82 28 similar similar JJ 10_1101-2020_12_31_424971 82 29 to to IN 10_1101-2020_12_31_424971 82 30 those those DT 10_1101-2020_12_31_424971 82 31 of of IN 10_1101-2020_12_31_424971 82 32 ProL ProL NNP 10_1101-2020_12_31_424971 82 33 - - HYPH 10_1101-2020_12_31_424971 82 34 TC TC NNP 10_1101-2020_12_31_424971 82 35 for for IN 10_1101-2020_12_31_424971 82 36 both both CC 10_1101-2020_12_31_424971 82 37 the the DT 10_1101-2020_12_31_424971 82 38 native- native- NN 10_1101-2020_12_31_424971 82 39 and and CC 10_1101-2020_12_31_424971 82 40 G37-state G37-state NNP 10_1101-2020_12_31_424971 82 41 tRNAs trnas CD 10_1101-2020_12_31_424971 82 42 ( ( -LRB- 10_1101-2020_12_31_424971 82 43 Figure Figure NNP 10_1101-2020_12_31_424971 82 44 3a 3a NNP 10_1101-2020_12_31_424971 82 45 ) ) -RRB- 10_1101-2020_12_31_424971 82 46 . . . 10_1101-2020_12_31_424971 83 1 217 217 CD 10_1101-2020_12_31_424971 83 2 We -PRON- PRP 10_1101-2020_12_31_424971 83 3 next next RB 10_1101-2020_12_31_424971 83 4 performed perform VBD 10_1101-2020_12_31_424971 83 5 a a DT 10_1101-2020_12_31_424971 83 6 dipeptide dipeptide NN 10_1101-2020_12_31_424971 83 7 formation formation NN 10_1101-2020_12_31_424971 83 8 assay assay NN 10_1101-2020_12_31_424971 83 9 that that WDT 10_1101-2020_12_31_424971 83 10 reports report VBZ 10_1101-2020_12_31_424971 83 11 on on IN 10_1101-2020_12_31_424971 83 12 the the DT 10_1101-2020_12_31_424971 83 13 synthesis synthesis NN 10_1101-2020_12_31_424971 83 14 of of IN 10_1101-2020_12_31_424971 83 15 a a DT 10_1101-2020_12_31_424971 83 16 peptide peptide JJ 10_1101-2020_12_31_424971 83 17 218 218 CD 10_1101-2020_12_31_424971 83 18 bond bond NN 10_1101-2020_12_31_424971 83 19 between between IN 10_1101-2020_12_31_424971 83 20 the the DT 10_1101-2020_12_31_424971 83 21 [ [ -LRB- 10_1101-2020_12_31_424971 83 22 35S]-fMet 35s]-fmet CD 10_1101-2020_12_31_424971 83 23 moiety moiety NN 10_1101-2020_12_31_424971 83 24 of of IN 10_1101-2020_12_31_424971 83 25 a a DT 10_1101-2020_12_31_424971 83 26 P p NN 10_1101-2020_12_31_424971 83 27 - - HYPH 10_1101-2020_12_31_424971 83 28 site site NN 10_1101-2020_12_31_424971 83 29 [ [ -LRB- 10_1101-2020_12_31_424971 83 30 35S]-fMet 35s]-fmet CD 10_1101-2020_12_31_424971 83 31 - - HYPH 10_1101-2020_12_31_424971 83 32 tRNAfMet trnafmet NN 10_1101-2020_12_31_424971 83 33 in in IN 10_1101-2020_12_31_424971 83 34 a a DT 10_1101-2020_12_31_424971 83 35 70S 70S NNP 10_1101-2020_12_31_424971 83 36 IC IC NNP 10_1101-2020_12_31_424971 83 37 and and CC 10_1101-2020_12_31_424971 83 38 the the DT 10_1101-2020_12_31_424971 83 39 Pro pro JJ 10_1101-2020_12_31_424971 83 40 moiety moiety NN 10_1101-2020_12_31_424971 83 41 219 219 CD 10_1101-2020_12_31_424971 83 42 of of IN 10_1101-2020_12_31_424971 83 43 a a DT 10_1101-2020_12_31_424971 83 44 SufB2- sufb2- NN 10_1101-2020_12_31_424971 83 45 or or CC 10_1101-2020_12_31_424971 83 46 ProL ProL NNP 10_1101-2020_12_31_424971 83 47 - - HYPH 10_1101-2020_12_31_424971 83 48 TC TC NNP 10_1101-2020_12_31_424971 83 49 delivered deliver VBD 10_1101-2020_12_31_424971 83 50 to to IN 10_1101-2020_12_31_424971 83 51 the the DT 10_1101-2020_12_31_424971 83 52 A A NNP 10_1101-2020_12_31_424971 83 53 site site NN 10_1101-2020_12_31_424971 83 54 . . . 10_1101-2020_12_31_424971 84 1 This this DT 10_1101-2020_12_31_424971 84 2 assay assay NN 10_1101-2020_12_31_424971 84 3 revealed reveal VBD 10_1101-2020_12_31_424971 84 4 that that IN 10_1101-2020_12_31_424971 84 5 the the DT 10_1101-2020_12_31_424971 84 6 rate rate NN 10_1101-2020_12_31_424971 84 7 of of IN 10_1101-2020_12_31_424971 84 8 [ [ -LRB- 10_1101-2020_12_31_424971 84 9 35S]-fMet 35s]-fmet CD 10_1101-2020_12_31_424971 84 10 - - HYPH 10_1101-2020_12_31_424971 84 11 Pro pro NN 10_1101-2020_12_31_424971 84 12 220 220 CD 10_1101-2020_12_31_424971 84 13 ( ( -LRB- 10_1101-2020_12_31_424971 84 14 fMP fmp NN 10_1101-2020_12_31_424971 84 15 ) ) -RRB- 10_1101-2020_12_31_424971 84 16 formation formation NN 10_1101-2020_12_31_424971 84 17 ( ( -LRB- 10_1101-2020_12_31_424971 84 18 kfMP kfMP NNP 10_1101-2020_12_31_424971 84 19 , , , 10_1101-2020_12_31_424971 84 20 obs obs NN 10_1101-2020_12_31_424971 84 21 ) ) -RRB- 10_1101-2020_12_31_424971 84 22 for for IN 10_1101-2020_12_31_424971 84 23 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 84 24 was be VBD 10_1101-2020_12_31_424971 84 25 within within IN 10_1101-2020_12_31_424971 84 26 2-fold 2-fold CD 10_1101-2020_12_31_424971 84 27 of of IN 10_1101-2020_12_31_424971 84 28 that that DT 10_1101-2020_12_31_424971 84 29 for for IN 10_1101-2020_12_31_424971 84 30 ProL ProL NNP 10_1101-2020_12_31_424971 84 31 - - HYPH 10_1101-2020_12_31_424971 84 32 TC TC NNP 10_1101-2020_12_31_424971 84 33 for for IN 10_1101-2020_12_31_424971 84 34 both both CC 10_1101-2020_12_31_424971 84 35 the the DT 10_1101-2020_12_31_424971 84 36 native- native- NN 10_1101-2020_12_31_424971 84 37 221 221 CD 10_1101-2020_12_31_424971 84 38 and and CC 10_1101-2020_12_31_424971 84 39 G37-state G37-state NNP 10_1101-2020_12_31_424971 84 40 tRNAs trnas CD 10_1101-2020_12_31_424971 84 41 ( ( -LRB- 10_1101-2020_12_31_424971 84 42 Figure figure NN 10_1101-2020_12_31_424971 84 43 3b 3b NNP 10_1101-2020_12_31_424971 84 44 , , , 10_1101-2020_12_31_424971 84 45 Table Table NNP 10_1101-2020_12_31_424971 84 46 S2 S2 NNP 10_1101-2020_12_31_424971 84 47 ) ) -RRB- 10_1101-2020_12_31_424971 84 48 . . . 10_1101-2020_12_31_424971 85 1 222 222 CD 10_1101-2020_12_31_424971 85 2 To to TO 10_1101-2020_12_31_424971 85 3 test test VB 10_1101-2020_12_31_424971 85 4 whether whether IN 10_1101-2020_12_31_424971 85 5 native native JJ 10_1101-2020_12_31_424971 85 6 - - HYPH 10_1101-2020_12_31_424971 85 7 state state NN 10_1101-2020_12_31_424971 85 8 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 85 9 can can MD 10_1101-2020_12_31_424971 85 10 effectively effectively RB 10_1101-2020_12_31_424971 85 11 compete compete VB 10_1101-2020_12_31_424971 85 12 with with IN 10_1101-2020_12_31_424971 85 13 ProL ProL NNP 10_1101-2020_12_31_424971 85 14 - - HYPH 10_1101-2020_12_31_424971 85 15 TC TC NNP 10_1101-2020_12_31_424971 85 16 for for IN 10_1101-2020_12_31_424971 85 17 delivery delivery NN 10_1101-2020_12_31_424971 85 18 to to IN 10_1101-2020_12_31_424971 85 19 223 223 CD 10_1101-2020_12_31_424971 85 20 the the DT 10_1101-2020_12_31_424971 85 21 A A NNP 10_1101-2020_12_31_424971 85 22 site site NN 10_1101-2020_12_31_424971 85 23 and and CC 10_1101-2020_12_31_424971 85 24 peptide peptide NN 10_1101-2020_12_31_424971 85 25 - - HYPH 10_1101-2020_12_31_424971 85 26 bond bond NN 10_1101-2020_12_31_424971 85 27 formation formation NN 10_1101-2020_12_31_424971 85 28 , , , 10_1101-2020_12_31_424971 85 29 we -PRON- PRP 10_1101-2020_12_31_424971 85 30 varied vary VBD 10_1101-2020_12_31_424971 85 31 the the DT 10_1101-2020_12_31_424971 85 32 dipeptide dipeptide NN 10_1101-2020_12_31_424971 85 33 formation formation NN 10_1101-2020_12_31_424971 85 34 assay assay NN 10_1101-2020_12_31_424971 85 35 such such JJ 10_1101-2020_12_31_424971 85 36 that that IN 10_1101-2020_12_31_424971 85 37 an an DT 10_1101-2020_12_31_424971 85 38 224 224 CD 10_1101-2020_12_31_424971 85 39 equimolar equimolar JJ 10_1101-2020_12_31_424971 85 40 mixture mixture NN 10_1101-2020_12_31_424971 85 41 of of IN 10_1101-2020_12_31_424971 85 42 each each DT 10_1101-2020_12_31_424971 85 43 TC TC NNP 10_1101-2020_12_31_424971 85 44 was be VBD 10_1101-2020_12_31_424971 85 45 used use VBN 10_1101-2020_12_31_424971 85 46 in in IN 10_1101-2020_12_31_424971 85 47 the the DT 10_1101-2020_12_31_424971 85 48 reaction reaction NN 10_1101-2020_12_31_424971 85 49 ( ( -LRB- 10_1101-2020_12_31_424971 85 50 Figure figure NN 10_1101-2020_12_31_424971 85 51 3c 3c NN 10_1101-2020_12_31_424971 85 52 ) ) -RRB- 10_1101-2020_12_31_424971 85 53 . . . 10_1101-2020_12_31_424971 86 1 Since since IN 10_1101-2020_12_31_424971 86 2 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 86 3 of of IN 10_1101-2020_12_31_424971 86 4 both both CC 10_1101-2020_12_31_424971 86 5 225 225 CD 10_1101-2020_12_31_424971 86 6 tRNAs trnas CD 10_1101-2020_12_31_424971 86 7 with with IN 10_1101-2020_12_31_424971 86 8 Pro Pro NNP 10_1101-2020_12_31_424971 86 9 would would MD 10_1101-2020_12_31_424971 86 10 create create VB 10_1101-2020_12_31_424971 86 11 dipeptides dipeptide NNS 10_1101-2020_12_31_424971 86 12 of of IN 10_1101-2020_12_31_424971 86 13 the the DT 10_1101-2020_12_31_424971 86 14 same same JJ 10_1101-2020_12_31_424971 86 15 identity identity NN 10_1101-2020_12_31_424971 86 16 ( ( -LRB- 10_1101-2020_12_31_424971 86 17 i.e. i.e. FW 10_1101-2020_12_31_424971 86 18 , , , 10_1101-2020_12_31_424971 86 19 fMP fmp CD 10_1101-2020_12_31_424971 86 20 ) ) -RRB- 10_1101-2020_12_31_424971 86 21 , , , 10_1101-2020_12_31_424971 86 22 we -PRON- PRP 10_1101-2020_12_31_424971 86 23 used use VBD 10_1101-2020_12_31_424971 86 24 a a DT 10_1101-2020_12_31_424971 86 25 Flexizyme Flexizyme NNS 10_1101-2020_12_31_424971 86 26 to to TO 10_1101-2020_12_31_424971 86 27 226 226 CD 10_1101-2020_12_31_424971 86 28 aminoacylate aminoacylate VB 10_1101-2020_12_31_424971 86 29 them -PRON- PRP 10_1101-2020_12_31_424971 86 30 with with IN 10_1101-2020_12_31_424971 86 31 different different JJ 10_1101-2020_12_31_424971 86 32 amino amino JJ 10_1101-2020_12_31_424971 86 33 acids acid NNS 10_1101-2020_12_31_424971 86 34 and and CC 10_1101-2020_12_31_424971 86 35 generate generate VB 10_1101-2020_12_31_424971 86 36 distinct distinct JJ 10_1101-2020_12_31_424971 86 37 dipeptides dipeptide NNS 10_1101-2020_12_31_424971 86 38 . . . 10_1101-2020_12_31_424971 87 1 Control control NN 10_1101-2020_12_31_424971 87 2 227 227 CD 10_1101-2020_12_31_424971 87 3 experiments experiment NNS 10_1101-2020_12_31_424971 87 4 showed show VBD 10_1101-2020_12_31_424971 87 5 that that IN 10_1101-2020_12_31_424971 87 6 ProL ProL NNP 10_1101-2020_12_31_424971 87 7 charged charge VBD 10_1101-2020_12_31_424971 87 8 with with IN 10_1101-2020_12_31_424971 87 9 Pro Pro NNP 10_1101-2020_12_31_424971 87 10 or or CC 10_1101-2020_12_31_424971 87 11 Arg Arg NNP 10_1101-2020_12_31_424971 87 12 ( ( -LRB- 10_1101-2020_12_31_424971 87 13 Figure Figure NNP 10_1101-2020_12_31_424971 87 14 3c 3c NN 10_1101-2020_12_31_424971 87 15 , , , 10_1101-2020_12_31_424971 87 16 Bars Bars NNP 10_1101-2020_12_31_424971 87 17 1 1 CD 10_1101-2020_12_31_424971 87 18 and and CC 10_1101-2020_12_31_424971 87 19 2 2 CD 10_1101-2020_12_31_424971 87 20 ) ) -RRB- 10_1101-2020_12_31_424971 87 21 and and CC 10_1101-2020_12_31_424971 87 22 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 87 23 228 228 CD 10_1101-2020_12_31_424971 87 24 .CC .CC : 10_1101-2020_12_31_424971 87 25 - - HYPH 10_1101-2020_12_31_424971 87 26 BY by IN 10_1101-2020_12_31_424971 87 27 - - HYPH 10_1101-2020_12_31_424971 87 28 NC NC NNP 10_1101-2020_12_31_424971 87 29 - - HYPH 10_1101-2020_12_31_424971 87 30 ND ND NNP 10_1101-2020_12_31_424971 87 31 4.0 4.0 CD 10_1101-2020_12_31_424971 87 32 International International NNP 10_1101-2020_12_31_424971 87 33 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 87 34 under under IN 10_1101-2020_12_31_424971 87 35 a a DT 10_1101-2020_12_31_424971 87 36 ( ( -LRB- 10_1101-2020_12_31_424971 87 37 which which WDT 10_1101-2020_12_31_424971 87 38 was be VBD 10_1101-2020_12_31_424971 87 39 not not RB 10_1101-2020_12_31_424971 87 40 certified certify VBN 10_1101-2020_12_31_424971 87 41 by by IN 10_1101-2020_12_31_424971 87 42 peer peer NN 10_1101-2020_12_31_424971 87 43 review review NN 10_1101-2020_12_31_424971 87 44 ) ) -RRB- 10_1101-2020_12_31_424971 87 45 is be VBZ 10_1101-2020_12_31_424971 87 46 the the DT 10_1101-2020_12_31_424971 87 47 author author NN 10_1101-2020_12_31_424971 87 48 / / SYM 10_1101-2020_12_31_424971 87 49 funder funder NN 10_1101-2020_12_31_424971 87 50 , , , 10_1101-2020_12_31_424971 87 51 who who WP 10_1101-2020_12_31_424971 87 52 has have VBZ 10_1101-2020_12_31_424971 87 53 granted grant VBN 10_1101-2020_12_31_424971 87 54 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 87 55 a a DT 10_1101-2020_12_31_424971 87 56 license license NN 10_1101-2020_12_31_424971 87 57 to to TO 10_1101-2020_12_31_424971 87 58 display display VB 10_1101-2020_12_31_424971 87 59 the the DT 10_1101-2020_12_31_424971 87 60 preprint preprint NN 10_1101-2020_12_31_424971 87 61 in in IN 10_1101-2020_12_31_424971 87 62 perpetuity perpetuity NN 10_1101-2020_12_31_424971 87 63 . . . 10_1101-2020_12_31_424971 88 1 It -PRON- PRP 10_1101-2020_12_31_424971 88 2 is be VBZ 10_1101-2020_12_31_424971 88 3 made make VBN 10_1101-2020_12_31_424971 88 4 The the DT 10_1101-2020_12_31_424971 88 5 copyright copyright NN 10_1101-2020_12_31_424971 88 6 holder holder NN 10_1101-2020_12_31_424971 88 7 for for IN 10_1101-2020_12_31_424971 88 8 this this DT 10_1101-2020_12_31_424971 88 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 88 10 version version NN 10_1101-2020_12_31_424971 88 11 posted post VBD 10_1101-2020_12_31_424971 88 12 January January NNP 10_1101-2020_12_31_424971 88 13 2 2 CD 10_1101-2020_12_31_424971 88 14 , , , 10_1101-2020_12_31_424971 88 15 2021 2021 CD 10_1101-2020_12_31_424971 88 16 . . . 10_1101-2020_12_31_424971 88 17 ; ; : 10_1101-2020_12_31_424971 88 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 88 19 : : : 10_1101-2020_12_31_424971 88 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 88 21 preprint preprint NN 10_1101-2020_12_31_424971 88 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NNP 10_1101-2020_12_31_424971 88 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 88 24 10 10 CD 10_1101-2020_12_31_424971 88 25 charged charge VBN 10_1101-2020_12_31_424971 88 26 with with IN 10_1101-2020_12_31_424971 88 27 Pro Pro NNP 10_1101-2020_12_31_424971 88 28 or or CC 10_1101-2020_12_31_424971 88 29 Arg Arg NNP 10_1101-2020_12_31_424971 88 30 ( ( -LRB- 10_1101-2020_12_31_424971 88 31 Bars Bars NNP 10_1101-2020_12_31_424971 88 32 3 3 CD 10_1101-2020_12_31_424971 88 33 and and CC 10_1101-2020_12_31_424971 88 34 4 4 CD 10_1101-2020_12_31_424971 88 35 ) ) -RRB- 10_1101-2020_12_31_424971 88 36 generated generate VBD 10_1101-2020_12_31_424971 88 37 the the DT 10_1101-2020_12_31_424971 88 38 same same JJ 10_1101-2020_12_31_424971 88 39 amount amount NN 10_1101-2020_12_31_424971 88 40 of of IN 10_1101-2020_12_31_424971 88 41 fMP fMP NNS 10_1101-2020_12_31_424971 88 42 and and CC 10_1101-2020_12_31_424971 88 43 fMR fmr NN 10_1101-2020_12_31_424971 88 44 , , , 10_1101-2020_12_31_424971 88 45 indicating indicate VBG 10_1101-2020_12_31_424971 88 46 229 229 CD 10_1101-2020_12_31_424971 88 47 that that IN 10_1101-2020_12_31_424971 88 48 the the DT 10_1101-2020_12_31_424971 88 49 amino amino JJ 10_1101-2020_12_31_424971 88 50 - - HYPH 10_1101-2020_12_31_424971 88 51 acid acid NN 10_1101-2020_12_31_424971 88 52 identity identity NN 10_1101-2020_12_31_424971 88 53 did do VBD 10_1101-2020_12_31_424971 88 54 not not RB 10_1101-2020_12_31_424971 88 55 affect affect VB 10_1101-2020_12_31_424971 88 56 the the DT 10_1101-2020_12_31_424971 88 57 level level NN 10_1101-2020_12_31_424971 88 58 of of IN 10_1101-2020_12_31_424971 88 59 dipeptide dipeptide NN 10_1101-2020_12_31_424971 88 60 formation formation NN 10_1101-2020_12_31_424971 88 61 . . . 10_1101-2020_12_31_424971 89 1 We -PRON- PRP 10_1101-2020_12_31_424971 89 2 found find VBD 10_1101-2020_12_31_424971 89 3 that that IN 10_1101-2020_12_31_424971 89 4 the the DT 10_1101-2020_12_31_424971 89 5 230 230 CD 10_1101-2020_12_31_424971 89 6 amount amount NN 10_1101-2020_12_31_424971 89 7 of of IN 10_1101-2020_12_31_424971 89 8 dipeptide dipeptide NN 10_1101-2020_12_31_424971 89 9 formed form VBN 10_1101-2020_12_31_424971 89 10 by by IN 10_1101-2020_12_31_424971 89 11 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 89 12 and and CC 10_1101-2020_12_31_424971 89 13 ProL ProL NNP 10_1101-2020_12_31_424971 89 14 - - HYPH 10_1101-2020_12_31_424971 89 15 TC TC NNP 10_1101-2020_12_31_424971 89 16 in in IN 10_1101-2020_12_31_424971 89 17 these these DT 10_1101-2020_12_31_424971 89 18 competition competition NN 10_1101-2020_12_31_424971 89 19 assays assays RB 10_1101-2020_12_31_424971 89 20 was be VBD 10_1101-2020_12_31_424971 89 21 similar similar JJ 10_1101-2020_12_31_424971 89 22 , , , 10_1101-2020_12_31_424971 89 23 231 231 CD 10_1101-2020_12_31_424971 89 24 although although IN 10_1101-2020_12_31_424971 89 25 the the DT 10_1101-2020_12_31_424971 89 26 amount amount NN 10_1101-2020_12_31_424971 89 27 formed form VBN 10_1101-2020_12_31_424971 89 28 by by IN 10_1101-2020_12_31_424971 89 29 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 89 30 was be VBD 10_1101-2020_12_31_424971 89 31 slightly slightly RB 10_1101-2020_12_31_424971 89 32 less less JJR 10_1101-2020_12_31_424971 89 33 ( ( -LRB- 10_1101-2020_12_31_424971 89 34 45 45 CD 10_1101-2020_12_31_424971 89 35 % % NN 10_1101-2020_12_31_424971 89 36 vs. vs. NN 10_1101-2020_12_31_424971 89 37 55 55 CD 10_1101-2020_12_31_424971 89 38 % % NN 10_1101-2020_12_31_424971 89 39 ) ) -RRB- 10_1101-2020_12_31_424971 89 40 , , , 10_1101-2020_12_31_424971 89 41 in in IN 10_1101-2020_12_31_424971 89 42 both both CC 10_1101-2020_12_31_424971 89 43 the the DT 10_1101-2020_12_31_424971 89 44 native- native- JJ 10_1101-2020_12_31_424971 89 45 232 232 CD 10_1101-2020_12_31_424971 89 46 ( ( -LRB- 10_1101-2020_12_31_424971 89 47 Bars Bars NNP 10_1101-2020_12_31_424971 89 48 5 5 CD 10_1101-2020_12_31_424971 89 49 - - SYM 10_1101-2020_12_31_424971 89 50 8 8 CD 10_1101-2020_12_31_424971 89 51 ) ) -RRB- 10_1101-2020_12_31_424971 89 52 and and CC 10_1101-2020_12_31_424971 89 53 G37-state G37-state NNP 10_1101-2020_12_31_424971 89 54 tRNAs trnas CD 10_1101-2020_12_31_424971 89 55 ( ( -LRB- 10_1101-2020_12_31_424971 89 56 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 89 57 Figure Figure NNP 10_1101-2020_12_31_424971 89 58 2a 2a NNP 10_1101-2020_12_31_424971 89 59 ) ) -RRB- 10_1101-2020_12_31_424971 89 60 . . . 10_1101-2020_12_31_424971 90 1 These these DT 10_1101-2020_12_31_424971 90 2 competition competition NN 10_1101-2020_12_31_424971 90 3 experiments experiment NNS 10_1101-2020_12_31_424971 90 4 233 233 CD 10_1101-2020_12_31_424971 90 5 provide provide VB 10_1101-2020_12_31_424971 90 6 direct direct JJ 10_1101-2020_12_31_424971 90 7 evidence evidence NN 10_1101-2020_12_31_424971 90 8 that that IN 10_1101-2020_12_31_424971 90 9 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 90 10 effectively effectively RB 10_1101-2020_12_31_424971 90 11 competes compete VBZ 10_1101-2020_12_31_424971 90 12 with with IN 10_1101-2020_12_31_424971 90 13 ProL ProL NNP 10_1101-2020_12_31_424971 90 14 - - HYPH 10_1101-2020_12_31_424971 90 15 TC TC NNP 10_1101-2020_12_31_424971 90 16 for for IN 10_1101-2020_12_31_424971 90 17 delivery delivery NN 10_1101-2020_12_31_424971 90 18 to to IN 10_1101-2020_12_31_424971 90 19 the the DT 10_1101-2020_12_31_424971 90 20 A a NN 10_1101-2020_12_31_424971 90 21 site site NN 10_1101-2020_12_31_424971 90 22 234 234 CD 10_1101-2020_12_31_424971 90 23 and and CC 10_1101-2020_12_31_424971 90 24 peptide peptide NN 10_1101-2020_12_31_424971 90 25 - - HYPH 10_1101-2020_12_31_424971 90 26 bond bond NN 10_1101-2020_12_31_424971 90 27 formation formation NN 10_1101-2020_12_31_424971 90 28 . . . 10_1101-2020_12_31_424971 91 1 235 235 CD 10_1101-2020_12_31_424971 91 2 Collectively collectively RB 10_1101-2020_12_31_424971 91 3 , , , 10_1101-2020_12_31_424971 91 4 the the DT 10_1101-2020_12_31_424971 91 5 results result NNS 10_1101-2020_12_31_424971 91 6 of of IN 10_1101-2020_12_31_424971 91 7 our -PRON- PRP$ 10_1101-2020_12_31_424971 91 8 GTPase- gtpase- JJ 10_1101-2020_12_31_424971 91 9 , , , 10_1101-2020_12_31_424971 91 10 dipeptide dipeptide NN 10_1101-2020_12_31_424971 91 11 formation- formation- NN 10_1101-2020_12_31_424971 91 12 , , , 10_1101-2020_12_31_424971 91 13 and and CC 10_1101-2020_12_31_424971 91 14 competition competition NN 10_1101-2020_12_31_424971 91 15 assays assays RB 10_1101-2020_12_31_424971 91 16 indicate indicate VBP 10_1101-2020_12_31_424971 91 17 236 236 CD 10_1101-2020_12_31_424971 91 18 that that WDT 10_1101-2020_12_31_424971 91 19 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 91 20 is be VBZ 10_1101-2020_12_31_424971 91 21 delivered deliver VBN 10_1101-2020_12_31_424971 91 22 to to IN 10_1101-2020_12_31_424971 91 23 the the DT 10_1101-2020_12_31_424971 91 24 A A NNP 10_1101-2020_12_31_424971 91 25 site site NN 10_1101-2020_12_31_424971 91 26 and and CC 10_1101-2020_12_31_424971 91 27 participates participate VBZ 10_1101-2020_12_31_424971 91 28 in in IN 10_1101-2020_12_31_424971 91 29 peptide peptide NN 10_1101-2020_12_31_424971 91 30 - - HYPH 10_1101-2020_12_31_424971 91 31 bond bond NN 10_1101-2020_12_31_424971 91 32 formation formation NN 10_1101-2020_12_31_424971 91 33 in in IN 10_1101-2020_12_31_424971 91 34 the the DT 10_1101-2020_12_31_424971 91 35 same same JJ 10_1101-2020_12_31_424971 91 36 237 237 CD 10_1101-2020_12_31_424971 91 37 way way NN 10_1101-2020_12_31_424971 91 38 as as IN 10_1101-2020_12_31_424971 91 39 ProL ProL NNP 10_1101-2020_12_31_424971 91 40 - - HYPH 10_1101-2020_12_31_424971 91 41 TC TC NNP 10_1101-2020_12_31_424971 91 42 , , , 10_1101-2020_12_31_424971 91 43 suggesting suggest VBG 10_1101-2020_12_31_424971 91 44 that that IN 10_1101-2020_12_31_424971 91 45 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 91 46 uses use VBZ 10_1101-2020_12_31_424971 91 47 triplet triplet NN 10_1101-2020_12_31_424971 91 48 pairing pair VBG 10_1101-2020_12_31_424971 91 49 in in IN 10_1101-2020_12_31_424971 91 50 the the DT 10_1101-2020_12_31_424971 91 51 0-frame 0-frame CD 10_1101-2020_12_31_424971 91 52 at at IN 10_1101-2020_12_31_424971 91 53 the the DT 10_1101-2020_12_31_424971 91 54 A A NNP 10_1101-2020_12_31_424971 91 55 site site NN 10_1101-2020_12_31_424971 91 56 that that IN 10_1101-2020_12_31_424971 91 57 238 238 CD 10_1101-2020_12_31_424971 91 58 successfully successfully RB 10_1101-2020_12_31_424971 91 59 competes compete VBZ 10_1101-2020_12_31_424971 91 60 with with IN 10_1101-2020_12_31_424971 91 61 triplet triplet NN 10_1101-2020_12_31_424971 91 62 pairing pair VBG 10_1101-2020_12_31_424971 91 63 by by IN 10_1101-2020_12_31_424971 91 64 ProL. ProL. NNP 10_1101-2020_12_31_424971 92 1 To to TO 10_1101-2020_12_31_424971 92 2 support support VB 10_1101-2020_12_31_424971 92 3 this this DT 10_1101-2020_12_31_424971 92 4 interpretation interpretation NN 10_1101-2020_12_31_424971 92 5 , , , 10_1101-2020_12_31_424971 92 6 we -PRON- PRP 10_1101-2020_12_31_424971 92 7 measured measure VBD 10_1101-2020_12_31_424971 92 8 239 239 CD 10_1101-2020_12_31_424971 92 9 kfMP kfMP NNP 10_1101-2020_12_31_424971 92 10 , , , 10_1101-2020_12_31_424971 92 11 obs obs NN 10_1101-2020_12_31_424971 92 12 in in IN 10_1101-2020_12_31_424971 92 13 our -PRON- PRP$ 10_1101-2020_12_31_424971 92 14 dipeptide dipeptide NN 10_1101-2020_12_31_424971 92 15 formation formation NN 10_1101-2020_12_31_424971 92 16 assay assay NN 10_1101-2020_12_31_424971 92 17 , , , 10_1101-2020_12_31_424971 92 18 using use VBG 10_1101-2020_12_31_424971 92 19 G37-state G37-state NNP 10_1101-2020_12_31_424971 92 20 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 92 21 and and CC 10_1101-2020_12_31_424971 92 22 a a DT 10_1101-2020_12_31_424971 92 23 series series NN 10_1101-2020_12_31_424971 92 24 of of IN 10_1101-2020_12_31_424971 92 25 mRNA mRNA NNP 10_1101-2020_12_31_424971 92 26 variants variant NNS 10_1101-2020_12_31_424971 92 27 240 240 CD 10_1101-2020_12_31_424971 92 28 in in IN 10_1101-2020_12_31_424971 92 29 which which WDT 10_1101-2020_12_31_424971 92 30 single single JJ 10_1101-2020_12_31_424971 92 31 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 92 32 in in IN 10_1101-2020_12_31_424971 92 33 the the DT 10_1101-2020_12_31_424971 92 34 CCC CCC NNP 10_1101-2020_12_31_424971 92 35 - - HYPH 10_1101-2020_12_31_424971 92 36 C C NNP 10_1101-2020_12_31_424971 92 37 motif motif NN 10_1101-2020_12_31_424971 92 38 were be VBD 10_1101-2020_12_31_424971 92 39 substituted substitute VBN 10_1101-2020_12_31_424971 92 40 . . . 10_1101-2020_12_31_424971 93 1 We -PRON- PRP 10_1101-2020_12_31_424971 93 2 showed show VBD 10_1101-2020_12_31_424971 93 3 that that IN 10_1101-2020_12_31_424971 93 4 kfMP kfMP NNP 10_1101-2020_12_31_424971 93 5 , , , 10_1101-2020_12_31_424971 93 6 obs obs NNP 10_1101-2020_12_31_424971 93 7 did do VBD 10_1101-2020_12_31_424971 93 8 not not RB 10_1101-2020_12_31_424971 93 9 241 241 CD 10_1101-2020_12_31_424971 93 10 decrease decrease NN 10_1101-2020_12_31_424971 93 11 upon upon IN 10_1101-2020_12_31_424971 93 12 substitution substitution NN 10_1101-2020_12_31_424971 93 13 of of IN 10_1101-2020_12_31_424971 93 14 the the DT 10_1101-2020_12_31_424971 93 15 4th 4th JJ 10_1101-2020_12_31_424971 93 16 nucleotide nucleotide NN 10_1101-2020_12_31_424971 93 17 of of IN 10_1101-2020_12_31_424971 93 18 the the DT 10_1101-2020_12_31_424971 93 19 CCC CCC NNP 10_1101-2020_12_31_424971 93 20 - - HYPH 10_1101-2020_12_31_424971 93 21 C C NNP 10_1101-2020_12_31_424971 93 22 motif motif NN 10_1101-2020_12_31_424971 93 23 , , , 10_1101-2020_12_31_424971 93 24 but but CC 10_1101-2020_12_31_424971 93 25 that that IN 10_1101-2020_12_31_424971 93 26 it -PRON- PRP 10_1101-2020_12_31_424971 93 27 decreased decrease VBD 10_1101-2020_12_31_424971 93 28 242 242 CD 10_1101-2020_12_31_424971 93 29 substantially substantially RB 10_1101-2020_12_31_424971 93 30 upon upon IN 10_1101-2020_12_31_424971 93 31 substitution substitution NN 10_1101-2020_12_31_424971 93 32 of of IN 10_1101-2020_12_31_424971 93 33 any any DT 10_1101-2020_12_31_424971 93 34 of of IN 10_1101-2020_12_31_424971 93 35 the the DT 10_1101-2020_12_31_424971 93 36 first first JJ 10_1101-2020_12_31_424971 93 37 three three CD 10_1101-2020_12_31_424971 93 38 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 93 39 of of IN 10_1101-2020_12_31_424971 93 40 the the DT 10_1101-2020_12_31_424971 93 41 motif motif NN 10_1101-2020_12_31_424971 93 42 ( ( -LRB- 10_1101-2020_12_31_424971 93 43 Figure Figure NNP 10_1101-2020_12_31_424971 93 44 3d 3d NN 10_1101-2020_12_31_424971 93 45 , , , 10_1101-2020_12_31_424971 93 46 243 243 CD 10_1101-2020_12_31_424971 93 47 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 93 48 Figure Figure NNP 10_1101-2020_12_31_424971 93 49 2b 2b NNP 10_1101-2020_12_31_424971 93 50 ) ) -RRB- 10_1101-2020_12_31_424971 93 51 . . . 10_1101-2020_12_31_424971 94 1 Thus thus RB 10_1101-2020_12_31_424971 94 2 , , , 10_1101-2020_12_31_424971 94 3 triplet triplet NN 10_1101-2020_12_31_424971 94 4 pairing pairing NN 10_1101-2020_12_31_424971 94 5 of of IN 10_1101-2020_12_31_424971 94 6 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 94 7 to to IN 10_1101-2020_12_31_424971 94 8 the the DT 10_1101-2020_12_31_424971 94 9 first first JJ 10_1101-2020_12_31_424971 94 10 three three CD 10_1101-2020_12_31_424971 94 11 Cs Cs NNP 10_1101-2020_12_31_424971 94 12 of of IN 10_1101-2020_12_31_424971 94 13 the the DT 10_1101-2020_12_31_424971 94 14 CCC CCC NNP 10_1101-2020_12_31_424971 94 15 - - HYPH 10_1101-2020_12_31_424971 94 16 C C NNP 10_1101-2020_12_31_424971 94 17 motif motif NN 10_1101-2020_12_31_424971 94 18 244 244 CD 10_1101-2020_12_31_424971 94 19 is be VBZ 10_1101-2020_12_31_424971 94 20 necessary necessary JJ 10_1101-2020_12_31_424971 94 21 and and CC 10_1101-2020_12_31_424971 94 22 sufficient sufficient JJ 10_1101-2020_12_31_424971 94 23 for for IN 10_1101-2020_12_31_424971 94 24 rapid rapid JJ 10_1101-2020_12_31_424971 94 25 delivery delivery NN 10_1101-2020_12_31_424971 94 26 of of IN 10_1101-2020_12_31_424971 94 27 the the DT 10_1101-2020_12_31_424971 94 28 tRNA trna NN 10_1101-2020_12_31_424971 94 29 to to IN 10_1101-2020_12_31_424971 94 30 the the DT 10_1101-2020_12_31_424971 94 31 A A NNP 10_1101-2020_12_31_424971 94 32 site site NN 10_1101-2020_12_31_424971 94 33 and and CC 10_1101-2020_12_31_424971 94 34 its -PRON- PRP$ 10_1101-2020_12_31_424971 94 35 participation participation NN 10_1101-2020_12_31_424971 94 36 in in IN 10_1101-2020_12_31_424971 94 37 245 245 CD 10_1101-2020_12_31_424971 94 38 peptide peptide NN 10_1101-2020_12_31_424971 94 39 - - HYPH 10_1101-2020_12_31_424971 94 40 bond bond NN 10_1101-2020_12_31_424971 94 41 formation formation NN 10_1101-2020_12_31_424971 94 42 . . . 10_1101-2020_12_31_424971 95 1 246 246 CD 10_1101-2020_12_31_424971 95 2 247 247 CD 10_1101-2020_12_31_424971 95 3 The the DT 10_1101-2020_12_31_424971 95 4 A A NNP 10_1101-2020_12_31_424971 95 5 - - HYPH 10_1101-2020_12_31_424971 95 6 site site NN 10_1101-2020_12_31_424971 95 7 activity activity NN 10_1101-2020_12_31_424971 95 8 of of IN 10_1101-2020_12_31_424971 95 9 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 95 10 depends depend VBZ 10_1101-2020_12_31_424971 95 11 on on IN 10_1101-2020_12_31_424971 95 12 the the DT 10_1101-2020_12_31_424971 95 13 sequence sequence NN 10_1101-2020_12_31_424971 95 14 of of IN 10_1101-2020_12_31_424971 95 15 the the DT 10_1101-2020_12_31_424971 95 16 anticodon anticodon NN 10_1101-2020_12_31_424971 95 17 loop loop NN 10_1101-2020_12_31_424971 95 18 248 248 CD 10_1101-2020_12_31_424971 95 19 We -PRON- PRP 10_1101-2020_12_31_424971 95 20 next next RB 10_1101-2020_12_31_424971 95 21 asked ask VBD 10_1101-2020_12_31_424971 95 22 how how WRB 10_1101-2020_12_31_424971 95 23 delivery delivery NN 10_1101-2020_12_31_424971 95 24 of of IN 10_1101-2020_12_31_424971 95 25 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 95 26 to to IN 10_1101-2020_12_31_424971 95 27 the the DT 10_1101-2020_12_31_424971 95 28 A a NN 10_1101-2020_12_31_424971 95 29 site site NN 10_1101-2020_12_31_424971 95 30 and and CC 10_1101-2020_12_31_424971 95 31 peptide peptide NN 10_1101-2020_12_31_424971 95 32 - - HYPH 10_1101-2020_12_31_424971 95 33 bond bond NN 10_1101-2020_12_31_424971 95 34 formation formation NN 10_1101-2020_12_31_424971 95 35 depend depend VBP 10_1101-2020_12_31_424971 95 36 on on IN 10_1101-2020_12_31_424971 95 37 249 249 CD 10_1101-2020_12_31_424971 95 38 the the DT 10_1101-2020_12_31_424971 95 39 sequence sequence NN 10_1101-2020_12_31_424971 95 40 of of IN 10_1101-2020_12_31_424971 95 41 the the DT 10_1101-2020_12_31_424971 95 42 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 95 43 anticodon anticodon NN 10_1101-2020_12_31_424971 95 44 loop loop NN 10_1101-2020_12_31_424971 95 45 . . . 10_1101-2020_12_31_424971 96 1 Starting start VBG 10_1101-2020_12_31_424971 96 2 from from IN 10_1101-2020_12_31_424971 96 3 G37-state G37-state NNP 10_1101-2020_12_31_424971 96 4 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 96 5 , , , 10_1101-2020_12_31_424971 96 6 we -PRON- PRP 10_1101-2020_12_31_424971 96 7 created create VBD 10_1101-2020_12_31_424971 96 8 two two CD 10_1101-2020_12_31_424971 96 9 250 250 CD 10_1101-2020_12_31_424971 96 10 variants variant NNS 10_1101-2020_12_31_424971 96 11 containing contain VBG 10_1101-2020_12_31_424971 96 12 a a DT 10_1101-2020_12_31_424971 96 13 G g NN 10_1101-2020_12_31_424971 96 14 - - HYPH 10_1101-2020_12_31_424971 96 15 to to IN 10_1101-2020_12_31_424971 96 16 - - HYPH 10_1101-2020_12_31_424971 96 17 C c NN 10_1101-2020_12_31_424971 96 18 substitution substitution NN 10_1101-2020_12_31_424971 96 19 in in IN 10_1101-2020_12_31_424971 96 20 nucleotide nucleotide JJ 10_1101-2020_12_31_424971 96 21 37 37 CD 10_1101-2020_12_31_424971 96 22 ( ( -LRB- 10_1101-2020_12_31_424971 96 23 G37C G37C NNP 10_1101-2020_12_31_424971 96 24 ) ) -RRB- 10_1101-2020_12_31_424971 96 25 or or CC 10_1101-2020_12_31_424971 96 26 34 34 CD 10_1101-2020_12_31_424971 96 27 ( ( -LRB- 10_1101-2020_12_31_424971 96 28 G34C g34c NN 10_1101-2020_12_31_424971 96 29 ) ) -RRB- 10_1101-2020_12_31_424971 96 30 within within IN 10_1101-2020_12_31_424971 96 31 the the DT 10_1101-2020_12_31_424971 96 32 251 251 CD 10_1101-2020_12_31_424971 96 33 anticodon anticodon NNP 10_1101-2020_12_31_424971 96 34 loop loop NN 10_1101-2020_12_31_424971 96 35 and and CC 10_1101-2020_12_31_424971 96 36 adapted adapt VBD 10_1101-2020_12_31_424971 96 37 our -PRON- PRP$ 10_1101-2020_12_31_424971 96 38 dipeptide dipeptide NN 10_1101-2020_12_31_424971 96 39 formation formation NN 10_1101-2020_12_31_424971 96 40 assay assay NN 10_1101-2020_12_31_424971 96 41 to to TO 10_1101-2020_12_31_424971 96 42 measure measure VB 10_1101-2020_12_31_424971 96 43 the the DT 10_1101-2020_12_31_424971 96 44 fMP fMP NNS 10_1101-2020_12_31_424971 96 45 yield yield NN 10_1101-2020_12_31_424971 96 46 and and CC 10_1101-2020_12_31_424971 96 47 kfMP kfMP NNP 10_1101-2020_12_31_424971 96 48 , , , 10_1101-2020_12_31_424971 96 49 obs obs NNP 10_1101-2020_12_31_424971 96 50 252 252 CD 10_1101-2020_12_31_424971 96 51 generated generate VBN 10_1101-2020_12_31_424971 96 52 by by IN 10_1101-2020_12_31_424971 96 53 each each DT 10_1101-2020_12_31_424971 96 54 variant variant NN 10_1101-2020_12_31_424971 96 55 at at IN 10_1101-2020_12_31_424971 96 56 the the DT 10_1101-2020_12_31_424971 96 57 CCC CCC NNP 10_1101-2020_12_31_424971 96 58 - - HYPH 10_1101-2020_12_31_424971 96 59 C C NNP 10_1101-2020_12_31_424971 96 60 motif motif NN 10_1101-2020_12_31_424971 96 61 at at IN 10_1101-2020_12_31_424971 96 62 the the DT 10_1101-2020_12_31_424971 96 63 A A NNP 10_1101-2020_12_31_424971 96 64 site site NN 10_1101-2020_12_31_424971 96 65 . . . 10_1101-2020_12_31_424971 97 1 We -PRON- PRP 10_1101-2020_12_31_424971 97 2 showed show VBD 10_1101-2020_12_31_424971 97 3 that that IN 10_1101-2020_12_31_424971 97 4 the the DT 10_1101-2020_12_31_424971 97 5 G37C G37C NNP 10_1101-2020_12_31_424971 97 6 variant variant NN 10_1101-2020_12_31_424971 97 7 253 253 CD 10_1101-2020_12_31_424971 97 8 resulted result VBD 10_1101-2020_12_31_424971 97 9 in in IN 10_1101-2020_12_31_424971 97 10 a a DT 10_1101-2020_12_31_424971 97 11 fMP fmp CD 10_1101-2020_12_31_424971 97 12 yield yield NN 10_1101-2020_12_31_424971 97 13 of of IN 10_1101-2020_12_31_424971 97 14 32 32 CD 10_1101-2020_12_31_424971 97 15 % % NN 10_1101-2020_12_31_424971 97 16 and and CC 10_1101-2020_12_31_424971 97 17 a a DT 10_1101-2020_12_31_424971 97 18 kfMP kfMP NNP 10_1101-2020_12_31_424971 97 19 , , , 10_1101-2020_12_31_424971 97 20 obs obs NN 10_1101-2020_12_31_424971 97 21 of of IN 10_1101-2020_12_31_424971 97 22 0.14 0.14 CD 10_1101-2020_12_31_424971 97 23 ± ± CD 10_1101-2020_12_31_424971 97 24 0.01 0.01 CD 10_1101-2020_12_31_424971 97 25 s–1 s–1 NNS 10_1101-2020_12_31_424971 97 26 , , , 10_1101-2020_12_31_424971 97 27 most most RBS 10_1101-2020_12_31_424971 97 28 likely likely JJ 10_1101-2020_12_31_424971 97 29 by by IN 10_1101-2020_12_31_424971 97 30 triplet triplet NN 10_1101-2020_12_31_424971 97 31 pairing pairing NN 10_1101-2020_12_31_424971 97 32 of of IN 10_1101-2020_12_31_424971 97 33 254 254 CD 10_1101-2020_12_31_424971 97 34 .CC .CC , 10_1101-2020_12_31_424971 97 35 - - HYPH 10_1101-2020_12_31_424971 97 36 BY by IN 10_1101-2020_12_31_424971 97 37 - - HYPH 10_1101-2020_12_31_424971 97 38 NC NC NNP 10_1101-2020_12_31_424971 97 39 - - HYPH 10_1101-2020_12_31_424971 97 40 ND ND NNP 10_1101-2020_12_31_424971 97 41 4.0 4.0 CD 10_1101-2020_12_31_424971 97 42 International International NNP 10_1101-2020_12_31_424971 97 43 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 97 44 under under IN 10_1101-2020_12_31_424971 97 45 a a DT 10_1101-2020_12_31_424971 97 46 ( ( -LRB- 10_1101-2020_12_31_424971 97 47 which which WDT 10_1101-2020_12_31_424971 97 48 was be VBD 10_1101-2020_12_31_424971 97 49 not not RB 10_1101-2020_12_31_424971 97 50 certified certify VBN 10_1101-2020_12_31_424971 97 51 by by IN 10_1101-2020_12_31_424971 97 52 peer peer NN 10_1101-2020_12_31_424971 97 53 review review NN 10_1101-2020_12_31_424971 97 54 ) ) -RRB- 10_1101-2020_12_31_424971 97 55 is be VBZ 10_1101-2020_12_31_424971 97 56 the the DT 10_1101-2020_12_31_424971 97 57 author author NN 10_1101-2020_12_31_424971 97 58 / / SYM 10_1101-2020_12_31_424971 97 59 funder funder NN 10_1101-2020_12_31_424971 97 60 , , , 10_1101-2020_12_31_424971 97 61 who who WP 10_1101-2020_12_31_424971 97 62 has have VBZ 10_1101-2020_12_31_424971 97 63 granted grant VBN 10_1101-2020_12_31_424971 97 64 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 97 65 a a DT 10_1101-2020_12_31_424971 97 66 license license NN 10_1101-2020_12_31_424971 97 67 to to TO 10_1101-2020_12_31_424971 97 68 display display VB 10_1101-2020_12_31_424971 97 69 the the DT 10_1101-2020_12_31_424971 97 70 preprint preprint NN 10_1101-2020_12_31_424971 97 71 in in IN 10_1101-2020_12_31_424971 97 72 perpetuity perpetuity NN 10_1101-2020_12_31_424971 97 73 . . . 10_1101-2020_12_31_424971 98 1 It -PRON- PRP 10_1101-2020_12_31_424971 98 2 is be VBZ 10_1101-2020_12_31_424971 98 3 made make VBN 10_1101-2020_12_31_424971 98 4 The the DT 10_1101-2020_12_31_424971 98 5 copyright copyright NN 10_1101-2020_12_31_424971 98 6 holder holder NN 10_1101-2020_12_31_424971 98 7 for for IN 10_1101-2020_12_31_424971 98 8 this this DT 10_1101-2020_12_31_424971 98 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 98 10 version version NN 10_1101-2020_12_31_424971 98 11 posted post VBD 10_1101-2020_12_31_424971 98 12 January January NNP 10_1101-2020_12_31_424971 98 13 2 2 CD 10_1101-2020_12_31_424971 98 14 , , , 10_1101-2020_12_31_424971 98 15 2021 2021 CD 10_1101-2020_12_31_424971 98 16 . . . 10_1101-2020_12_31_424971 98 17 ; ; : 10_1101-2020_12_31_424971 98 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 98 19 : : : 10_1101-2020_12_31_424971 98 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 98 21 preprint preprint NN 10_1101-2020_12_31_424971 98 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 98 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 98 24 11 11 CD 10_1101-2020_12_31_424971 98 25 nucleotides nucleotide VBZ 10_1101-2020_12_31_424971 98 26 34 34 CD 10_1101-2020_12_31_424971 98 27 - - SYM 10_1101-2020_12_31_424971 98 28 36 36 CD 10_1101-2020_12_31_424971 98 29 of of IN 10_1101-2020_12_31_424971 98 30 the the DT 10_1101-2020_12_31_424971 98 31 anticodon anticodon NN 10_1101-2020_12_31_424971 98 32 loop loop NN 10_1101-2020_12_31_424971 98 33 with with IN 10_1101-2020_12_31_424971 98 34 the the DT 10_1101-2020_12_31_424971 98 35 0-frame 0-frame CD 10_1101-2020_12_31_424971 98 36 of of IN 10_1101-2020_12_31_424971 98 37 the the DT 10_1101-2020_12_31_424971 98 38 CCC CCC NNP 10_1101-2020_12_31_424971 98 39 - - HYPH 10_1101-2020_12_31_424971 98 40 C C NNP 10_1101-2020_12_31_424971 98 41 motif motif NN 10_1101-2020_12_31_424971 98 42 ( ( -LRB- 10_1101-2020_12_31_424971 98 43 Figure Figure NNP 10_1101-2020_12_31_424971 98 44 4a 4a CD 10_1101-2020_12_31_424971 98 45 ) ) -RRB- 10_1101-2020_12_31_424971 98 46 . . . 10_1101-2020_12_31_424971 99 1 In in IN 10_1101-2020_12_31_424971 99 2 255 255 CD 10_1101-2020_12_31_424971 99 3 contrast contrast NN 10_1101-2020_12_31_424971 99 4 , , , 10_1101-2020_12_31_424971 99 5 the the DT 10_1101-2020_12_31_424971 99 6 G34C G34C NNP 10_1101-2020_12_31_424971 99 7 variant variant NN 10_1101-2020_12_31_424971 99 8 resulted result VBD 10_1101-2020_12_31_424971 99 9 in in IN 10_1101-2020_12_31_424971 99 10 a a DT 10_1101-2020_12_31_424971 99 11 fMP fmp CD 10_1101-2020_12_31_424971 99 12 yield yield NN 10_1101-2020_12_31_424971 99 13 of of IN 10_1101-2020_12_31_424971 99 14 30 30 CD 10_1101-2020_12_31_424971 99 15 % % NN 10_1101-2020_12_31_424971 99 16 and and CC 10_1101-2020_12_31_424971 99 17 a a DT 10_1101-2020_12_31_424971 99 18 kfMP kfMP NNP 10_1101-2020_12_31_424971 99 19 , , , 10_1101-2020_12_31_424971 99 20 obs obs NN 10_1101-2020_12_31_424971 99 21 of of IN 10_1101-2020_12_31_424971 99 22 0.28 0.28 CD 10_1101-2020_12_31_424971 99 23 ± ± CD 10_1101-2020_12_31_424971 99 24 0.04 0.04 CD 10_1101-2020_12_31_424971 99 25 s–1 s–1 NNS 10_1101-2020_12_31_424971 99 26 , , , 10_1101-2020_12_31_424971 99 27 most most JJS 10_1101-2020_12_31_424971 99 28 256 256 CD 10_1101-2020_12_31_424971 99 29 likely likely JJ 10_1101-2020_12_31_424971 99 30 by by IN 10_1101-2020_12_31_424971 99 31 triplet triplet NN 10_1101-2020_12_31_424971 99 32 pairing pairing NN 10_1101-2020_12_31_424971 99 33 of of IN 10_1101-2020_12_31_424971 99 34 nucleotides nucleotide VBZ 10_1101-2020_12_31_424971 99 35 35 35 CD 10_1101-2020_12_31_424971 99 36 - - SYM 10_1101-2020_12_31_424971 99 37 37 37 CD 10_1101-2020_12_31_424971 99 38 of of IN 10_1101-2020_12_31_424971 99 39 the the DT 10_1101-2020_12_31_424971 99 40 anticodon anticodon NN 10_1101-2020_12_31_424971 99 41 loop loop NN 10_1101-2020_12_31_424971 99 42 with with IN 10_1101-2020_12_31_424971 99 43 the the DT 10_1101-2020_12_31_424971 99 44 0-frame 0-frame CD 10_1101-2020_12_31_424971 99 45 of of IN 10_1101-2020_12_31_424971 99 46 the the DT 10_1101-2020_12_31_424971 99 47 CCC CCC NNP 10_1101-2020_12_31_424971 99 48 - - HYPH 10_1101-2020_12_31_424971 99 49 C C NNP 10_1101-2020_12_31_424971 99 50 257 257 CD 10_1101-2020_12_31_424971 99 51 motif motif NN 10_1101-2020_12_31_424971 99 52 ( ( -LRB- 10_1101-2020_12_31_424971 99 53 Figure Figure NNP 10_1101-2020_12_31_424971 99 54 4b 4b NNP 10_1101-2020_12_31_424971 99 55 ) ) -RRB- 10_1101-2020_12_31_424971 99 56 . . . 10_1101-2020_12_31_424971 100 1 Our -PRON- PRP$ 10_1101-2020_12_31_424971 100 2 interpretation interpretation NN 10_1101-2020_12_31_424971 100 3 that that WDT 10_1101-2020_12_31_424971 100 4 nucleotides nucleotide VBZ 10_1101-2020_12_31_424971 100 5 35 35 CD 10_1101-2020_12_31_424971 100 6 - - SYM 10_1101-2020_12_31_424971 100 7 37 37 CD 10_1101-2020_12_31_424971 100 8 of of IN 10_1101-2020_12_31_424971 100 9 the the DT 10_1101-2020_12_31_424971 100 10 anticodon anticodon NNP 10_1101-2020_12_31_424971 100 11 loop loop NN 10_1101-2020_12_31_424971 100 12 of of IN 10_1101-2020_12_31_424971 100 13 the the DT 10_1101-2020_12_31_424971 100 14 G34C G34C NNP 10_1101-2020_12_31_424971 100 15 258 258 CD 10_1101-2020_12_31_424971 100 16 variant variant JJ 10_1101-2020_12_31_424971 100 17 most most RBS 10_1101-2020_12_31_424971 100 18 likely likely JJ 10_1101-2020_12_31_424971 100 19 triplet triplet NN 10_1101-2020_12_31_424971 100 20 pair pair NN 10_1101-2020_12_31_424971 100 21 with with IN 10_1101-2020_12_31_424971 100 22 the the DT 10_1101-2020_12_31_424971 100 23 0-frame 0-frame CD 10_1101-2020_12_31_424971 100 24 of of IN 10_1101-2020_12_31_424971 100 25 the the DT 10_1101-2020_12_31_424971 100 26 CCC CCC NNP 10_1101-2020_12_31_424971 100 27 - - HYPH 10_1101-2020_12_31_424971 100 28 C C NNP 10_1101-2020_12_31_424971 100 29 motif motif NN 10_1101-2020_12_31_424971 100 30 is be VBZ 10_1101-2020_12_31_424971 100 31 consistent consistent JJ 10_1101-2020_12_31_424971 100 32 with with IN 10_1101-2020_12_31_424971 100 33 the the DT 10_1101-2020_12_31_424971 100 34 259 259 CD 10_1101-2020_12_31_424971 100 35 observations observation NNS 10_1101-2020_12_31_424971 100 36 that that WDT 10_1101-2020_12_31_424971 100 37 the the DT 10_1101-2020_12_31_424971 100 38 fMP fMP NNS 10_1101-2020_12_31_424971 100 39 yield yield NN 10_1101-2020_12_31_424971 100 40 and and CC 10_1101-2020_12_31_424971 100 41 kfMP kfMP NNP 10_1101-2020_12_31_424971 100 42 , , , 10_1101-2020_12_31_424971 100 43 obs obs NN 10_1101-2020_12_31_424971 100 44 of of IN 10_1101-2020_12_31_424971 100 45 the the DT 10_1101-2020_12_31_424971 100 46 G34C G34C NNP 10_1101-2020_12_31_424971 100 47 variant variant NN 10_1101-2020_12_31_424971 100 48 are be VBP 10_1101-2020_12_31_424971 100 49 similar similar JJ 10_1101-2020_12_31_424971 100 50 and and CC 10_1101-2020_12_31_424971 100 51 2-fold 2-fold CD 10_1101-2020_12_31_424971 100 52 higher high JJR 10_1101-2020_12_31_424971 100 53 , , , 10_1101-2020_12_31_424971 100 54 260 260 CD 10_1101-2020_12_31_424971 100 55 respectively respectively RB 10_1101-2020_12_31_424971 100 56 , , , 10_1101-2020_12_31_424971 100 57 than than IN 10_1101-2020_12_31_424971 100 58 those those DT 10_1101-2020_12_31_424971 100 59 of of IN 10_1101-2020_12_31_424971 100 60 the the DT 10_1101-2020_12_31_424971 100 61 G37C G37C NNP 10_1101-2020_12_31_424971 100 62 variant variant NN 10_1101-2020_12_31_424971 100 63 . . . 10_1101-2020_12_31_424971 101 1 If if IN 10_1101-2020_12_31_424971 101 2 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 101 3 34 34 CD 10_1101-2020_12_31_424971 101 4 - - SYM 10_1101-2020_12_31_424971 101 5 36 36 CD 10_1101-2020_12_31_424971 101 6 of of IN 10_1101-2020_12_31_424971 101 7 the the DT 10_1101-2020_12_31_424971 101 8 anticodon anticodon NNP 10_1101-2020_12_31_424971 101 9 loop loop NN 10_1101-2020_12_31_424971 101 10 of of IN 10_1101-2020_12_31_424971 101 11 the the DT 10_1101-2020_12_31_424971 101 12 261 261 CD 10_1101-2020_12_31_424971 101 13 G34C G34C NNP 10_1101-2020_12_31_424971 101 14 variant variant NN 10_1101-2020_12_31_424971 101 15 were be VBD 10_1101-2020_12_31_424971 101 16 to to TO 10_1101-2020_12_31_424971 101 17 form form VB 10_1101-2020_12_31_424971 101 18 a a DT 10_1101-2020_12_31_424971 101 19 triplet triplet NN 10_1101-2020_12_31_424971 101 20 pair pair NN 10_1101-2020_12_31_424971 101 21 with with IN 10_1101-2020_12_31_424971 101 22 the the DT 10_1101-2020_12_31_424971 101 23 CCC CCC NNP 10_1101-2020_12_31_424971 101 24 - - HYPH 10_1101-2020_12_31_424971 101 25 C C NNP 10_1101-2020_12_31_424971 101 26 motif motif NN 10_1101-2020_12_31_424971 101 27 , , , 10_1101-2020_12_31_424971 101 28 we -PRON- PRP 10_1101-2020_12_31_424971 101 29 would would MD 10_1101-2020_12_31_424971 101 30 have have VB 10_1101-2020_12_31_424971 101 31 expected expect VBN 10_1101-2020_12_31_424971 101 32 it -PRON- PRP 10_1101-2020_12_31_424971 101 33 to to TO 10_1101-2020_12_31_424971 101 34 pair pair VB 10_1101-2020_12_31_424971 101 35 262 262 CD 10_1101-2020_12_31_424971 101 36 in in IN 10_1101-2020_12_31_424971 101 37 the the DT 10_1101-2020_12_31_424971 101 38 +2-frame +2-frame NN 10_1101-2020_12_31_424971 101 39 , , , 10_1101-2020_12_31_424971 101 40 which which WDT 10_1101-2020_12_31_424971 101 41 would would MD 10_1101-2020_12_31_424971 101 42 have have VB 10_1101-2020_12_31_424971 101 43 most most RBS 10_1101-2020_12_31_424971 101 44 likely likely RB 10_1101-2020_12_31_424971 101 45 reduced reduce VBN 10_1101-2020_12_31_424971 101 46 the the DT 10_1101-2020_12_31_424971 101 47 fMP fmp CD 10_1101-2020_12_31_424971 101 48 yield yield NN 10_1101-2020_12_31_424971 101 49 and and CC 10_1101-2020_12_31_424971 101 50 kfMP kfMP NNP 10_1101-2020_12_31_424971 101 51 , , , 10_1101-2020_12_31_424971 101 52 obs obs NN 10_1101-2020_12_31_424971 101 53 of of IN 10_1101-2020_12_31_424971 101 54 the the DT 10_1101-2020_12_31_424971 101 55 G34C G34C NNP 10_1101-2020_12_31_424971 101 56 263 263 CD 10_1101-2020_12_31_424971 101 57 variant variant JJ 10_1101-2020_12_31_424971 101 58 relative relative NN 10_1101-2020_12_31_424971 101 59 to to IN 10_1101-2020_12_31_424971 101 60 the the DT 10_1101-2020_12_31_424971 101 61 G37C G37C NNP 10_1101-2020_12_31_424971 101 62 variant variant NN 10_1101-2020_12_31_424971 101 63 . . . 10_1101-2020_12_31_424971 102 1 These these DT 10_1101-2020_12_31_424971 102 2 results result NNS 10_1101-2020_12_31_424971 102 3 suggest suggest VBP 10_1101-2020_12_31_424971 102 4 that that IN 10_1101-2020_12_31_424971 102 5 G37-state G37-state NNP 10_1101-2020_12_31_424971 102 6 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 102 7 exhibits exhibit VBZ 10_1101-2020_12_31_424971 102 8 some some DT 10_1101-2020_12_31_424971 102 9 264 264 CD 10_1101-2020_12_31_424971 102 10 plasticity plasticity NN 10_1101-2020_12_31_424971 102 11 as as IN 10_1101-2020_12_31_424971 102 12 to to IN 10_1101-2020_12_31_424971 102 13 whether whether IN 10_1101-2020_12_31_424971 102 14 it -PRON- PRP 10_1101-2020_12_31_424971 102 15 can can MD 10_1101-2020_12_31_424971 102 16 undergo undergo VB 10_1101-2020_12_31_424971 102 17 triplet triplet NN 10_1101-2020_12_31_424971 102 18 pairing pair VBG 10_1101-2020_12_31_424971 102 19 with with IN 10_1101-2020_12_31_424971 102 20 anticodon anticodon NNP 10_1101-2020_12_31_424971 102 21 loop loop NNP 10_1101-2020_12_31_424971 102 22 nucleotides nucleotide VBZ 10_1101-2020_12_31_424971 102 23 34 34 CD 10_1101-2020_12_31_424971 102 24 - - SYM 10_1101-2020_12_31_424971 102 25 36 36 CD 10_1101-2020_12_31_424971 102 26 or or CC 10_1101-2020_12_31_424971 102 27 35 35 CD 10_1101-2020_12_31_424971 102 28 - - SYM 10_1101-2020_12_31_424971 102 29 265 265 CD 10_1101-2020_12_31_424971 102 30 37 37 CD 10_1101-2020_12_31_424971 102 31 , , , 10_1101-2020_12_31_424971 102 32 consistent consistent JJ 10_1101-2020_12_31_424971 102 33 with with IN 10_1101-2020_12_31_424971 102 34 a a DT 10_1101-2020_12_31_424971 102 35 previous previous JJ 10_1101-2020_12_31_424971 102 36 study33 study33 NN 10_1101-2020_12_31_424971 102 37 . . . 10_1101-2020_12_31_424971 103 1 266 266 CD 10_1101-2020_12_31_424971 103 2 267 267 CD 10_1101-2020_12_31_424971 103 3 SufB2 sufb2 CD 10_1101-2020_12_31_424971 103 4 shifts shift NNS 10_1101-2020_12_31_424971 103 5 to to IN 10_1101-2020_12_31_424971 103 6 the the DT 10_1101-2020_12_31_424971 103 7 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 103 8 during during IN 10_1101-2020_12_31_424971 103 9 translocation translocation NN 10_1101-2020_12_31_424971 103 10 268 268 CD 10_1101-2020_12_31_424971 103 11 Although although IN 10_1101-2020_12_31_424971 103 12 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 103 13 uses use VBZ 10_1101-2020_12_31_424971 103 14 triplet triplet NN 10_1101-2020_12_31_424971 103 15 pairing pair VBG 10_1101-2020_12_31_424971 103 16 in in IN 10_1101-2020_12_31_424971 103 17 the the DT 10_1101-2020_12_31_424971 103 18 0-frame 0-frame CD 10_1101-2020_12_31_424971 103 19 when when WRB 10_1101-2020_12_31_424971 103 20 it -PRON- PRP 10_1101-2020_12_31_424971 103 21 is be VBZ 10_1101-2020_12_31_424971 103 22 delivered deliver VBN 10_1101-2020_12_31_424971 103 23 to to IN 10_1101-2020_12_31_424971 103 24 the the DT 10_1101-2020_12_31_424971 103 25 A A NNP 10_1101-2020_12_31_424971 103 26 site site NN 10_1101-2020_12_31_424971 103 27 , , , 10_1101-2020_12_31_424971 103 28 it -PRON- PRP 10_1101-2020_12_31_424971 103 29 is be VBZ 10_1101-2020_12_31_424971 103 30 a a DT 10_1101-2020_12_31_424971 103 31 269 269 CD 10_1101-2020_12_31_424971 103 32 highly highly RB 10_1101-2020_12_31_424971 103 33 efficient efficient JJ 10_1101-2020_12_31_424971 103 34 +1-frameshifting +1-frameshifting NN 10_1101-2020_12_31_424971 103 35 tRNA trna NN 10_1101-2020_12_31_424971 103 36 ( ( -LRB- 10_1101-2020_12_31_424971 103 37 Figure figure NN 10_1101-2020_12_31_424971 103 38 2 2 CD 10_1101-2020_12_31_424971 103 39 ) ) -RRB- 10_1101-2020_12_31_424971 103 40 . . . 10_1101-2020_12_31_424971 104 1 We -PRON- PRP 10_1101-2020_12_31_424971 104 2 therefore therefore RB 10_1101-2020_12_31_424971 104 3 asked ask VBD 10_1101-2020_12_31_424971 104 4 whether whether IN 10_1101-2020_12_31_424971 104 5 +1 +1 NNS 10_1101-2020_12_31_424971 104 6 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 104 7 270 270 CD 10_1101-2020_12_31_424971 104 8 occurs occur NNS 10_1101-2020_12_31_424971 104 9 during during IN 10_1101-2020_12_31_424971 104 10 or or CC 10_1101-2020_12_31_424971 104 11 after after IN 10_1101-2020_12_31_424971 104 12 translocation translocation NN 10_1101-2020_12_31_424971 104 13 of of IN 10_1101-2020_12_31_424971 104 14 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 104 15 into into IN 10_1101-2020_12_31_424971 104 16 the the DT 10_1101-2020_12_31_424971 104 17 P p NN 10_1101-2020_12_31_424971 104 18 site site NN 10_1101-2020_12_31_424971 104 19 . . . 10_1101-2020_12_31_424971 105 1 We -PRON- PRP 10_1101-2020_12_31_424971 105 2 addressed address VBD 10_1101-2020_12_31_424971 105 3 this this DT 10_1101-2020_12_31_424971 105 4 question question NN 10_1101-2020_12_31_424971 105 5 by by IN 10_1101-2020_12_31_424971 105 6 271 271 CD 10_1101-2020_12_31_424971 105 7 adapting adapt VBG 10_1101-2020_12_31_424971 105 8 our -PRON- PRP$ 10_1101-2020_12_31_424971 105 9 previously previously RB 10_1101-2020_12_31_424971 105 10 developed develop VBN 10_1101-2020_12_31_424971 105 11 tripeptide tripeptide NN 10_1101-2020_12_31_424971 105 12 formation formation NN 10_1101-2020_12_31_424971 105 13 assays29 assays29 NN 10_1101-2020_12_31_424971 105 14 . . . 10_1101-2020_12_31_424971 106 1 We -PRON- PRP 10_1101-2020_12_31_424971 106 2 rapidly rapidly RB 10_1101-2020_12_31_424971 106 3 delivered deliver VBD 10_1101-2020_12_31_424971 106 4 EF EF NNP 10_1101-2020_12_31_424971 106 5 - - HYPH 10_1101-2020_12_31_424971 106 6 G G NNP 10_1101-2020_12_31_424971 106 7 and and CC 10_1101-2020_12_31_424971 106 8 272 272 CD 10_1101-2020_12_31_424971 106 9 an an DT 10_1101-2020_12_31_424971 106 10 equimolar equimolar JJ 10_1101-2020_12_31_424971 106 11 mixture mixture NN 10_1101-2020_12_31_424971 106 12 of of IN 10_1101-2020_12_31_424971 106 13 G37-state G37-state NNP 10_1101-2020_12_31_424971 106 14 SufB2- sufb2- NN 10_1101-2020_12_31_424971 106 15 , , , 10_1101-2020_12_31_424971 106 16 tRNAVal- tRNAVal- NNP 10_1101-2020_12_31_424971 106 17 , , , 10_1101-2020_12_31_424971 106 18 and and CC 10_1101-2020_12_31_424971 106 19 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 106 20 - - HYPH 10_1101-2020_12_31_424971 106 21 TCs TCs NNP 10_1101-2020_12_31_424971 106 22 to to IN 10_1101-2020_12_31_424971 106 23 70S 70S NNP 10_1101-2020_12_31_424971 106 24 ICs ic NNS 10_1101-2020_12_31_424971 106 25 assembled assemble VBD 10_1101-2020_12_31_424971 106 26 on on IN 10_1101-2020_12_31_424971 106 27 273 273 CD 10_1101-2020_12_31_424971 106 28 an an DT 10_1101-2020_12_31_424971 106 29 mRNA mrna NN 10_1101-2020_12_31_424971 106 30 in in IN 10_1101-2020_12_31_424971 106 31 which which WDT 10_1101-2020_12_31_424971 106 32 the the DT 10_1101-2020_12_31_424971 106 33 2nd 2nd JJ 10_1101-2020_12_31_424971 106 34 codon codon NN 10_1101-2020_12_31_424971 106 35 was be VBD 10_1101-2020_12_31_424971 106 36 a a DT 10_1101-2020_12_31_424971 106 37 CCC CCC NNP 10_1101-2020_12_31_424971 106 38 - - HYPH 10_1101-2020_12_31_424971 106 39 C C NNP 10_1101-2020_12_31_424971 106 40 motif motif NN 10_1101-2020_12_31_424971 106 41 and and CC 10_1101-2020_12_31_424971 106 42 the the DT 10_1101-2020_12_31_424971 106 43 3rd 3rd JJ 10_1101-2020_12_31_424971 106 44 codon codon NN 10_1101-2020_12_31_424971 106 45 was be VBD 10_1101-2020_12_31_424971 106 46 either either CC 10_1101-2020_12_31_424971 106 47 a a DT 10_1101-2020_12_31_424971 106 48 GUU GUU NNP 10_1101-2020_12_31_424971 106 49 codon codon NN 10_1101-2020_12_31_424971 106 50 274 274 CD 10_1101-2020_12_31_424971 106 51 encoding encode VBG 10_1101-2020_12_31_424971 106 52 Val Val NNP 10_1101-2020_12_31_424971 106 53 in in IN 10_1101-2020_12_31_424971 106 54 the the DT 10_1101-2020_12_31_424971 106 55 +1 +1 NNS 10_1101-2020_12_31_424971 106 56 frame frame NN 10_1101-2020_12_31_424971 106 57 or or CC 10_1101-2020_12_31_424971 106 58 a a DT 10_1101-2020_12_31_424971 106 59 CGU cgu NN 10_1101-2020_12_31_424971 106 60 codon codon NN 10_1101-2020_12_31_424971 106 61 encoding encode VBG 10_1101-2020_12_31_424971 106 62 Arg arg NN 10_1101-2020_12_31_424971 106 63 in in IN 10_1101-2020_12_31_424971 106 64 the the DT 10_1101-2020_12_31_424971 106 65 0-frame 0-frame CD 10_1101-2020_12_31_424971 106 66 . . . 10_1101-2020_12_31_424971 107 1 As as RB 10_1101-2020_12_31_424971 107 2 soon soon RB 10_1101-2020_12_31_424971 107 3 as as IN 10_1101-2020_12_31_424971 107 4 275 275 CD 10_1101-2020_12_31_424971 107 5 translocation translocation NN 10_1101-2020_12_31_424971 107 6 of of IN 10_1101-2020_12_31_424971 107 7 the the DT 10_1101-2020_12_31_424971 107 8 PRE PRE NNP 10_1101-2020_12_31_424971 107 9 complex complex NN 10_1101-2020_12_31_424971 107 10 and and CC 10_1101-2020_12_31_424971 107 11 the the DT 10_1101-2020_12_31_424971 107 12 associated associated JJ 10_1101-2020_12_31_424971 107 13 movement movement NN 10_1101-2020_12_31_424971 107 14 of of IN 10_1101-2020_12_31_424971 107 15 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 107 16 from from IN 10_1101-2020_12_31_424971 107 17 the the DT 10_1101-2020_12_31_424971 107 18 P p NN 10_1101-2020_12_31_424971 107 19 to to IN 10_1101-2020_12_31_424971 107 20 A a DT 10_1101-2020_12_31_424971 107 21 sites site NNS 10_1101-2020_12_31_424971 107 22 276 276 CD 10_1101-2020_12_31_424971 107 23 formed form VBD 10_1101-2020_12_31_424971 107 24 a a DT 10_1101-2020_12_31_424971 107 25 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 107 26 post post NN 10_1101-2020_12_31_424971 107 27 - - JJ 10_1101-2020_12_31_424971 107 28 translocation translocation JJ 10_1101-2020_12_31_424971 107 29 ( ( -LRB- 10_1101-2020_12_31_424971 107 30 POST POST NNP 10_1101-2020_12_31_424971 107 31 ) ) -RRB- 10_1101-2020_12_31_424971 107 32 complex complex NN 10_1101-2020_12_31_424971 107 33 with with IN 10_1101-2020_12_31_424971 107 34 an an DT 10_1101-2020_12_31_424971 107 35 empty empty JJ 10_1101-2020_12_31_424971 107 36 A a DT 10_1101-2020_12_31_424971 107 37 site site NN 10_1101-2020_12_31_424971 107 38 in in IN 10_1101-2020_12_31_424971 107 39 these these DT 10_1101-2020_12_31_424971 107 40 experiments experiment NNS 10_1101-2020_12_31_424971 107 41 , , , 10_1101-2020_12_31_424971 107 42 277 277 CD 10_1101-2020_12_31_424971 107 43 tRNAVal- tRNAVal- NNP 10_1101-2020_12_31_424971 107 44 and and CC 10_1101-2020_12_31_424971 107 45 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 107 46 - - HYPH 10_1101-2020_12_31_424971 107 47 TC TC NNP 10_1101-2020_12_31_424971 107 48 would would MD 10_1101-2020_12_31_424971 107 49 compete compete VB 10_1101-2020_12_31_424971 107 50 for for IN 10_1101-2020_12_31_424971 107 51 the the DT 10_1101-2020_12_31_424971 107 52 codon codon NN 10_1101-2020_12_31_424971 107 53 at at IN 10_1101-2020_12_31_424971 107 54 the the DT 10_1101-2020_12_31_424971 107 55 A A NNP 10_1101-2020_12_31_424971 107 56 site site NN 10_1101-2020_12_31_424971 107 57 to to TO 10_1101-2020_12_31_424971 107 58 promote promote VB 10_1101-2020_12_31_424971 107 59 formation formation NN 10_1101-2020_12_31_424971 107 60 of of IN 10_1101-2020_12_31_424971 107 61 an an DT 10_1101-2020_12_31_424971 107 62 278 278 CD 10_1101-2020_12_31_424971 107 63 fMPV fmpv NN 10_1101-2020_12_31_424971 107 64 tripeptide tripeptide NN 10_1101-2020_12_31_424971 107 65 or or CC 10_1101-2020_12_31_424971 107 66 an an DT 10_1101-2020_12_31_424971 107 67 fMPR fmpr JJ 10_1101-2020_12_31_424971 107 68 tripeptide tripeptide NN 10_1101-2020_12_31_424971 107 69 . . . 10_1101-2020_12_31_424971 108 1 Thus thus RB 10_1101-2020_12_31_424971 108 2 , , , 10_1101-2020_12_31_424971 108 3 the the DT 10_1101-2020_12_31_424971 108 4 fMPV fmpv JJ 10_1101-2020_12_31_424971 108 5 yield yield NN 10_1101-2020_12_31_424971 108 6 and and CC 10_1101-2020_12_31_424971 108 7 kfMPV kfmpv NN 10_1101-2020_12_31_424971 108 8 , , , 10_1101-2020_12_31_424971 108 9 obs obs NN 10_1101-2020_12_31_424971 108 10 report report NN 10_1101-2020_12_31_424971 108 11 on on IN 10_1101-2020_12_31_424971 108 12 the the DT 10_1101-2020_12_31_424971 108 13 sub-279 sub-279 NNP 10_1101-2020_12_31_424971 108 14 population population NN 10_1101-2020_12_31_424971 108 15 of of IN 10_1101-2020_12_31_424971 108 16 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 108 17 that that WDT 10_1101-2020_12_31_424971 108 18 shifted shift VBD 10_1101-2020_12_31_424971 108 19 to to IN 10_1101-2020_12_31_424971 108 20 the the DT 10_1101-2020_12_31_424971 108 21 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 108 22 , , , 10_1101-2020_12_31_424971 108 23 whereas whereas IN 10_1101-2020_12_31_424971 108 24 the the DT 10_1101-2020_12_31_424971 108 25 fMPR fmpr JJ 10_1101-2020_12_31_424971 108 26 yield yield NN 10_1101-2020_12_31_424971 108 27 and and CC 10_1101-2020_12_31_424971 108 28 kfMPR kfmpr NN 10_1101-2020_12_31_424971 108 29 , , , 10_1101-2020_12_31_424971 108 30 obs obs NN 10_1101-2020_12_31_424971 108 31 report report NN 10_1101-2020_12_31_424971 108 32 on on IN 10_1101-2020_12_31_424971 108 33 280 280 CD 10_1101-2020_12_31_424971 108 34 .CC .CC NFP 10_1101-2020_12_31_424971 108 35 - - HYPH 10_1101-2020_12_31_424971 108 36 BY by IN 10_1101-2020_12_31_424971 108 37 - - HYPH 10_1101-2020_12_31_424971 108 38 NC NC NNP 10_1101-2020_12_31_424971 108 39 - - HYPH 10_1101-2020_12_31_424971 108 40 ND ND NNP 10_1101-2020_12_31_424971 108 41 4.0 4.0 CD 10_1101-2020_12_31_424971 108 42 International International NNP 10_1101-2020_12_31_424971 108 43 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 108 44 under under IN 10_1101-2020_12_31_424971 108 45 a a DT 10_1101-2020_12_31_424971 108 46 ( ( -LRB- 10_1101-2020_12_31_424971 108 47 which which WDT 10_1101-2020_12_31_424971 108 48 was be VBD 10_1101-2020_12_31_424971 108 49 not not RB 10_1101-2020_12_31_424971 108 50 certified certify VBN 10_1101-2020_12_31_424971 108 51 by by IN 10_1101-2020_12_31_424971 108 52 peer peer NN 10_1101-2020_12_31_424971 108 53 review review NN 10_1101-2020_12_31_424971 108 54 ) ) -RRB- 10_1101-2020_12_31_424971 108 55 is be VBZ 10_1101-2020_12_31_424971 108 56 the the DT 10_1101-2020_12_31_424971 108 57 author author NN 10_1101-2020_12_31_424971 108 58 / / SYM 10_1101-2020_12_31_424971 108 59 funder funder NN 10_1101-2020_12_31_424971 108 60 , , , 10_1101-2020_12_31_424971 108 61 who who WP 10_1101-2020_12_31_424971 108 62 has have VBZ 10_1101-2020_12_31_424971 108 63 granted grant VBN 10_1101-2020_12_31_424971 108 64 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 108 65 a a DT 10_1101-2020_12_31_424971 108 66 license license NN 10_1101-2020_12_31_424971 108 67 to to TO 10_1101-2020_12_31_424971 108 68 display display VB 10_1101-2020_12_31_424971 108 69 the the DT 10_1101-2020_12_31_424971 108 70 preprint preprint NN 10_1101-2020_12_31_424971 108 71 in in IN 10_1101-2020_12_31_424971 108 72 perpetuity perpetuity NN 10_1101-2020_12_31_424971 108 73 . . . 10_1101-2020_12_31_424971 109 1 It -PRON- PRP 10_1101-2020_12_31_424971 109 2 is be VBZ 10_1101-2020_12_31_424971 109 3 made make VBN 10_1101-2020_12_31_424971 109 4 The the DT 10_1101-2020_12_31_424971 109 5 copyright copyright NN 10_1101-2020_12_31_424971 109 6 holder holder NN 10_1101-2020_12_31_424971 109 7 for for IN 10_1101-2020_12_31_424971 109 8 this this DT 10_1101-2020_12_31_424971 109 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 109 10 version version NN 10_1101-2020_12_31_424971 109 11 posted post VBD 10_1101-2020_12_31_424971 109 12 January January NNP 10_1101-2020_12_31_424971 109 13 2 2 CD 10_1101-2020_12_31_424971 109 14 , , , 10_1101-2020_12_31_424971 109 15 2021 2021 CD 10_1101-2020_12_31_424971 109 16 . . . 10_1101-2020_12_31_424971 109 17 ; ; : 10_1101-2020_12_31_424971 109 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 109 19 : : : 10_1101-2020_12_31_424971 109 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 109 21 preprint preprint NN 10_1101-2020_12_31_424971 109 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NNP 10_1101-2020_12_31_424971 109 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 109 24 12 12 CD 10_1101-2020_12_31_424971 109 25 the the DT 10_1101-2020_12_31_424971 109 26 sub sub NN 10_1101-2020_12_31_424971 109 27 - - NN 10_1101-2020_12_31_424971 109 28 population population NN 10_1101-2020_12_31_424971 109 29 that that WDT 10_1101-2020_12_31_424971 109 30 remained remain VBD 10_1101-2020_12_31_424971 109 31 in in IN 10_1101-2020_12_31_424971 109 32 the the DT 10_1101-2020_12_31_424971 109 33 0-frame29,34 0-frame29,34 CD 10_1101-2020_12_31_424971 109 34 . . . 10_1101-2020_12_31_424971 110 1 The the DT 10_1101-2020_12_31_424971 110 2 results result NNS 10_1101-2020_12_31_424971 110 3 showed show VBD 10_1101-2020_12_31_424971 110 4 that that IN 10_1101-2020_12_31_424971 110 5 the the DT 10_1101-2020_12_31_424971 110 6 yield yield NN 10_1101-2020_12_31_424971 110 7 of of IN 10_1101-2020_12_31_424971 110 8 fMPV fMPV NNP 10_1101-2020_12_31_424971 110 9 281 281 CD 10_1101-2020_12_31_424971 110 10 was be VBD 10_1101-2020_12_31_424971 110 11 much much RB 10_1101-2020_12_31_424971 110 12 higher high JJR 10_1101-2020_12_31_424971 110 13 than than IN 10_1101-2020_12_31_424971 110 14 that that DT 10_1101-2020_12_31_424971 110 15 of of IN 10_1101-2020_12_31_424971 110 16 fMPR fmpr JJ 10_1101-2020_12_31_424971 110 17 ( ( -LRB- 10_1101-2020_12_31_424971 110 18 90 90 CD 10_1101-2020_12_31_424971 110 19 % % NN 10_1101-2020_12_31_424971 110 20 vs. vs. NN 10_1101-2020_12_31_424971 110 21 10 10 CD 10_1101-2020_12_31_424971 110 22 % % NN 10_1101-2020_12_31_424971 110 23 , , , 10_1101-2020_12_31_424971 110 24 Figure Figure NNP 10_1101-2020_12_31_424971 110 25 5a 5a NNP 10_1101-2020_12_31_424971 110 26 ) ) -RRB- 10_1101-2020_12_31_424971 110 27 , , , 10_1101-2020_12_31_424971 110 28 demonstrating demonstrate VBG 10_1101-2020_12_31_424971 110 29 the the DT 10_1101-2020_12_31_424971 110 30 high high JJ 10_1101-2020_12_31_424971 110 31 efficiency efficiency NN 10_1101-2020_12_31_424971 110 32 282 282 CD 10_1101-2020_12_31_424971 110 33 with with IN 10_1101-2020_12_31_424971 110 34 which which WDT 10_1101-2020_12_31_424971 110 35 G37-state G37-state NNP 10_1101-2020_12_31_424971 110 36 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 110 37 induces induce VBZ 10_1101-2020_12_31_424971 110 38 +1 +1 : 10_1101-2020_12_31_424971 110 39 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 110 40 . . . 10_1101-2020_12_31_424971 111 1 Notably notably RB 10_1101-2020_12_31_424971 111 2 , , , 10_1101-2020_12_31_424971 111 3 relative relative JJ 10_1101-2020_12_31_424971 111 4 to to IN 10_1101-2020_12_31_424971 111 5 the the DT 10_1101-2020_12_31_424971 111 6 +1 +1 NNS 10_1101-2020_12_31_424971 111 7 frameshifting frameshifting NN 10_1101-2020_12_31_424971 111 8 of of IN 10_1101-2020_12_31_424971 111 9 283 283 CD 10_1101-2020_12_31_424971 111 10 ProL ProL NNP 10_1101-2020_12_31_424971 111 11 we -PRON- PRP 10_1101-2020_12_31_424971 111 12 have have VBP 10_1101-2020_12_31_424971 111 13 previously previously RB 10_1101-2020_12_31_424971 111 14 reported29 reported29 NNP 10_1101-2020_12_31_424971 111 15 , , , 10_1101-2020_12_31_424971 111 16 kfMPV kfmpv NN 10_1101-2020_12_31_424971 111 17 , , , 10_1101-2020_12_31_424971 111 18 obs obs NN 10_1101-2020_12_31_424971 111 19 of of IN 10_1101-2020_12_31_424971 111 20 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 111 21 ( ( -LRB- 10_1101-2020_12_31_424971 111 22 0.09 0.09 CD 10_1101-2020_12_31_424971 111 23 s–1 s–1 NNS 10_1101-2020_12_31_424971 111 24 ) ) -RRB- 10_1101-2020_12_31_424971 111 25 was be VBD 10_1101-2020_12_31_424971 111 26 comparable comparable JJ 10_1101-2020_12_31_424971 111 27 to to IN 10_1101-2020_12_31_424971 111 28 the the DT 10_1101-2020_12_31_424971 111 29 rate rate NN 10_1101-2020_12_31_424971 111 30 of of IN 10_1101-2020_12_31_424971 111 31 +1 +1 NNS 10_1101-2020_12_31_424971 111 32 284 284 CD 10_1101-2020_12_31_424971 111 33 frameshifting frameshifting NN 10_1101-2020_12_31_424971 111 34 of of IN 10_1101-2020_12_31_424971 111 35 ProL prol NN 10_1101-2020_12_31_424971 111 36 during during IN 10_1101-2020_12_31_424971 111 37 translocation translocation NN 10_1101-2020_12_31_424971 111 38 ( ( -LRB- 10_1101-2020_12_31_424971 111 39 0.1 0.1 CD 10_1101-2020_12_31_424971 111 40 s–1 s–1 NNS 10_1101-2020_12_31_424971 111 41 ) ) -RRB- 10_1101-2020_12_31_424971 111 42 rather rather RB 10_1101-2020_12_31_424971 111 43 than than IN 10_1101-2020_12_31_424971 111 44 that that DT 10_1101-2020_12_31_424971 111 45 of of IN 10_1101-2020_12_31_424971 111 46 +1 +1 NNS 10_1101-2020_12_31_424971 111 47 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 111 48 after after IN 10_1101-2020_12_31_424971 111 49 285 285 CD 10_1101-2020_12_31_424971 111 50 translocation translocation NN 10_1101-2020_12_31_424971 111 51 into into IN 10_1101-2020_12_31_424971 111 52 the the DT 10_1101-2020_12_31_424971 111 53 P p NN 10_1101-2020_12_31_424971 111 54 site site NN 10_1101-2020_12_31_424971 111 55 ( ( -LRB- 10_1101-2020_12_31_424971 111 56 ~10–3 ~10–3 NFP 10_1101-2020_12_31_424971 111 57 s–1)29 s–1)29 NN 10_1101-2020_12_31_424971 111 58 , , , 10_1101-2020_12_31_424971 111 59 indicating indicate VBG 10_1101-2020_12_31_424971 111 60 that that IN 10_1101-2020_12_31_424971 111 61 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 111 62 underwent undergo VBD 10_1101-2020_12_31_424971 111 63 +1 +1 NNS 10_1101-2020_12_31_424971 111 64 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 111 65 during during IN 10_1101-2020_12_31_424971 111 66 286 286 CD 10_1101-2020_12_31_424971 111 67 translocation translocation NN 10_1101-2020_12_31_424971 111 68 . . . 10_1101-2020_12_31_424971 112 1 Our -PRON- PRP$ 10_1101-2020_12_31_424971 112 2 observation observation NN 10_1101-2020_12_31_424971 112 3 that that IN 10_1101-2020_12_31_424971 112 4 the the DT 10_1101-2020_12_31_424971 112 5 fMPV fMPV NNP 10_1101-2020_12_31_424971 112 6 yield yield NN 10_1101-2020_12_31_424971 112 7 plateaus plateaus NN 10_1101-2020_12_31_424971 112 8 at at IN 10_1101-2020_12_31_424971 112 9 90 90 CD 10_1101-2020_12_31_424971 112 10 % % NN 10_1101-2020_12_31_424971 112 11 at at IN 10_1101-2020_12_31_424971 112 12 long long JJ 10_1101-2020_12_31_424971 112 13 reaction reaction NN 10_1101-2020_12_31_424971 112 14 times times NNP 10_1101-2020_12_31_424971 112 15 287 287 CD 10_1101-2020_12_31_424971 112 16 suggests suggest VBZ 10_1101-2020_12_31_424971 112 17 that that IN 10_1101-2020_12_31_424971 112 18 the the DT 10_1101-2020_12_31_424971 112 19 sub sub NN 10_1101-2020_12_31_424971 112 20 - - NNS 10_1101-2020_12_31_424971 112 21 populations population NNS 10_1101-2020_12_31_424971 112 22 of of IN 10_1101-2020_12_31_424971 112 23 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 112 24 that that WDT 10_1101-2020_12_31_424971 112 25 will will MD 10_1101-2020_12_31_424971 112 26 shift shift VB 10_1101-2020_12_31_424971 112 27 to to IN 10_1101-2020_12_31_424971 112 28 the the DT 10_1101-2020_12_31_424971 112 29 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 112 30 and and CC 10_1101-2020_12_31_424971 112 31 remain remain VBP 10_1101-2020_12_31_424971 112 32 in in IN 10_1101-2020_12_31_424971 112 33 the the DT 10_1101-2020_12_31_424971 112 34 0 0 CD 10_1101-2020_12_31_424971 112 35 - - SYM 10_1101-2020_12_31_424971 112 36 288 288 CD 10_1101-2020_12_31_424971 112 37 frame frame NN 10_1101-2020_12_31_424971 112 38 are be VBP 10_1101-2020_12_31_424971 112 39 likely likely RB 10_1101-2020_12_31_424971 112 40 established establish VBN 10_1101-2020_12_31_424971 112 41 in in IN 10_1101-2020_12_31_424971 112 42 the the DT 10_1101-2020_12_31_424971 112 43 A A NNP 10_1101-2020_12_31_424971 112 44 site site NN 10_1101-2020_12_31_424971 112 45 , , , 10_1101-2020_12_31_424971 112 46 even even RB 10_1101-2020_12_31_424971 112 47 before before IN 10_1101-2020_12_31_424971 112 48 EF EF NNP 10_1101-2020_12_31_424971 112 49 - - HYPH 10_1101-2020_12_31_424971 112 50 G G NNP 10_1101-2020_12_31_424971 112 51 binds bind NNS 10_1101-2020_12_31_424971 112 52 to to IN 10_1101-2020_12_31_424971 112 53 the the DT 10_1101-2020_12_31_424971 112 54 PRE PRE NNP 10_1101-2020_12_31_424971 112 55 complex complex NN 10_1101-2020_12_31_424971 112 56 . . . 10_1101-2020_12_31_424971 113 1 Given give VBN 10_1101-2020_12_31_424971 113 2 that that IN 10_1101-2020_12_31_424971 113 3 289 289 CD 10_1101-2020_12_31_424971 113 4 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 113 5 exhibits exhibit NNS 10_1101-2020_12_31_424971 113 6 triplet triplet NN 10_1101-2020_12_31_424971 113 7 pairing pair VBG 10_1101-2020_12_31_424971 113 8 in in IN 10_1101-2020_12_31_424971 113 9 the the DT 10_1101-2020_12_31_424971 113 10 0-frame 0-frame CD 10_1101-2020_12_31_424971 113 11 at at IN 10_1101-2020_12_31_424971 113 12 the the DT 10_1101-2020_12_31_424971 113 13 A a NN 10_1101-2020_12_31_424971 113 14 site site NN 10_1101-2020_12_31_424971 113 15 ( ( -LRB- 10_1101-2020_12_31_424971 113 16 Figures Figures NNP 10_1101-2020_12_31_424971 113 17 3a 3a NNP 10_1101-2020_12_31_424971 113 18 - - HYPH 10_1101-2020_12_31_424971 113 19 c c NNP 10_1101-2020_12_31_424971 113 20 , , , 10_1101-2020_12_31_424971 113 21 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 113 22 Table Table NNP 10_1101-2020_12_31_424971 113 23 2 2 CD 10_1101-2020_12_31_424971 113 24 , , , 10_1101-2020_12_31_424971 113 25 290 290 CD 10_1101-2020_12_31_424971 113 26 and and CC 10_1101-2020_12_31_424971 113 27 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 113 28 Figure Figure NNP 10_1101-2020_12_31_424971 113 29 2a 2a NNP 10_1101-2020_12_31_424971 113 30 ) ) -RRB- 10_1101-2020_12_31_424971 113 31 and and CC 10_1101-2020_12_31_424971 113 32 shifts shift NNS 10_1101-2020_12_31_424971 113 33 into into IN 10_1101-2020_12_31_424971 113 34 the the DT 10_1101-2020_12_31_424971 113 35 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 113 36 during during IN 10_1101-2020_12_31_424971 113 37 translocation translocation NN 10_1101-2020_12_31_424971 113 38 ( ( -LRB- 10_1101-2020_12_31_424971 113 39 Figure Figure NNP 10_1101-2020_12_31_424971 113 40 5a 5a NNP 10_1101-2020_12_31_424971 113 41 ) ) -RRB- 10_1101-2020_12_31_424971 113 42 , , , 10_1101-2020_12_31_424971 113 43 the the DT 10_1101-2020_12_31_424971 113 44 291 291 CD 10_1101-2020_12_31_424971 113 45 two two CD 10_1101-2020_12_31_424971 113 46 sub sub NN 10_1101-2020_12_31_424971 113 47 - - NNS 10_1101-2020_12_31_424971 113 48 populations population NNS 10_1101-2020_12_31_424971 113 49 of of IN 10_1101-2020_12_31_424971 113 50 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 113 51 in in IN 10_1101-2020_12_31_424971 113 52 the the DT 10_1101-2020_12_31_424971 113 53 A A NNP 10_1101-2020_12_31_424971 113 54 site site NN 10_1101-2020_12_31_424971 113 55 seem seem VBP 10_1101-2020_12_31_424971 113 56 to to TO 10_1101-2020_12_31_424971 113 57 differ differ VB 10_1101-2020_12_31_424971 113 58 primarily primarily RB 10_1101-2020_12_31_424971 113 59 in in IN 10_1101-2020_12_31_424971 113 60 their -PRON- PRP$ 10_1101-2020_12_31_424971 113 61 propensity propensity NN 10_1101-2020_12_31_424971 113 62 to to TO 10_1101-2020_12_31_424971 113 63 undergo undergo VB 10_1101-2020_12_31_424971 113 64 292 292 CD 10_1101-2020_12_31_424971 113 65 +1 +1 NNS 10_1101-2020_12_31_424971 113 66 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 113 67 during during IN 10_1101-2020_12_31_424971 113 68 translocation translocation NN 10_1101-2020_12_31_424971 113 69 . . . 10_1101-2020_12_31_424971 114 1 The the DT 10_1101-2020_12_31_424971 114 2 sub sub JJ 10_1101-2020_12_31_424971 114 3 - - NN 10_1101-2020_12_31_424971 114 4 population population NN 10_1101-2020_12_31_424971 114 5 that that WDT 10_1101-2020_12_31_424971 114 6 encompasses encompass VBZ 10_1101-2020_12_31_424971 114 7 90 90 CD 10_1101-2020_12_31_424971 114 8 % % NN 10_1101-2020_12_31_424971 114 9 of of IN 10_1101-2020_12_31_424971 114 10 the the DT 10_1101-2020_12_31_424971 114 11 total total JJ 10_1101-2020_12_31_424971 114 12 293 293 CD 10_1101-2020_12_31_424971 114 13 would would MD 10_1101-2020_12_31_424971 114 14 exhibit exhibit VB 10_1101-2020_12_31_424971 114 15 a a DT 10_1101-2020_12_31_424971 114 16 high high JJ 10_1101-2020_12_31_424971 114 17 propensity propensity NN 10_1101-2020_12_31_424971 114 18 of of IN 10_1101-2020_12_31_424971 114 19 undergoing undergoing JJ 10_1101-2020_12_31_424971 114 20 +1 +1 NNS 10_1101-2020_12_31_424971 114 21 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 114 22 during during IN 10_1101-2020_12_31_424971 114 23 translocation translocation NN 10_1101-2020_12_31_424971 114 24 , , , 10_1101-2020_12_31_424971 114 25 whereas whereas IN 10_1101-2020_12_31_424971 114 26 the the DT 10_1101-2020_12_31_424971 114 27 294 294 CD 10_1101-2020_12_31_424971 114 28 sub sub NN 10_1101-2020_12_31_424971 114 29 - - NN 10_1101-2020_12_31_424971 114 30 population population NN 10_1101-2020_12_31_424971 114 31 that that WDT 10_1101-2020_12_31_424971 114 32 encompasses encompass VBZ 10_1101-2020_12_31_424971 114 33 10 10 CD 10_1101-2020_12_31_424971 114 34 % % NN 10_1101-2020_12_31_424971 114 35 of of IN 10_1101-2020_12_31_424971 114 36 the the DT 10_1101-2020_12_31_424971 114 37 total total NN 10_1101-2020_12_31_424971 114 38 would would MD 10_1101-2020_12_31_424971 114 39 exhibit exhibit VB 10_1101-2020_12_31_424971 114 40 a a DT 10_1101-2020_12_31_424971 114 41 low low JJ 10_1101-2020_12_31_424971 114 42 propensity propensity NN 10_1101-2020_12_31_424971 114 43 of of IN 10_1101-2020_12_31_424971 114 44 undergoing undergo VBG 10_1101-2020_12_31_424971 114 45 295 295 CD 10_1101-2020_12_31_424971 114 46 +1 +1 NNS 10_1101-2020_12_31_424971 114 47 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 114 48 during during IN 10_1101-2020_12_31_424971 114 49 translocation translocation NN 10_1101-2020_12_31_424971 114 50 , , , 10_1101-2020_12_31_424971 114 51 preferring prefer VBG 10_1101-2020_12_31_424971 114 52 instead instead RB 10_1101-2020_12_31_424971 114 53 to to TO 10_1101-2020_12_31_424971 114 54 remain remain VB 10_1101-2020_12_31_424971 114 55 in in IN 10_1101-2020_12_31_424971 114 56 the the DT 10_1101-2020_12_31_424971 114 57 0-frame 0-frame CD 10_1101-2020_12_31_424971 114 58 . . . 10_1101-2020_12_31_424971 115 1 296 296 CD 10_1101-2020_12_31_424971 115 2 We -PRON- PRP 10_1101-2020_12_31_424971 115 3 next next RB 10_1101-2020_12_31_424971 115 4 determined determine VBD 10_1101-2020_12_31_424971 115 5 whether whether IN 10_1101-2020_12_31_424971 115 6 the the DT 10_1101-2020_12_31_424971 115 7 10 10 CD 10_1101-2020_12_31_424971 115 8 % % NN 10_1101-2020_12_31_424971 115 9 sub sub NN 10_1101-2020_12_31_424971 115 10 - - NN 10_1101-2020_12_31_424971 115 11 population population NN 10_1101-2020_12_31_424971 115 12 of of IN 10_1101-2020_12_31_424971 115 13 G37-state G37-state NNP 10_1101-2020_12_31_424971 115 14 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 115 15 that that WDT 10_1101-2020_12_31_424971 115 16 remained remain VBD 10_1101-2020_12_31_424971 115 17 in in IN 10_1101-2020_12_31_424971 115 18 the the DT 10_1101-2020_12_31_424971 115 19 297 297 CD 10_1101-2020_12_31_424971 115 20 0-frame 0-frame CD 10_1101-2020_12_31_424971 115 21 during during IN 10_1101-2020_12_31_424971 115 22 translocation translocation NN 10_1101-2020_12_31_424971 115 23 could could MD 10_1101-2020_12_31_424971 115 24 undergo undergo VB 10_1101-2020_12_31_424971 115 25 +1 +1 NNS 10_1101-2020_12_31_424971 115 26 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 115 27 after after IN 10_1101-2020_12_31_424971 115 28 arrival arrival NN 10_1101-2020_12_31_424971 115 29 at at IN 10_1101-2020_12_31_424971 115 30 the the DT 10_1101-2020_12_31_424971 115 31 P p NN 10_1101-2020_12_31_424971 115 32 site site NN 10_1101-2020_12_31_424971 115 33 . . . 10_1101-2020_12_31_424971 116 1 We -PRON- PRP 10_1101-2020_12_31_424971 116 2 varied vary VBD 10_1101-2020_12_31_424971 116 3 298 298 CD 10_1101-2020_12_31_424971 116 4 our -PRON- PRP$ 10_1101-2020_12_31_424971 116 5 tripeptide tripeptide NN 10_1101-2020_12_31_424971 116 6 formation formation NN 10_1101-2020_12_31_424971 116 7 assay assay NN 10_1101-2020_12_31_424971 116 8 so so IN 10_1101-2020_12_31_424971 116 9 as as IN 10_1101-2020_12_31_424971 116 10 to to TO 10_1101-2020_12_31_424971 116 11 deliver deliver VB 10_1101-2020_12_31_424971 116 12 the the DT 10_1101-2020_12_31_424971 116 13 TCs tc NNS 10_1101-2020_12_31_424971 116 14 in in IN 10_1101-2020_12_31_424971 116 15 two two CD 10_1101-2020_12_31_424971 116 16 steps step NNS 10_1101-2020_12_31_424971 116 17 separated separate VBN 10_1101-2020_12_31_424971 116 18 by by IN 10_1101-2020_12_31_424971 116 19 a a DT 10_1101-2020_12_31_424971 116 20 defined define VBN 10_1101-2020_12_31_424971 116 21 time time NN 10_1101-2020_12_31_424971 116 22 299 299 CD 10_1101-2020_12_31_424971 116 23 interval interval NN 10_1101-2020_12_31_424971 116 24 ( ( -LRB- 10_1101-2020_12_31_424971 116 25 Figure Figure NNP 10_1101-2020_12_31_424971 116 26 5b 5b NN 10_1101-2020_12_31_424971 116 27 ) ) -RRB- 10_1101-2020_12_31_424971 116 28 . . . 10_1101-2020_12_31_424971 117 1 In in IN 10_1101-2020_12_31_424971 117 2 the the DT 10_1101-2020_12_31_424971 117 3 first first JJ 10_1101-2020_12_31_424971 117 4 step step NN 10_1101-2020_12_31_424971 117 5 , , , 10_1101-2020_12_31_424971 117 6 G37-state G37-state NNP 10_1101-2020_12_31_424971 117 7 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 117 8 and and CC 10_1101-2020_12_31_424971 117 9 EF EF NNP 10_1101-2020_12_31_424971 117 10 - - HYPH 10_1101-2020_12_31_424971 117 11 G G NNP 10_1101-2020_12_31_424971 117 12 were be VBD 10_1101-2020_12_31_424971 117 13 delivered deliver VBN 10_1101-2020_12_31_424971 117 14 to to IN 10_1101-2020_12_31_424971 117 15 the the DT 10_1101-2020_12_31_424971 117 16 70S 70S NNP 10_1101-2020_12_31_424971 117 17 IC IC NNP 10_1101-2020_12_31_424971 117 18 300 300 CD 10_1101-2020_12_31_424971 117 19 to to TO 10_1101-2020_12_31_424971 117 20 form form VB 10_1101-2020_12_31_424971 117 21 a a DT 10_1101-2020_12_31_424971 117 22 POST POST NNP 10_1101-2020_12_31_424971 117 23 complex complex NN 10_1101-2020_12_31_424971 117 24 , , , 10_1101-2020_12_31_424971 117 25 which which WDT 10_1101-2020_12_31_424971 117 26 was be VBD 10_1101-2020_12_31_424971 117 27 then then RB 10_1101-2020_12_31_424971 117 28 allowed allow VBN 10_1101-2020_12_31_424971 117 29 the the DT 10_1101-2020_12_31_424971 117 30 opportunity opportunity NN 10_1101-2020_12_31_424971 117 31 to to TO 10_1101-2020_12_31_424971 117 32 shift shift VB 10_1101-2020_12_31_424971 117 33 to to IN 10_1101-2020_12_31_424971 117 34 the the DT 10_1101-2020_12_31_424971 117 35 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 117 36 over over IN 10_1101-2020_12_31_424971 117 37 a a DT 10_1101-2020_12_31_424971 117 38 301 301 CD 10_1101-2020_12_31_424971 117 39 systematically systematically RB 10_1101-2020_12_31_424971 117 40 increasing increasing JJ 10_1101-2020_12_31_424971 117 41 time time NN 10_1101-2020_12_31_424971 117 42 interval interval NN 10_1101-2020_12_31_424971 117 43 . . . 10_1101-2020_12_31_424971 118 1 In in IN 10_1101-2020_12_31_424971 118 2 the the DT 10_1101-2020_12_31_424971 118 3 second second JJ 10_1101-2020_12_31_424971 118 4 step step NN 10_1101-2020_12_31_424971 118 5 , , , 10_1101-2020_12_31_424971 118 6 an an DT 10_1101-2020_12_31_424971 118 7 equimolar equimolar JJ 10_1101-2020_12_31_424971 118 8 mixture mixture NN 10_1101-2020_12_31_424971 118 9 of of IN 10_1101-2020_12_31_424971 118 10 tRNAArg- trnaarg- NN 10_1101-2020_12_31_424971 118 11 and and CC 10_1101-2020_12_31_424971 118 12 302 302 CD 10_1101-2020_12_31_424971 118 13 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 118 14 - - HYPH 10_1101-2020_12_31_424971 118 15 TCs TCs NNP 10_1101-2020_12_31_424971 118 16 was be VBD 10_1101-2020_12_31_424971 118 17 delivered deliver VBN 10_1101-2020_12_31_424971 118 18 to to IN 10_1101-2020_12_31_424971 118 19 the the DT 10_1101-2020_12_31_424971 118 20 POST POST NNP 10_1101-2020_12_31_424971 118 21 complex complex NN 10_1101-2020_12_31_424971 118 22 . . . 10_1101-2020_12_31_424971 119 1 The the DT 10_1101-2020_12_31_424971 119 2 results result NNS 10_1101-2020_12_31_424971 119 3 showed show VBD 10_1101-2020_12_31_424971 119 4 that that IN 10_1101-2020_12_31_424971 119 5 fMPV fmpv NN 10_1101-2020_12_31_424971 119 6 was be VBD 10_1101-2020_12_31_424971 119 7 rapidly rapidly RB 10_1101-2020_12_31_424971 119 8 303 303 CD 10_1101-2020_12_31_424971 119 9 formed form VBN 10_1101-2020_12_31_424971 119 10 at at IN 10_1101-2020_12_31_424971 119 11 a a DT 10_1101-2020_12_31_424971 119 12 high high JJ 10_1101-2020_12_31_424971 119 13 yield yield NN 10_1101-2020_12_31_424971 119 14 and and CC 10_1101-2020_12_31_424971 119 15 exhibited exhibit VBD 10_1101-2020_12_31_424971 119 16 a a DT 10_1101-2020_12_31_424971 119 17 kfMP+V kfMP+V NNP 10_1101-2020_12_31_424971 119 18 , , , 10_1101-2020_12_31_424971 119 19 obs obs NNP 10_1101-2020_12_31_424971 119 20 ( ( -LRB- 10_1101-2020_12_31_424971 119 21 where where WRB 10_1101-2020_12_31_424971 119 22 the the DT 10_1101-2020_12_31_424971 119 23 “ " `` 10_1101-2020_12_31_424971 119 24 + + NNS 10_1101-2020_12_31_424971 119 25 ” " '' 10_1101-2020_12_31_424971 119 26 denotes denote VBZ 10_1101-2020_12_31_424971 119 27 the the DT 10_1101-2020_12_31_424971 119 28 time time NN 10_1101-2020_12_31_424971 119 29 interval interval NN 10_1101-2020_12_31_424971 119 30 between between IN 10_1101-2020_12_31_424971 119 31 304 304 CD 10_1101-2020_12_31_424971 119 32 the the DT 10_1101-2020_12_31_424971 119 33 delivery delivery NN 10_1101-2020_12_31_424971 119 34 of of IN 10_1101-2020_12_31_424971 119 35 translation translation NN 10_1101-2020_12_31_424971 119 36 components component NNS 10_1101-2020_12_31_424971 119 37 ) ) -RRB- 10_1101-2020_12_31_424971 119 38 that that WDT 10_1101-2020_12_31_424971 119 39 did do VBD 10_1101-2020_12_31_424971 119 40 not not RB 10_1101-2020_12_31_424971 119 41 increase increase VB 10_1101-2020_12_31_424971 119 42 as as IN 10_1101-2020_12_31_424971 119 43 a a DT 10_1101-2020_12_31_424971 119 44 function function NN 10_1101-2020_12_31_424971 119 45 of of IN 10_1101-2020_12_31_424971 119 46 time time NN 10_1101-2020_12_31_424971 119 47 . . . 10_1101-2020_12_31_424971 120 1 In in IN 10_1101-2020_12_31_424971 120 2 contrast contrast NN 10_1101-2020_12_31_424971 120 3 , , , 10_1101-2020_12_31_424971 120 4 305 305 CD 10_1101-2020_12_31_424971 120 5 fMPR fmpr NN 10_1101-2020_12_31_424971 120 6 was be VBD 10_1101-2020_12_31_424971 120 7 formed form VBN 10_1101-2020_12_31_424971 120 8 at at IN 10_1101-2020_12_31_424971 120 9 a a DT 10_1101-2020_12_31_424971 120 10 low low JJ 10_1101-2020_12_31_424971 120 11 yield yield NN 10_1101-2020_12_31_424971 120 12 and and CC 10_1101-2020_12_31_424971 120 13 exhibited exhibit VBD 10_1101-2020_12_31_424971 120 14 a a DT 10_1101-2020_12_31_424971 120 15 kfMP+R kfmp+r NN 10_1101-2020_12_31_424971 120 16 , , , 10_1101-2020_12_31_424971 120 17 obs obs NN 10_1101-2020_12_31_424971 120 18 that that WDT 10_1101-2020_12_31_424971 120 19 did do VBD 10_1101-2020_12_31_424971 120 20 not not RB 10_1101-2020_12_31_424971 120 21 decrease decrease VB 10_1101-2020_12_31_424971 120 22 as as IN 10_1101-2020_12_31_424971 120 23 a a DT 10_1101-2020_12_31_424971 120 24 function function NN 10_1101-2020_12_31_424971 120 25 of of IN 10_1101-2020_12_31_424971 120 26 306 306 CD 10_1101-2020_12_31_424971 120 27 .CC .CC , 10_1101-2020_12_31_424971 120 28 - - HYPH 10_1101-2020_12_31_424971 120 29 BY by IN 10_1101-2020_12_31_424971 120 30 - - HYPH 10_1101-2020_12_31_424971 120 31 NC NC NNP 10_1101-2020_12_31_424971 120 32 - - HYPH 10_1101-2020_12_31_424971 120 33 ND ND NNP 10_1101-2020_12_31_424971 120 34 4.0 4.0 CD 10_1101-2020_12_31_424971 120 35 International International NNP 10_1101-2020_12_31_424971 120 36 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 120 37 under under IN 10_1101-2020_12_31_424971 120 38 a a DT 10_1101-2020_12_31_424971 120 39 ( ( -LRB- 10_1101-2020_12_31_424971 120 40 which which WDT 10_1101-2020_12_31_424971 120 41 was be VBD 10_1101-2020_12_31_424971 120 42 not not RB 10_1101-2020_12_31_424971 120 43 certified certify VBN 10_1101-2020_12_31_424971 120 44 by by IN 10_1101-2020_12_31_424971 120 45 peer peer NN 10_1101-2020_12_31_424971 120 46 review review NN 10_1101-2020_12_31_424971 120 47 ) ) -RRB- 10_1101-2020_12_31_424971 120 48 is be VBZ 10_1101-2020_12_31_424971 120 49 the the DT 10_1101-2020_12_31_424971 120 50 author author NN 10_1101-2020_12_31_424971 120 51 / / SYM 10_1101-2020_12_31_424971 120 52 funder funder NN 10_1101-2020_12_31_424971 120 53 , , , 10_1101-2020_12_31_424971 120 54 who who WP 10_1101-2020_12_31_424971 120 55 has have VBZ 10_1101-2020_12_31_424971 120 56 granted grant VBN 10_1101-2020_12_31_424971 120 57 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 120 58 a a DT 10_1101-2020_12_31_424971 120 59 license license NN 10_1101-2020_12_31_424971 120 60 to to TO 10_1101-2020_12_31_424971 120 61 display display VB 10_1101-2020_12_31_424971 120 62 the the DT 10_1101-2020_12_31_424971 120 63 preprint preprint NN 10_1101-2020_12_31_424971 120 64 in in IN 10_1101-2020_12_31_424971 120 65 perpetuity perpetuity NN 10_1101-2020_12_31_424971 120 66 . . . 10_1101-2020_12_31_424971 121 1 It -PRON- PRP 10_1101-2020_12_31_424971 121 2 is be VBZ 10_1101-2020_12_31_424971 121 3 made make VBN 10_1101-2020_12_31_424971 121 4 The the DT 10_1101-2020_12_31_424971 121 5 copyright copyright NN 10_1101-2020_12_31_424971 121 6 holder holder NN 10_1101-2020_12_31_424971 121 7 for for IN 10_1101-2020_12_31_424971 121 8 this this DT 10_1101-2020_12_31_424971 121 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 121 10 version version NN 10_1101-2020_12_31_424971 121 11 posted post VBD 10_1101-2020_12_31_424971 121 12 January January NNP 10_1101-2020_12_31_424971 121 13 2 2 CD 10_1101-2020_12_31_424971 121 14 , , , 10_1101-2020_12_31_424971 121 15 2021 2021 CD 10_1101-2020_12_31_424971 121 16 . . . 10_1101-2020_12_31_424971 121 17 ; ; : 10_1101-2020_12_31_424971 121 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 121 19 : : : 10_1101-2020_12_31_424971 121 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 121 21 preprint preprint NN 10_1101-2020_12_31_424971 121 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 121 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 121 24 13 13 CD 10_1101-2020_12_31_424971 121 25 time time NN 10_1101-2020_12_31_424971 121 26 . . . 10_1101-2020_12_31_424971 122 1 Together together RB 10_1101-2020_12_31_424971 122 2 , , , 10_1101-2020_12_31_424971 122 3 these these DT 10_1101-2020_12_31_424971 122 4 results result NNS 10_1101-2020_12_31_424971 122 5 indicate indicate VBP 10_1101-2020_12_31_424971 122 6 that that IN 10_1101-2020_12_31_424971 122 7 the the DT 10_1101-2020_12_31_424971 122 8 sub sub NN 10_1101-2020_12_31_424971 122 9 - - NN 10_1101-2020_12_31_424971 122 10 population population NN 10_1101-2020_12_31_424971 122 11 of of IN 10_1101-2020_12_31_424971 122 12 P p NN 10_1101-2020_12_31_424971 122 13 site site NN 10_1101-2020_12_31_424971 122 14 - - HYPH 10_1101-2020_12_31_424971 122 15 bound bind VBN 10_1101-2020_12_31_424971 122 16 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 122 17 in in IN 10_1101-2020_12_31_424971 122 18 the the DT 10_1101-2020_12_31_424971 122 19 0-frame 0-frame CD 10_1101-2020_12_31_424971 122 20 307 307 CD 10_1101-2020_12_31_424971 122 21 does do VBZ 10_1101-2020_12_31_424971 122 22 not not RB 10_1101-2020_12_31_424971 122 23 undergo undergo VB 10_1101-2020_12_31_424971 122 24 +1 +1 NNS 10_1101-2020_12_31_424971 122 25 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 122 26 . . . 10_1101-2020_12_31_424971 123 1 This this DT 10_1101-2020_12_31_424971 123 2 interpretation interpretation NN 10_1101-2020_12_31_424971 123 3 is be VBZ 10_1101-2020_12_31_424971 123 4 supported support VBN 10_1101-2020_12_31_424971 123 5 by by IN 10_1101-2020_12_31_424971 123 6 the the DT 10_1101-2020_12_31_424971 123 7 observation observation NN 10_1101-2020_12_31_424971 123 8 that that IN 10_1101-2020_12_31_424971 123 9 EF EF NNP 10_1101-2020_12_31_424971 123 10 - - HYPH 10_1101-2020_12_31_424971 123 11 P P NNP 10_1101-2020_12_31_424971 123 12 , , , 10_1101-2020_12_31_424971 123 13 308 308 CD 10_1101-2020_12_31_424971 123 14 an an DT 10_1101-2020_12_31_424971 123 15 elongation elongation NN 10_1101-2020_12_31_424971 123 16 factor factor NN 10_1101-2020_12_31_424971 123 17 which which WDT 10_1101-2020_12_31_424971 123 18 we -PRON- PRP 10_1101-2020_12_31_424971 123 19 showed show VBD 10_1101-2020_12_31_424971 123 20 suppresses suppresse NNS 10_1101-2020_12_31_424971 123 21 +1 +1 : 10_1101-2020_12_31_424971 123 22 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 123 23 within within IN 10_1101-2020_12_31_424971 123 24 the the DT 10_1101-2020_12_31_424971 123 25 P p NN 10_1101-2020_12_31_424971 123 26 site29 site29 NN 10_1101-2020_12_31_424971 123 27 , , , 10_1101-2020_12_31_424971 123 28 had have VBD 10_1101-2020_12_31_424971 123 29 no no DT 10_1101-2020_12_31_424971 123 30 309 309 CD 10_1101-2020_12_31_424971 123 31 effect effect NN 10_1101-2020_12_31_424971 123 32 on on IN 10_1101-2020_12_31_424971 123 33 the the DT 10_1101-2020_12_31_424971 123 34 yield yield NN 10_1101-2020_12_31_424971 123 35 of of IN 10_1101-2020_12_31_424971 123 36 fMPV fmpv JJ 10_1101-2020_12_31_424971 123 37 yield yield NN 10_1101-2020_12_31_424971 123 38 ( ( -LRB- 10_1101-2020_12_31_424971 123 39 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 123 40 Figure Figure NNP 10_1101-2020_12_31_424971 123 41 2c 2c NN 10_1101-2020_12_31_424971 123 42 and and CC 10_1101-2020_12_31_424971 123 43 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 123 44 Table Table NNP 10_1101-2020_12_31_424971 123 45 3 3 CD 10_1101-2020_12_31_424971 123 46 ) ) -RRB- 10_1101-2020_12_31_424971 123 47 . . . 10_1101-2020_12_31_424971 124 1 310 310 CD 10_1101-2020_12_31_424971 124 2 Having have VBG 10_1101-2020_12_31_424971 124 3 shown show VBN 10_1101-2020_12_31_424971 124 4 that that IN 10_1101-2020_12_31_424971 124 5 +1 +1 NNS 10_1101-2020_12_31_424971 124 6 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 124 7 of of IN 10_1101-2020_12_31_424971 124 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 124 9 occurs occur VBZ 10_1101-2020_12_31_424971 124 10 only only RB 10_1101-2020_12_31_424971 124 11 during during IN 10_1101-2020_12_31_424971 124 12 translocation translocation NN 10_1101-2020_12_31_424971 124 13 , , , 10_1101-2020_12_31_424971 124 14 we -PRON- PRP 10_1101-2020_12_31_424971 124 15 evaluated evaluate VBD 10_1101-2020_12_31_424971 124 16 311 311 CD 10_1101-2020_12_31_424971 124 17 the the DT 10_1101-2020_12_31_424971 124 18 effect effect NN 10_1101-2020_12_31_424971 124 19 of of IN 10_1101-2020_12_31_424971 124 20 m1G37 m1g37 NN 10_1101-2020_12_31_424971 124 21 on on IN 10_1101-2020_12_31_424971 124 22 the the DT 10_1101-2020_12_31_424971 124 23 frequency frequency NN 10_1101-2020_12_31_424971 124 24 of of IN 10_1101-2020_12_31_424971 124 25 this this DT 10_1101-2020_12_31_424971 124 26 event event NN 10_1101-2020_12_31_424971 124 27 . . . 10_1101-2020_12_31_424971 125 1 We -PRON- PRP 10_1101-2020_12_31_424971 125 2 began begin VBD 10_1101-2020_12_31_424971 125 3 by by IN 10_1101-2020_12_31_424971 125 4 delivering deliver VBG 10_1101-2020_12_31_424971 125 5 G37- G37- NNP 10_1101-2020_12_31_424971 125 6 , , , 10_1101-2020_12_31_424971 125 7 m1G37- m1g37- CD 10_1101-2020_12_31_424971 125 8 , , , 10_1101-2020_12_31_424971 125 9 or or CC 10_1101-2020_12_31_424971 125 10 312 312 CD 10_1101-2020_12_31_424971 125 11 native native JJ 10_1101-2020_12_31_424971 125 12 - - HYPH 10_1101-2020_12_31_424971 125 13 state state NN 10_1101-2020_12_31_424971 125 14 SufB2-TCs sufb2-tc NNS 10_1101-2020_12_31_424971 125 15 together together RB 10_1101-2020_12_31_424971 125 16 with with IN 10_1101-2020_12_31_424971 125 17 EF EF NNP 10_1101-2020_12_31_424971 125 18 - - HYPH 10_1101-2020_12_31_424971 125 19 G G NNP 10_1101-2020_12_31_424971 125 20 to to IN 10_1101-2020_12_31_424971 125 21 70S 70S NNP 10_1101-2020_12_31_424971 125 22 ICs ic NNS 10_1101-2020_12_31_424971 125 23 to to TO 10_1101-2020_12_31_424971 125 24 form form VB 10_1101-2020_12_31_424971 125 25 the the DT 10_1101-2020_12_31_424971 125 26 corresponding corresponding JJ 10_1101-2020_12_31_424971 125 27 POST POST NNP 10_1101-2020_12_31_424971 125 28 313 313 CD 10_1101-2020_12_31_424971 125 29 complexes complex NNS 10_1101-2020_12_31_424971 125 30 and and CC 10_1101-2020_12_31_424971 125 31 then then RB 10_1101-2020_12_31_424971 125 32 delivered deliver VBD 10_1101-2020_12_31_424971 125 33 an an DT 10_1101-2020_12_31_424971 125 34 equimolar equimolar JJ 10_1101-2020_12_31_424971 125 35 mixture mixture NN 10_1101-2020_12_31_424971 125 36 of of IN 10_1101-2020_12_31_424971 125 37 tRNAArg- tRNAArg- NNP 10_1101-2020_12_31_424971 125 38 and and CC 10_1101-2020_12_31_424971 125 39 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 125 40 - - HYPH 10_1101-2020_12_31_424971 125 41 TCs TCs NNP 10_1101-2020_12_31_424971 125 42 to to IN 10_1101-2020_12_31_424971 125 43 each each DT 10_1101-2020_12_31_424971 125 44 POST POST NNP 10_1101-2020_12_31_424971 125 45 314 314 CD 10_1101-2020_12_31_424971 125 46 complex complex NN 10_1101-2020_12_31_424971 125 47 to to TO 10_1101-2020_12_31_424971 125 48 determine determine VB 10_1101-2020_12_31_424971 125 49 the the DT 10_1101-2020_12_31_424971 125 50 relative relative JJ 10_1101-2020_12_31_424971 125 51 formation formation NN 10_1101-2020_12_31_424971 125 52 of of IN 10_1101-2020_12_31_424971 125 53 fMPV fmpv NN 10_1101-2020_12_31_424971 125 54 and and CC 10_1101-2020_12_31_424971 125 55 fMPR fmpr NN 10_1101-2020_12_31_424971 125 56 . . . 10_1101-2020_12_31_424971 126 1 The the DT 10_1101-2020_12_31_424971 126 2 results result NNS 10_1101-2020_12_31_424971 126 3 showed show VBD 10_1101-2020_12_31_424971 126 4 that that IN 10_1101-2020_12_31_424971 126 5 m1G37- m1g37- CD 10_1101-2020_12_31_424971 126 6 315 315 CD 10_1101-2020_12_31_424971 126 7 and and CC 10_1101-2020_12_31_424971 126 8 native native JJ 10_1101-2020_12_31_424971 126 9 - - HYPH 10_1101-2020_12_31_424971 126 10 state state NN 10_1101-2020_12_31_424971 126 11 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 126 12 displayed display VBD 10_1101-2020_12_31_424971 126 13 a a DT 10_1101-2020_12_31_424971 126 14 reduced reduce VBN 10_1101-2020_12_31_424971 126 15 fMPV fmpv NN 10_1101-2020_12_31_424971 126 16 yield yield NN 10_1101-2020_12_31_424971 126 17 and and CC 10_1101-2020_12_31_424971 126 18 a a DT 10_1101-2020_12_31_424971 126 19 concomitantly concomitantly RB 10_1101-2020_12_31_424971 126 20 increased increase VBN 10_1101-2020_12_31_424971 126 21 fMPR fmpr CD 10_1101-2020_12_31_424971 126 22 yield yield NN 10_1101-2020_12_31_424971 126 23 316 316 CD 10_1101-2020_12_31_424971 126 24 relative relative JJ 10_1101-2020_12_31_424971 126 25 to to IN 10_1101-2020_12_31_424971 126 26 G37-state G37-state NNP 10_1101-2020_12_31_424971 126 27 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 126 28 ( ( -LRB- 10_1101-2020_12_31_424971 126 29 Figures figure NNS 10_1101-2020_12_31_424971 126 30 5c 5c CD 10_1101-2020_12_31_424971 126 31 , , , 10_1101-2020_12_31_424971 126 32 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 126 33 Figures Figures NNP 10_1101-2020_12_31_424971 126 34 2d 2d NNP 10_1101-2020_12_31_424971 126 35 - - HYPH 10_1101-2020_12_31_424971 126 36 f f NN 10_1101-2020_12_31_424971 126 37 ) ) -RRB- 10_1101-2020_12_31_424971 126 38 , , , 10_1101-2020_12_31_424971 126 39 consistent consistent JJ 10_1101-2020_12_31_424971 126 40 with with IN 10_1101-2020_12_31_424971 126 41 the the DT 10_1101-2020_12_31_424971 126 42 notion notion NN 10_1101-2020_12_31_424971 126 43 317 317 CD 10_1101-2020_12_31_424971 126 44 that that IN 10_1101-2020_12_31_424971 126 45 the the DT 10_1101-2020_12_31_424971 126 46 presence presence NN 10_1101-2020_12_31_424971 126 47 of of IN 10_1101-2020_12_31_424971 126 48 m1G37 m1g37 JJ 10_1101-2020_12_31_424971 126 49 compromises compromise NNS 10_1101-2020_12_31_424971 126 50 +1 +1 NNS 10_1101-2020_12_31_424971 126 51 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 126 52 . . . 10_1101-2020_12_31_424971 127 1 318 318 CD 10_1101-2020_12_31_424971 127 2 We -PRON- PRP 10_1101-2020_12_31_424971 127 3 then then RB 10_1101-2020_12_31_424971 127 4 used use VBD 10_1101-2020_12_31_424971 127 5 the the DT 10_1101-2020_12_31_424971 127 6 same same JJ 10_1101-2020_12_31_424971 127 7 tripeptide tripeptide NN 10_1101-2020_12_31_424971 127 8 formation formation NN 10_1101-2020_12_31_424971 127 9 assay assay NN 10_1101-2020_12_31_424971 127 10 to to TO 10_1101-2020_12_31_424971 127 11 determine determine VB 10_1101-2020_12_31_424971 127 12 how how WRB 10_1101-2020_12_31_424971 127 13 +1 +1 NNS 10_1101-2020_12_31_424971 127 14 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 127 15 during during IN 10_1101-2020_12_31_424971 127 16 319 319 CD 10_1101-2020_12_31_424971 127 17 translocation translocation NN 10_1101-2020_12_31_424971 127 18 of of IN 10_1101-2020_12_31_424971 127 19 G37-state G37-state NNP 10_1101-2020_12_31_424971 127 20 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 127 21 depends depend VBZ 10_1101-2020_12_31_424971 127 22 on on IN 10_1101-2020_12_31_424971 127 23 the the DT 10_1101-2020_12_31_424971 127 24 identity identity NN 10_1101-2020_12_31_424971 127 25 of of IN 10_1101-2020_12_31_424971 127 26 the the DT 10_1101-2020_12_31_424971 127 27 4th 4th JJ 10_1101-2020_12_31_424971 127 28 nucleotide nucleotide NN 10_1101-2020_12_31_424971 127 29 of of IN 10_1101-2020_12_31_424971 127 30 the the DT 10_1101-2020_12_31_424971 127 31 CCC CCC NNP 10_1101-2020_12_31_424971 127 32 - - HYPH 10_1101-2020_12_31_424971 127 33 C C NNP 10_1101-2020_12_31_424971 127 34 motif motif NN 10_1101-2020_12_31_424971 127 35 . . . 10_1101-2020_12_31_424971 128 1 320 320 CD 10_1101-2020_12_31_424971 128 2 A a DT 10_1101-2020_12_31_424971 128 3 series series NN 10_1101-2020_12_31_424971 128 4 of of IN 10_1101-2020_12_31_424971 128 5 POST POST NNP 10_1101-2020_12_31_424971 128 6 complexes complex NNS 10_1101-2020_12_31_424971 128 7 were be VBD 10_1101-2020_12_31_424971 128 8 generated generate VBN 10_1101-2020_12_31_424971 128 9 by by IN 10_1101-2020_12_31_424971 128 10 delivering deliver VBG 10_1101-2020_12_31_424971 128 11 G37-state G37-state NNP 10_1101-2020_12_31_424971 128 12 SufB2-TCs SufB2-TCs NNP 10_1101-2020_12_31_424971 128 13 and and CC 10_1101-2020_12_31_424971 128 14 EF EF NNP 10_1101-2020_12_31_424971 128 15 - - HYPH 10_1101-2020_12_31_424971 128 16 G G NNP 10_1101-2020_12_31_424971 128 17 to to IN 10_1101-2020_12_31_424971 128 18 321 321 CD 10_1101-2020_12_31_424971 128 19 70S 70S NNP 10_1101-2020_12_31_424971 128 20 ICs ic NNS 10_1101-2020_12_31_424971 128 21 programmed program VBD 10_1101-2020_12_31_424971 128 22 with with IN 10_1101-2020_12_31_424971 128 23 a a DT 10_1101-2020_12_31_424971 128 24 CCC CCC NNP 10_1101-2020_12_31_424971 128 25 - - HYPH 10_1101-2020_12_31_424971 128 26 N n NN 10_1101-2020_12_31_424971 128 27 motif motif NN 10_1101-2020_12_31_424971 128 28 at at IN 10_1101-2020_12_31_424971 128 29 the the DT 10_1101-2020_12_31_424971 128 30 2nd 2nd JJ 10_1101-2020_12_31_424971 128 31 codon codon NNP 10_1101-2020_12_31_424971 128 32 position position NN 10_1101-2020_12_31_424971 128 33 . . . 10_1101-2020_12_31_424971 129 1 Each each DT 10_1101-2020_12_31_424971 129 2 POST POST NNP 10_1101-2020_12_31_424971 129 3 complex complex NN 10_1101-2020_12_31_424971 129 4 was be VBD 10_1101-2020_12_31_424971 129 5 322 322 CD 10_1101-2020_12_31_424971 129 6 then then RB 10_1101-2020_12_31_424971 129 7 rapidly rapidly RB 10_1101-2020_12_31_424971 129 8 mixed mixed JJ 10_1101-2020_12_31_424971 129 9 with with IN 10_1101-2020_12_31_424971 129 10 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 129 11 - - HYPH 10_1101-2020_12_31_424971 129 12 TC TC NNP 10_1101-2020_12_31_424971 129 13 to to TO 10_1101-2020_12_31_424971 129 14 monitor monitor VB 10_1101-2020_12_31_424971 129 15 the the DT 10_1101-2020_12_31_424971 129 16 yield yield NN 10_1101-2020_12_31_424971 129 17 of of IN 10_1101-2020_12_31_424971 129 18 fMPV fMPV NNP 10_1101-2020_12_31_424971 129 19 and and CC 10_1101-2020_12_31_424971 129 20 kfMP+V kfMP+V NNP 10_1101-2020_12_31_424971 129 21 , , , 10_1101-2020_12_31_424971 129 22 obs obs NNP 10_1101-2020_12_31_424971 129 23 ( ( -LRB- 10_1101-2020_12_31_424971 129 24 Figure Figure NNP 10_1101-2020_12_31_424971 129 25 5d 5d NNP 10_1101-2020_12_31_424971 129 26 ) ) -RRB- 10_1101-2020_12_31_424971 129 27 . . . 10_1101-2020_12_31_424971 130 1 The the DT 10_1101-2020_12_31_424971 130 2 323 323 CD 10_1101-2020_12_31_424971 130 3 results result NNS 10_1101-2020_12_31_424971 130 4 showed show VBD 10_1101-2020_12_31_424971 130 5 a a DT 10_1101-2020_12_31_424971 130 6 high high JJ 10_1101-2020_12_31_424971 130 7 fMPV fmpv NN 10_1101-2020_12_31_424971 130 8 yield yield NN 10_1101-2020_12_31_424971 130 9 and and CC 10_1101-2020_12_31_424971 130 10 high high JJ 10_1101-2020_12_31_424971 130 11 kfMP+V kfMP+V NNP 10_1101-2020_12_31_424971 130 12 , , , 10_1101-2020_12_31_424971 130 13 obs obs NNP 10_1101-2020_12_31_424971 130 14 at at IN 10_1101-2020_12_31_424971 130 15 the the DT 10_1101-2020_12_31_424971 130 16 CCC-[C CCC-[C NNP 10_1101-2020_12_31_424971 130 17 / / SYM 10_1101-2020_12_31_424971 130 18 U U NNP 10_1101-2020_12_31_424971 130 19 ] ] -RRB- 10_1101-2020_12_31_424971 130 20 motifs motif NNS 10_1101-2020_12_31_424971 130 21 , , , 10_1101-2020_12_31_424971 130 22 but but CC 10_1101-2020_12_31_424971 130 23 a a DT 10_1101-2020_12_31_424971 130 24 low low JJ 10_1101-2020_12_31_424971 130 25 yield yield NN 10_1101-2020_12_31_424971 130 26 and and CC 10_1101-2020_12_31_424971 130 27 324 324 CD 10_1101-2020_12_31_424971 130 28 low low JJ 10_1101-2020_12_31_424971 130 29 kfMP+V kfMP+V NNP 10_1101-2020_12_31_424971 130 30 , , , 10_1101-2020_12_31_424971 130 31 obs obs NNP 10_1101-2020_12_31_424971 130 32 at at IN 10_1101-2020_12_31_424971 130 33 the the DT 10_1101-2020_12_31_424971 130 34 CCC-[A CCC-[A NNP 10_1101-2020_12_31_424971 130 35 / / SYM 10_1101-2020_12_31_424971 130 36 G G NNP 10_1101-2020_12_31_424971 130 37 ] ] -RRB- 10_1101-2020_12_31_424971 130 38 motifs motif NNS 10_1101-2020_12_31_424971 130 39 . . . 10_1101-2020_12_31_424971 131 1 This this DT 10_1101-2020_12_31_424971 131 2 indicates indicate VBZ 10_1101-2020_12_31_424971 131 3 that that IN 10_1101-2020_12_31_424971 131 4 high high JJ 10_1101-2020_12_31_424971 131 5 - - HYPH 10_1101-2020_12_31_424971 131 6 efficiency efficiency NN 10_1101-2020_12_31_424971 131 7 of of IN 10_1101-2020_12_31_424971 131 8 SufB2-induced SufB2-induced NNP 10_1101-2020_12_31_424971 131 9 +1 +1 NNS 10_1101-2020_12_31_424971 131 10 325 325 CD 10_1101-2020_12_31_424971 131 11 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 131 12 during during IN 10_1101-2020_12_31_424971 131 13 translocation translocation NN 10_1101-2020_12_31_424971 131 14 requires require VBZ 10_1101-2020_12_31_424971 131 15 the the DT 10_1101-2020_12_31_424971 131 16 presence presence NN 10_1101-2020_12_31_424971 131 17 of of IN 10_1101-2020_12_31_424971 131 18 a a DT 10_1101-2020_12_31_424971 131 19 [ [ -LRB- 10_1101-2020_12_31_424971 131 20 C C NNP 10_1101-2020_12_31_424971 131 21 / / SYM 10_1101-2020_12_31_424971 131 22 U u NN 10_1101-2020_12_31_424971 131 23 ] ] -RRB- 10_1101-2020_12_31_424971 131 24 at at IN 10_1101-2020_12_31_424971 131 25 the the DT 10_1101-2020_12_31_424971 131 26 4th 4th JJ 10_1101-2020_12_31_424971 131 27 nucleotide nucleotide NN 10_1101-2020_12_31_424971 131 28 of of IN 10_1101-2020_12_31_424971 131 29 the the DT 10_1101-2020_12_31_424971 131 30 326 326 CD 10_1101-2020_12_31_424971 131 31 CCC CCC NNP 10_1101-2020_12_31_424971 131 32 - - HYPH 10_1101-2020_12_31_424971 131 33 C C NNP 10_1101-2020_12_31_424971 131 34 motif motif NN 10_1101-2020_12_31_424971 131 35 . . . 10_1101-2020_12_31_424971 132 1 Because because IN 10_1101-2020_12_31_424971 132 2 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 132 3 in in IN 10_1101-2020_12_31_424971 132 4 these these DT 10_1101-2020_12_31_424971 132 5 experiments experiment NNS 10_1101-2020_12_31_424971 132 6 was be VBD 10_1101-2020_12_31_424971 132 7 in in IN 10_1101-2020_12_31_424971 132 8 the the DT 10_1101-2020_12_31_424971 132 9 G37-state G37-state NNP 10_1101-2020_12_31_424971 132 10 , , , 10_1101-2020_12_31_424971 132 11 it -PRON- PRP 10_1101-2020_12_31_424971 132 12 is be VBZ 10_1101-2020_12_31_424971 132 13 possible possible JJ 10_1101-2020_12_31_424971 132 14 that that IN 10_1101-2020_12_31_424971 132 15 a a DT 10_1101-2020_12_31_424971 132 16 sub-327 sub-327 NNP 10_1101-2020_12_31_424971 132 17 population population NN 10_1101-2020_12_31_424971 132 18 underwent undergo VBD 10_1101-2020_12_31_424971 132 19 +1 +1 NNS 10_1101-2020_12_31_424971 132 20 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 132 21 via via IN 10_1101-2020_12_31_424971 132 22 quadruplet quadruplet NN 10_1101-2020_12_31_424971 132 23 - - HYPH 10_1101-2020_12_31_424971 132 24 pairing pair VBG 10_1101-2020_12_31_424971 132 25 with with IN 10_1101-2020_12_31_424971 132 26 the the DT 10_1101-2020_12_31_424971 132 27 [ [ -LRB- 10_1101-2020_12_31_424971 132 28 C C NNP 10_1101-2020_12_31_424971 132 29 / / SYM 10_1101-2020_12_31_424971 132 30 U u NN 10_1101-2020_12_31_424971 132 31 ] ] -RRB- 10_1101-2020_12_31_424971 132 32 at at IN 10_1101-2020_12_31_424971 132 33 the the DT 10_1101-2020_12_31_424971 132 34 4th 4th NNP 10_1101-2020_12_31_424971 132 35 nucleotide nucleotide NNP 10_1101-2020_12_31_424971 132 36 328 328 CD 10_1101-2020_12_31_424971 132 37 of of IN 10_1101-2020_12_31_424971 132 38 the the DT 10_1101-2020_12_31_424971 132 39 CCC-[C CCC-[C NNP 10_1101-2020_12_31_424971 132 40 / / SYM 10_1101-2020_12_31_424971 132 41 U U NNP 10_1101-2020_12_31_424971 132 42 ] ] -RRB- 10_1101-2020_12_31_424971 132 43 motif motif NN 10_1101-2020_12_31_424971 132 44 during during IN 10_1101-2020_12_31_424971 132 45 translocation translocation NN 10_1101-2020_12_31_424971 132 46 . . . 10_1101-2020_12_31_424971 133 1 It -PRON- PRP 10_1101-2020_12_31_424971 133 2 is be VBZ 10_1101-2020_12_31_424971 133 3 also also RB 10_1101-2020_12_31_424971 133 4 possible possible JJ 10_1101-2020_12_31_424971 133 5 that that IN 10_1101-2020_12_31_424971 133 6 a a DT 10_1101-2020_12_31_424971 133 7 sub sub JJ 10_1101-2020_12_31_424971 133 8 - - NN 10_1101-2020_12_31_424971 133 9 population population NN 10_1101-2020_12_31_424971 133 10 underwent undergo VBD 10_1101-2020_12_31_424971 133 11 329 329 CD 10_1101-2020_12_31_424971 133 12 +1 +1 NNS 10_1101-2020_12_31_424971 133 13 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 133 14 via via IN 10_1101-2020_12_31_424971 133 15 triplet triplet NN 10_1101-2020_12_31_424971 133 16 - - HYPH 10_1101-2020_12_31_424971 133 17 slippage slippage NN 10_1101-2020_12_31_424971 133 18 , , , 10_1101-2020_12_31_424971 133 19 which which WDT 10_1101-2020_12_31_424971 133 20 could could MD 10_1101-2020_12_31_424971 133 21 potentially potentially RB 10_1101-2020_12_31_424971 133 22 be be VB 10_1101-2020_12_31_424971 133 23 inhibited inhibit VBN 10_1101-2020_12_31_424971 133 24 by by IN 10_1101-2020_12_31_424971 133 25 the the DT 10_1101-2020_12_31_424971 133 26 presence presence NN 10_1101-2020_12_31_424971 133 27 of of IN 10_1101-2020_12_31_424971 133 28 [ [ -LRB- 10_1101-2020_12_31_424971 133 29 G G NNP 10_1101-2020_12_31_424971 133 30 / / , 10_1101-2020_12_31_424971 133 31 A a NN 10_1101-2020_12_31_424971 133 32 ] ] -RRB- 10_1101-2020_12_31_424971 133 33 330 330 CD 10_1101-2020_12_31_424971 133 34 at at IN 10_1101-2020_12_31_424971 133 35 the the DT 10_1101-2020_12_31_424971 133 36 4th 4th JJ 10_1101-2020_12_31_424971 133 37 nucleotide nucleotide NN 10_1101-2020_12_31_424971 133 38 of of IN 10_1101-2020_12_31_424971 133 39 the the DT 10_1101-2020_12_31_424971 133 40 motif motif NN 10_1101-2020_12_31_424971 133 41 . . . 10_1101-2020_12_31_424971 134 1 To to TO 10_1101-2020_12_31_424971 134 2 verify verify VB 10_1101-2020_12_31_424971 134 3 that that IN 10_1101-2020_12_31_424971 134 4 the the DT 10_1101-2020_12_31_424971 134 5 POST POST NNP 10_1101-2020_12_31_424971 134 6 complex complex NN 10_1101-2020_12_31_424971 134 7 formed form VBN 10_1101-2020_12_31_424971 134 8 with with IN 10_1101-2020_12_31_424971 134 9 the the DT 10_1101-2020_12_31_424971 134 10 CCC CCC NNP 10_1101-2020_12_31_424971 134 11 - - HYPH 10_1101-2020_12_31_424971 134 12 A a DT 10_1101-2020_12_31_424971 134 13 331 331 CD 10_1101-2020_12_31_424971 134 14 sequence sequence NN 10_1101-2020_12_31_424971 134 15 was be VBD 10_1101-2020_12_31_424971 134 16 largely largely RB 10_1101-2020_12_31_424971 134 17 in in IN 10_1101-2020_12_31_424971 134 18 the the DT 10_1101-2020_12_31_424971 134 19 0-frame 0-frame CD 10_1101-2020_12_31_424971 134 20 , , , 10_1101-2020_12_31_424971 134 21 we -PRON- PRP 10_1101-2020_12_31_424971 134 22 rapidly rapidly RB 10_1101-2020_12_31_424971 134 23 mixed mix VBD 10_1101-2020_12_31_424971 134 24 the the DT 10_1101-2020_12_31_424971 134 25 complex complex NN 10_1101-2020_12_31_424971 134 26 with with IN 10_1101-2020_12_31_424971 134 27 an an DT 10_1101-2020_12_31_424971 134 28 equimolar equimolar JJ 10_1101-2020_12_31_424971 134 29 mixture mixture NN 10_1101-2020_12_31_424971 134 30 of of IN 10_1101-2020_12_31_424971 134 31 332 332 CD 10_1101-2020_12_31_424971 134 32 .CC .CC : 10_1101-2020_12_31_424971 134 33 - - HYPH 10_1101-2020_12_31_424971 134 34 BY by IN 10_1101-2020_12_31_424971 134 35 - - HYPH 10_1101-2020_12_31_424971 134 36 NC NC NNP 10_1101-2020_12_31_424971 134 37 - - HYPH 10_1101-2020_12_31_424971 134 38 ND ND NNP 10_1101-2020_12_31_424971 134 39 4.0 4.0 CD 10_1101-2020_12_31_424971 134 40 International International NNP 10_1101-2020_12_31_424971 134 41 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 134 42 under under IN 10_1101-2020_12_31_424971 134 43 a a DT 10_1101-2020_12_31_424971 134 44 ( ( -LRB- 10_1101-2020_12_31_424971 134 45 which which WDT 10_1101-2020_12_31_424971 134 46 was be VBD 10_1101-2020_12_31_424971 134 47 not not RB 10_1101-2020_12_31_424971 134 48 certified certify VBN 10_1101-2020_12_31_424971 134 49 by by IN 10_1101-2020_12_31_424971 134 50 peer peer NN 10_1101-2020_12_31_424971 134 51 review review NN 10_1101-2020_12_31_424971 134 52 ) ) -RRB- 10_1101-2020_12_31_424971 134 53 is be VBZ 10_1101-2020_12_31_424971 134 54 the the DT 10_1101-2020_12_31_424971 134 55 author author NN 10_1101-2020_12_31_424971 134 56 / / SYM 10_1101-2020_12_31_424971 134 57 funder funder NN 10_1101-2020_12_31_424971 134 58 , , , 10_1101-2020_12_31_424971 134 59 who who WP 10_1101-2020_12_31_424971 134 60 has have VBZ 10_1101-2020_12_31_424971 134 61 granted grant VBN 10_1101-2020_12_31_424971 134 62 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 134 63 a a DT 10_1101-2020_12_31_424971 134 64 license license NN 10_1101-2020_12_31_424971 134 65 to to TO 10_1101-2020_12_31_424971 134 66 display display VB 10_1101-2020_12_31_424971 134 67 the the DT 10_1101-2020_12_31_424971 134 68 preprint preprint NN 10_1101-2020_12_31_424971 134 69 in in IN 10_1101-2020_12_31_424971 134 70 perpetuity perpetuity NN 10_1101-2020_12_31_424971 134 71 . . . 10_1101-2020_12_31_424971 135 1 It -PRON- PRP 10_1101-2020_12_31_424971 135 2 is be VBZ 10_1101-2020_12_31_424971 135 3 made make VBN 10_1101-2020_12_31_424971 135 4 The the DT 10_1101-2020_12_31_424971 135 5 copyright copyright NN 10_1101-2020_12_31_424971 135 6 holder holder NN 10_1101-2020_12_31_424971 135 7 for for IN 10_1101-2020_12_31_424971 135 8 this this DT 10_1101-2020_12_31_424971 135 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 135 10 version version NN 10_1101-2020_12_31_424971 135 11 posted post VBD 10_1101-2020_12_31_424971 135 12 January January NNP 10_1101-2020_12_31_424971 135 13 2 2 CD 10_1101-2020_12_31_424971 135 14 , , , 10_1101-2020_12_31_424971 135 15 2021 2021 CD 10_1101-2020_12_31_424971 135 16 . . . 10_1101-2020_12_31_424971 135 17 ; ; : 10_1101-2020_12_31_424971 135 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 135 19 : : : 10_1101-2020_12_31_424971 135 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 135 21 preprint preprint NN 10_1101-2020_12_31_424971 135 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 135 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 135 24 14 14 CD 10_1101-2020_12_31_424971 135 25 tRNASer tRNASer NNP 10_1101-2020_12_31_424971 135 26 - - HYPH 10_1101-2020_12_31_424971 135 27 TC TC NNP 10_1101-2020_12_31_424971 135 28 , , , 10_1101-2020_12_31_424971 135 29 cognate cognate VB 10_1101-2020_12_31_424971 135 30 to to IN 10_1101-2020_12_31_424971 135 31 the the DT 10_1101-2020_12_31_424971 135 32 next next JJ 10_1101-2020_12_31_424971 135 33 A A NNP 10_1101-2020_12_31_424971 135 34 - - HYPH 10_1101-2020_12_31_424971 135 35 site site NN 10_1101-2020_12_31_424971 135 36 codon codon NN 10_1101-2020_12_31_424971 135 37 in in IN 10_1101-2020_12_31_424971 135 38 the the DT 10_1101-2020_12_31_424971 135 39 0-frame 0-frame CD 10_1101-2020_12_31_424971 135 40 ( ( -LRB- 10_1101-2020_12_31_424971 135 41 AGU AGU NNP 10_1101-2020_12_31_424971 135 42 ) ) -RRB- 10_1101-2020_12_31_424971 135 43 , , , 10_1101-2020_12_31_424971 135 44 and and CC 10_1101-2020_12_31_424971 135 45 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 135 46 - - HYPH 10_1101-2020_12_31_424971 135 47 TC TC NNP 10_1101-2020_12_31_424971 135 48 , , , 10_1101-2020_12_31_424971 135 49 cognate cognate VBP 10_1101-2020_12_31_424971 135 50 to to IN 10_1101-2020_12_31_424971 135 51 333 333 CD 10_1101-2020_12_31_424971 135 52 the the DT 10_1101-2020_12_31_424971 135 53 next next JJ 10_1101-2020_12_31_424971 135 54 A a NN 10_1101-2020_12_31_424971 135 55 - - HYPH 10_1101-2020_12_31_424971 135 56 site site NN 10_1101-2020_12_31_424971 135 57 codon codon NN 10_1101-2020_12_31_424971 135 58 in in IN 10_1101-2020_12_31_424971 135 59 the the DT 10_1101-2020_12_31_424971 135 60 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 135 61 ( ( -LRB- 10_1101-2020_12_31_424971 135 62 GUU GUU NNP 10_1101-2020_12_31_424971 135 63 ) ) -RRB- 10_1101-2020_12_31_424971 135 64 ( ( -LRB- 10_1101-2020_12_31_424971 135 65 Figure figure NN 10_1101-2020_12_31_424971 135 66 5e 5e NN 10_1101-2020_12_31_424971 135 67 ) ) -RRB- 10_1101-2020_12_31_424971 135 68 . . . 10_1101-2020_12_31_424971 136 1 The the DT 10_1101-2020_12_31_424971 136 2 results result NNS 10_1101-2020_12_31_424971 136 3 showed show VBD 10_1101-2020_12_31_424971 136 4 a a DT 10_1101-2020_12_31_424971 136 5 high high JJ 10_1101-2020_12_31_424971 136 6 yield yield NN 10_1101-2020_12_31_424971 136 7 and and CC 10_1101-2020_12_31_424971 136 8 334 334 CD 10_1101-2020_12_31_424971 136 9 high high JJ 10_1101-2020_12_31_424971 136 10 kfMP+S kfmp+s NN 10_1101-2020_12_31_424971 136 11 , , , 10_1101-2020_12_31_424971 136 12 obs obs NN 10_1101-2020_12_31_424971 136 13 , , , 10_1101-2020_12_31_424971 136 14 supporting support VBG 10_1101-2020_12_31_424971 136 15 the the DT 10_1101-2020_12_31_424971 136 16 notion notion NN 10_1101-2020_12_31_424971 136 17 that that IN 10_1101-2020_12_31_424971 136 18 the the DT 10_1101-2020_12_31_424971 136 19 POST POST NNP 10_1101-2020_12_31_424971 136 20 complex complex NN 10_1101-2020_12_31_424971 136 21 formed form VBN 10_1101-2020_12_31_424971 136 22 with with IN 10_1101-2020_12_31_424971 136 23 the the DT 10_1101-2020_12_31_424971 136 24 CCC CCC NNP 10_1101-2020_12_31_424971 136 25 - - HYPH 10_1101-2020_12_31_424971 136 26 A a DT 10_1101-2020_12_31_424971 136 27 motif motif NN 10_1101-2020_12_31_424971 136 28 was be VBD 10_1101-2020_12_31_424971 136 29 335 335 CD 10_1101-2020_12_31_424971 136 30 largely largely RB 10_1101-2020_12_31_424971 136 31 in in IN 10_1101-2020_12_31_424971 136 32 the the DT 10_1101-2020_12_31_424971 136 33 0-frame 0-frame CD 10_1101-2020_12_31_424971 136 34 . . . 10_1101-2020_12_31_424971 137 1 Thus thus RB 10_1101-2020_12_31_424971 137 2 , , , 10_1101-2020_12_31_424971 137 3 the the DT 10_1101-2020_12_31_424971 137 4 4th 4th JJ 10_1101-2020_12_31_424971 137 5 nucleotide nucleotide NN 10_1101-2020_12_31_424971 137 6 of of IN 10_1101-2020_12_31_424971 137 7 the the DT 10_1101-2020_12_31_424971 137 8 CCC CCC NNP 10_1101-2020_12_31_424971 137 9 - - HYPH 10_1101-2020_12_31_424971 137 10 C C NNP 10_1101-2020_12_31_424971 137 11 motif motif NN 10_1101-2020_12_31_424971 137 12 plays play VBZ 10_1101-2020_12_31_424971 137 13 a a DT 10_1101-2020_12_31_424971 137 14 role role NN 10_1101-2020_12_31_424971 137 15 in in IN 10_1101-2020_12_31_424971 137 16 determining determine VBG 10_1101-2020_12_31_424971 137 17 +1 +1 NNS 10_1101-2020_12_31_424971 137 18 336 336 CD 10_1101-2020_12_31_424971 137 19 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 137 20 during during IN 10_1101-2020_12_31_424971 137 21 translocation translocation NN 10_1101-2020_12_31_424971 137 22 of of IN 10_1101-2020_12_31_424971 137 23 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 137 24 from from IN 10_1101-2020_12_31_424971 137 25 the the DT 10_1101-2020_12_31_424971 137 26 A A NNP 10_1101-2020_12_31_424971 137 27 site site NN 10_1101-2020_12_31_424971 137 28 to to IN 10_1101-2020_12_31_424971 137 29 the the DT 10_1101-2020_12_31_424971 137 30 P p NN 10_1101-2020_12_31_424971 137 31 site site NN 10_1101-2020_12_31_424971 137 32 . . . 10_1101-2020_12_31_424971 138 1 337 337 CD 10_1101-2020_12_31_424971 138 2 338 338 CD 10_1101-2020_12_31_424971 138 3 The the DT 10_1101-2020_12_31_424971 138 4 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 138 5 efficiency efficiency NN 10_1101-2020_12_31_424971 138 6 of of IN 10_1101-2020_12_31_424971 138 7 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 138 8 depends depend VBZ 10_1101-2020_12_31_424971 138 9 on on IN 10_1101-2020_12_31_424971 138 10 sequences sequence NNS 10_1101-2020_12_31_424971 138 11 of of IN 10_1101-2020_12_31_424971 138 12 the the DT 10_1101-2020_12_31_424971 138 13 anticodon anticodon NNP 10_1101-2020_12_31_424971 138 14 loop loop NN 10_1101-2020_12_31_424971 138 15 and and CC 10_1101-2020_12_31_424971 138 16 339 339 CD 10_1101-2020_12_31_424971 138 17 the the DT 10_1101-2020_12_31_424971 138 18 CCC CCC NNP 10_1101-2020_12_31_424971 138 19 - - HYPH 10_1101-2020_12_31_424971 138 20 C C NNP 10_1101-2020_12_31_424971 138 21 motif motif NN 10_1101-2020_12_31_424971 138 22 340 340 CD 10_1101-2020_12_31_424971 138 23 To to TO 10_1101-2020_12_31_424971 138 24 determine determine VB 10_1101-2020_12_31_424971 138 25 whether whether IN 10_1101-2020_12_31_424971 138 26 the the DT 10_1101-2020_12_31_424971 138 27 +1-frameshifting +1-frameshifting JJ 10_1101-2020_12_31_424971 138 28 efficiency efficiency NN 10_1101-2020_12_31_424971 138 29 of of IN 10_1101-2020_12_31_424971 138 30 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 138 31 during during IN 10_1101-2020_12_31_424971 138 32 translocation translocation NN 10_1101-2020_12_31_424971 138 33 is be VBZ 10_1101-2020_12_31_424971 138 34 influenced influence VBN 10_1101-2020_12_31_424971 138 35 341 341 CD 10_1101-2020_12_31_424971 138 36 by by IN 10_1101-2020_12_31_424971 138 37 sequences sequence NNS 10_1101-2020_12_31_424971 138 38 of of IN 10_1101-2020_12_31_424971 138 39 the the DT 10_1101-2020_12_31_424971 138 40 anticodon anticodon NNP 10_1101-2020_12_31_424971 138 41 loop loop NN 10_1101-2020_12_31_424971 138 42 and and CC 10_1101-2020_12_31_424971 138 43 the the DT 10_1101-2020_12_31_424971 138 44 CCC CCC NNP 10_1101-2020_12_31_424971 138 45 - - HYPH 10_1101-2020_12_31_424971 138 46 C C NNP 10_1101-2020_12_31_424971 138 47 motif motif NN 10_1101-2020_12_31_424971 138 48 , , , 10_1101-2020_12_31_424971 138 49 we -PRON- PRP 10_1101-2020_12_31_424971 138 50 performed perform VBD 10_1101-2020_12_31_424971 138 51 tripeptide tripeptide NN 10_1101-2020_12_31_424971 138 52 formation formation NN 10_1101-2020_12_31_424971 138 53 342 342 CD 10_1101-2020_12_31_424971 138 54 assays assay NNS 10_1101-2020_12_31_424971 138 55 and and CC 10_1101-2020_12_31_424971 138 56 monitored monitor VBD 10_1101-2020_12_31_424971 138 57 the the DT 10_1101-2020_12_31_424971 138 58 yield yield NN 10_1101-2020_12_31_424971 138 59 of of IN 10_1101-2020_12_31_424971 138 60 fMPV fmpv NN 10_1101-2020_12_31_424971 138 61 . . . 10_1101-2020_12_31_424971 139 1 In in IN 10_1101-2020_12_31_424971 139 2 these these DT 10_1101-2020_12_31_424971 139 3 experiments experiment NNS 10_1101-2020_12_31_424971 139 4 , , , 10_1101-2020_12_31_424971 139 5 we -PRON- PRP 10_1101-2020_12_31_424971 139 6 varied vary VBD 10_1101-2020_12_31_424971 139 7 the the DT 10_1101-2020_12_31_424971 139 8 sequence sequence NN 10_1101-2020_12_31_424971 139 9 of of IN 10_1101-2020_12_31_424971 139 10 the the DT 10_1101-2020_12_31_424971 139 11 343 343 CD 10_1101-2020_12_31_424971 139 12 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 139 13 anticodon anticodon NN 10_1101-2020_12_31_424971 139 14 loop loop NN 10_1101-2020_12_31_424971 139 15 and/or and/or CC 10_1101-2020_12_31_424971 139 16 the the DT 10_1101-2020_12_31_424971 139 17 CCC CCC NNP 10_1101-2020_12_31_424971 139 18 - - HYPH 10_1101-2020_12_31_424971 139 19 C C NNP 10_1101-2020_12_31_424971 139 20 motif motif NN 10_1101-2020_12_31_424971 139 21 at at IN 10_1101-2020_12_31_424971 139 22 the the DT 10_1101-2020_12_31_424971 139 23 2nd 2nd JJ 10_1101-2020_12_31_424971 139 24 codon codon NNP 10_1101-2020_12_31_424971 139 25 position position NN 10_1101-2020_12_31_424971 139 26 of of IN 10_1101-2020_12_31_424971 139 27 the the DT 10_1101-2020_12_31_424971 139 28 mRNA mrna NN 10_1101-2020_12_31_424971 139 29 . . . 10_1101-2020_12_31_424971 140 1 To to TO 10_1101-2020_12_31_424971 140 2 explore explore VB 10_1101-2020_12_31_424971 140 3 344 344 CD 10_1101-2020_12_31_424971 140 4 the the DT 10_1101-2020_12_31_424971 140 5 possibilities possibility NNS 10_1101-2020_12_31_424971 140 6 of of IN 10_1101-2020_12_31_424971 140 7 both both CC 10_1101-2020_12_31_424971 140 8 triplet triplet NN 10_1101-2020_12_31_424971 140 9 - - HYPH 10_1101-2020_12_31_424971 140 10 slippage slippage NN 10_1101-2020_12_31_424971 140 11 and and CC 10_1101-2020_12_31_424971 140 12 quadruplet quadruplet NN 10_1101-2020_12_31_424971 140 13 - - HYPH 10_1101-2020_12_31_424971 140 14 pairing pair VBG 10_1101-2020_12_31_424971 140 15 , , , 10_1101-2020_12_31_424971 140 16 we -PRON- PRP 10_1101-2020_12_31_424971 140 17 used use VBD 10_1101-2020_12_31_424971 140 18 variants variant NNS 10_1101-2020_12_31_424971 140 19 of of IN 10_1101-2020_12_31_424971 140 20 G37-state G37-state NNP 10_1101-2020_12_31_424971 140 21 345 345 CD 10_1101-2020_12_31_424971 140 22 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 140 23 . . . 10_1101-2020_12_31_424971 141 1 We -PRON- PRP 10_1101-2020_12_31_424971 141 2 showed show VBD 10_1101-2020_12_31_424971 141 3 that that IN 10_1101-2020_12_31_424971 141 4 variants variant NNS 10_1101-2020_12_31_424971 141 5 with with IN 10_1101-2020_12_31_424971 141 6 the the DT 10_1101-2020_12_31_424971 141 7 potential potential NN 10_1101-2020_12_31_424971 141 8 to to TO 10_1101-2020_12_31_424971 141 9 undergo undergo VB 10_1101-2020_12_31_424971 141 10 quadruplet quadruplet NN 10_1101-2020_12_31_424971 141 11 - - HYPH 10_1101-2020_12_31_424971 141 12 pairing pair VBG 10_1101-2020_12_31_424971 141 13 with with IN 10_1101-2020_12_31_424971 141 14 the the DT 10_1101-2020_12_31_424971 141 15 CCC CCC NNP 10_1101-2020_12_31_424971 141 16 - - HYPH 10_1101-2020_12_31_424971 141 17 C C NNP 10_1101-2020_12_31_424971 141 18 346 346 CD 10_1101-2020_12_31_424971 141 19 motif motif NN 10_1101-2020_12_31_424971 141 20 resulted result VBD 10_1101-2020_12_31_424971 141 21 in in IN 10_1101-2020_12_31_424971 141 22 fMPV fmpv JJ 10_1101-2020_12_31_424971 141 23 yields yield NNS 10_1101-2020_12_31_424971 141 24 of of IN 10_1101-2020_12_31_424971 141 25 87 87 CD 10_1101-2020_12_31_424971 141 26 % % NN 10_1101-2020_12_31_424971 141 27 and and CC 10_1101-2020_12_31_424971 141 28 62 62 CD 10_1101-2020_12_31_424971 141 29 % % NN 10_1101-2020_12_31_424971 141 30 ( ( -LRB- 10_1101-2020_12_31_424971 141 31 Figures Figures NNP 10_1101-2020_12_31_424971 141 32 4c 4c NNP 10_1101-2020_12_31_424971 141 33 , , , 10_1101-2020_12_31_424971 141 34 d d NN 10_1101-2020_12_31_424971 141 35 ) ) -RRB- 10_1101-2020_12_31_424971 141 36 . . . 10_1101-2020_12_31_424971 142 1 The the DT 10_1101-2020_12_31_424971 142 2 different different JJ 10_1101-2020_12_31_424971 142 3 yields yield NNS 10_1101-2020_12_31_424971 142 4 suggest suggest VBP 10_1101-2020_12_31_424971 142 5 that that IN 10_1101-2020_12_31_424971 142 6 347 347 CD 10_1101-2020_12_31_424971 142 7 G37-state G37-state NNP 10_1101-2020_12_31_424971 142 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 142 9 variants variant NNS 10_1101-2020_12_31_424971 142 10 can can MD 10_1101-2020_12_31_424971 142 11 induce induce VB 10_1101-2020_12_31_424971 142 12 triplet triplet NN 10_1101-2020_12_31_424971 142 13 - - HYPH 10_1101-2020_12_31_424971 142 14 slippage slippage NN 10_1101-2020_12_31_424971 142 15 and/or and/or CC 10_1101-2020_12_31_424971 142 16 engage engage VB 10_1101-2020_12_31_424971 142 17 in in IN 10_1101-2020_12_31_424971 142 18 quadruplet quadruplet NN 10_1101-2020_12_31_424971 142 19 - - HYPH 10_1101-2020_12_31_424971 142 20 pairing pair VBG 10_1101-2020_12_31_424971 142 21 with with IN 10_1101-2020_12_31_424971 142 22 348 348 CD 10_1101-2020_12_31_424971 142 23 different different JJ 10_1101-2020_12_31_424971 142 24 efficiencies efficiency NNS 10_1101-2020_12_31_424971 142 25 during during IN 10_1101-2020_12_31_424971 142 26 translocation translocation NN 10_1101-2020_12_31_424971 142 27 . . . 10_1101-2020_12_31_424971 143 1 Analogous analogous JJ 10_1101-2020_12_31_424971 143 2 experiments experiment NNS 10_1101-2020_12_31_424971 143 3 showed show VBD 10_1101-2020_12_31_424971 143 4 that that IN 10_1101-2020_12_31_424971 143 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 143 6 variants variant NNS 10_1101-2020_12_31_424971 143 7 349 349 CD 10_1101-2020_12_31_424971 143 8 that that WDT 10_1101-2020_12_31_424971 143 9 were be VBD 10_1101-2020_12_31_424971 143 10 restricted restrict VBN 10_1101-2020_12_31_424971 143 11 to to IN 10_1101-2020_12_31_424971 143 12 triplet triplet NN 10_1101-2020_12_31_424971 143 13 - - HYPH 10_1101-2020_12_31_424971 143 14 pairing pairing NN 10_1101-2020_12_31_424971 143 15 resulted result VBD 10_1101-2020_12_31_424971 143 16 in in IN 10_1101-2020_12_31_424971 143 17 reduced reduce VBN 10_1101-2020_12_31_424971 143 18 fMPV fmpv NN 10_1101-2020_12_31_424971 143 19 yields yield NNS 10_1101-2020_12_31_424971 143 20 ( ( -LRB- 10_1101-2020_12_31_424971 143 21 26 26 CD 10_1101-2020_12_31_424971 143 22 % % NN 10_1101-2020_12_31_424971 143 23 and and CC 10_1101-2020_12_31_424971 143 24 20 20 CD 10_1101-2020_12_31_424971 143 25 % % NN 10_1101-2020_12_31_424971 143 26 , , , 10_1101-2020_12_31_424971 143 27 respectively respectively RB 10_1101-2020_12_31_424971 143 28 ) ) -RRB- 10_1101-2020_12_31_424971 143 29 350 350 CD 10_1101-2020_12_31_424971 143 30 upon upon IN 10_1101-2020_12_31_424971 143 31 pairing pair VBG 10_1101-2020_12_31_424971 143 32 with with IN 10_1101-2020_12_31_424971 143 33 a a DT 10_1101-2020_12_31_424971 143 34 CCC CCC NNP 10_1101-2020_12_31_424971 143 35 - - HYPH 10_1101-2020_12_31_424971 143 36 C C NNP 10_1101-2020_12_31_424971 143 37 motif motif NN 10_1101-2020_12_31_424971 143 38 ( ( -LRB- 10_1101-2020_12_31_424971 143 39 Figures Figures NNPS 10_1101-2020_12_31_424971 143 40 4e 4e NNP 10_1101-2020_12_31_424971 143 41 , , , 10_1101-2020_12_31_424971 143 42 f f NN 10_1101-2020_12_31_424971 143 43 ) ) -RRB- 10_1101-2020_12_31_424971 143 44 . . . 10_1101-2020_12_31_424971 144 1 Collectively collectively RB 10_1101-2020_12_31_424971 144 2 , , , 10_1101-2020_12_31_424971 144 3 these these DT 10_1101-2020_12_31_424971 144 4 results result NNS 10_1101-2020_12_31_424971 144 5 suggest suggest VBP 10_1101-2020_12_31_424971 144 6 that that IN 10_1101-2020_12_31_424971 144 7 there there EX 10_1101-2020_12_31_424971 144 8 is be VBZ 10_1101-2020_12_31_424971 144 9 351 351 CD 10_1101-2020_12_31_424971 144 10 considerable considerable JJ 10_1101-2020_12_31_424971 144 11 plasticity plasticity NN 10_1101-2020_12_31_424971 144 12 in in IN 10_1101-2020_12_31_424971 144 13 the the DT 10_1101-2020_12_31_424971 144 14 mechanisms mechanism NNS 10_1101-2020_12_31_424971 144 15 that that WDT 10_1101-2020_12_31_424971 144 16 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 144 17 uses use VBZ 10_1101-2020_12_31_424971 144 18 to to TO 10_1101-2020_12_31_424971 144 19 induce induce VB 10_1101-2020_12_31_424971 144 20 +1 +1 NNS 10_1101-2020_12_31_424971 144 21 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 144 22 during during IN 10_1101-2020_12_31_424971 144 23 352 352 CD 10_1101-2020_12_31_424971 144 24 translocation translocation NN 10_1101-2020_12_31_424971 144 25 and and CC 10_1101-2020_12_31_424971 144 26 in in IN 10_1101-2020_12_31_424971 144 27 the the DT 10_1101-2020_12_31_424971 144 28 efficiencies efficiency NNS 10_1101-2020_12_31_424971 144 29 of of IN 10_1101-2020_12_31_424971 144 30 these these DT 10_1101-2020_12_31_424971 144 31 mechanisms mechanism NNS 10_1101-2020_12_31_424971 144 32 . . . 10_1101-2020_12_31_424971 145 1 353 353 CD 10_1101-2020_12_31_424971 145 2 354 354 CD 10_1101-2020_12_31_424971 145 3 An an DT 10_1101-2020_12_31_424971 145 4 smFRET smFRET NNP 10_1101-2020_12_31_424971 145 5 signal signal NN 10_1101-2020_12_31_424971 145 6 that that WDT 10_1101-2020_12_31_424971 145 7 reports report VBZ 10_1101-2020_12_31_424971 145 8 on on IN 10_1101-2020_12_31_424971 145 9 ribosome ribosome JJ 10_1101-2020_12_31_424971 145 10 dynamics dynamic NNS 10_1101-2020_12_31_424971 145 11 during during IN 10_1101-2020_12_31_424971 145 12 individual individual JJ 10_1101-2020_12_31_424971 145 13 elongation elongation NN 10_1101-2020_12_31_424971 145 14 cycles cycle NNS 10_1101-2020_12_31_424971 145 15 355 355 CD 10_1101-2020_12_31_424971 145 16 To to TO 10_1101-2020_12_31_424971 145 17 address address VB 10_1101-2020_12_31_424971 145 18 the the DT 10_1101-2020_12_31_424971 145 19 mechanism mechanism NN 10_1101-2020_12_31_424971 145 20 of of IN 10_1101-2020_12_31_424971 145 21 SufB2-induced SufB2-induced NNP 10_1101-2020_12_31_424971 145 22 +1 +1 NNS 10_1101-2020_12_31_424971 145 23 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 145 24 during during IN 10_1101-2020_12_31_424971 145 25 translocation translocation NN 10_1101-2020_12_31_424971 145 26 , , , 10_1101-2020_12_31_424971 145 27 we -PRON- PRP 10_1101-2020_12_31_424971 145 28 used use VBD 10_1101-2020_12_31_424971 145 29 356 356 CD 10_1101-2020_12_31_424971 145 30 a a DT 10_1101-2020_12_31_424971 145 31 previously previously RB 10_1101-2020_12_31_424971 145 32 developed develop VBN 10_1101-2020_12_31_424971 145 33 smFRET smfret NN 10_1101-2020_12_31_424971 145 34 signal signal NN 10_1101-2020_12_31_424971 145 35 to to TO 10_1101-2020_12_31_424971 145 36 determine determine VB 10_1101-2020_12_31_424971 145 37 whether whether IN 10_1101-2020_12_31_424971 145 38 and and CC 10_1101-2020_12_31_424971 145 39 how how WRB 10_1101-2020_12_31_424971 145 40 SufB2 sufb2 JJ 10_1101-2020_12_31_424971 145 41 alters alter VBZ 10_1101-2020_12_31_424971 145 42 the the DT 10_1101-2020_12_31_424971 145 43 rates rate NNS 10_1101-2020_12_31_424971 145 44 with with IN 10_1101-2020_12_31_424971 145 45 357 357 CD 10_1101-2020_12_31_424971 145 46 which which WDT 10_1101-2020_12_31_424971 145 47 the the DT 10_1101-2020_12_31_424971 145 48 ribosome ribosome NN 10_1101-2020_12_31_424971 145 49 undergoes undergo VBZ 10_1101-2020_12_31_424971 145 50 a a DT 10_1101-2020_12_31_424971 145 51 series series NN 10_1101-2020_12_31_424971 145 52 of of IN 10_1101-2020_12_31_424971 145 53 conformational conformational JJ 10_1101-2020_12_31_424971 145 54 changes change NNS 10_1101-2020_12_31_424971 145 55 that that WDT 10_1101-2020_12_31_424971 145 56 drive drive VBP 10_1101-2020_12_31_424971 145 57 and and CC 10_1101-2020_12_31_424971 145 58 regulate regulate VB 10_1101-2020_12_31_424971 145 59 the the DT 10_1101-2020_12_31_424971 145 60 358 358 CD 10_1101-2020_12_31_424971 145 61 .CC .CC NFP 10_1101-2020_12_31_424971 145 62 - - HYPH 10_1101-2020_12_31_424971 145 63 BY by IN 10_1101-2020_12_31_424971 145 64 - - HYPH 10_1101-2020_12_31_424971 145 65 NC NC NNP 10_1101-2020_12_31_424971 145 66 - - HYPH 10_1101-2020_12_31_424971 145 67 ND ND NNP 10_1101-2020_12_31_424971 145 68 4.0 4.0 CD 10_1101-2020_12_31_424971 145 69 International International NNP 10_1101-2020_12_31_424971 145 70 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 145 71 under under IN 10_1101-2020_12_31_424971 145 72 a a DT 10_1101-2020_12_31_424971 145 73 ( ( -LRB- 10_1101-2020_12_31_424971 145 74 which which WDT 10_1101-2020_12_31_424971 145 75 was be VBD 10_1101-2020_12_31_424971 145 76 not not RB 10_1101-2020_12_31_424971 145 77 certified certify VBN 10_1101-2020_12_31_424971 145 78 by by IN 10_1101-2020_12_31_424971 145 79 peer peer NN 10_1101-2020_12_31_424971 145 80 review review NN 10_1101-2020_12_31_424971 145 81 ) ) -RRB- 10_1101-2020_12_31_424971 145 82 is be VBZ 10_1101-2020_12_31_424971 145 83 the the DT 10_1101-2020_12_31_424971 145 84 author author NN 10_1101-2020_12_31_424971 145 85 / / SYM 10_1101-2020_12_31_424971 145 86 funder funder NN 10_1101-2020_12_31_424971 145 87 , , , 10_1101-2020_12_31_424971 145 88 who who WP 10_1101-2020_12_31_424971 145 89 has have VBZ 10_1101-2020_12_31_424971 145 90 granted grant VBN 10_1101-2020_12_31_424971 145 91 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 145 92 a a DT 10_1101-2020_12_31_424971 145 93 license license NN 10_1101-2020_12_31_424971 145 94 to to TO 10_1101-2020_12_31_424971 145 95 display display VB 10_1101-2020_12_31_424971 145 96 the the DT 10_1101-2020_12_31_424971 145 97 preprint preprint NN 10_1101-2020_12_31_424971 145 98 in in IN 10_1101-2020_12_31_424971 145 99 perpetuity perpetuity NN 10_1101-2020_12_31_424971 145 100 . . . 10_1101-2020_12_31_424971 146 1 It -PRON- PRP 10_1101-2020_12_31_424971 146 2 is be VBZ 10_1101-2020_12_31_424971 146 3 made make VBN 10_1101-2020_12_31_424971 146 4 The the DT 10_1101-2020_12_31_424971 146 5 copyright copyright NN 10_1101-2020_12_31_424971 146 6 holder holder NN 10_1101-2020_12_31_424971 146 7 for for IN 10_1101-2020_12_31_424971 146 8 this this DT 10_1101-2020_12_31_424971 146 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 146 10 version version NN 10_1101-2020_12_31_424971 146 11 posted post VBD 10_1101-2020_12_31_424971 146 12 January January NNP 10_1101-2020_12_31_424971 146 13 2 2 CD 10_1101-2020_12_31_424971 146 14 , , , 10_1101-2020_12_31_424971 146 15 2021 2021 CD 10_1101-2020_12_31_424971 146 16 . . . 10_1101-2020_12_31_424971 146 17 ; ; : 10_1101-2020_12_31_424971 146 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 146 19 : : : 10_1101-2020_12_31_424971 146 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 146 21 preprint preprint NN 10_1101-2020_12_31_424971 146 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 146 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 146 24 15 15 CD 10_1101-2020_12_31_424971 146 25 elongation elongation NN 10_1101-2020_12_31_424971 146 26 cycle35 cycle35 VBZ 10_1101-2020_12_31_424971 146 27 ( ( -LRB- 10_1101-2020_12_31_424971 146 28 Figures Figures NNP 10_1101-2020_12_31_424971 146 29 6a 6a NNP 10_1101-2020_12_31_424971 146 30 - - HYPH 10_1101-2020_12_31_424971 146 31 c c NNP 10_1101-2020_12_31_424971 146 32 ) ) -RRB- 10_1101-2020_12_31_424971 146 33 . . . 10_1101-2020_12_31_424971 147 1 This this DT 10_1101-2020_12_31_424971 147 2 signal signal NN 10_1101-2020_12_31_424971 147 3 is be VBZ 10_1101-2020_12_31_424971 147 4 generated generate VBN 10_1101-2020_12_31_424971 147 5 using use VBG 10_1101-2020_12_31_424971 147 6 a a DT 10_1101-2020_12_31_424971 147 7 ribosomal ribosomal NN 10_1101-2020_12_31_424971 147 8 large large JJ 10_1101-2020_12_31_424971 147 9 , , , 10_1101-2020_12_31_424971 147 10 or or CC 10_1101-2020_12_31_424971 147 11 50S 50S NNP 10_1101-2020_12_31_424971 147 12 , , , 10_1101-2020_12_31_424971 147 13 subunit subunit NN 10_1101-2020_12_31_424971 147 14 359 359 CD 10_1101-2020_12_31_424971 147 15 that that WDT 10_1101-2020_12_31_424971 147 16 has have VBZ 10_1101-2020_12_31_424971 147 17 been be VBN 10_1101-2020_12_31_424971 147 18 Cy3- cy3- CD 10_1101-2020_12_31_424971 147 19 and and CC 10_1101-2020_12_31_424971 147 20 Cy5-labeled cy5-labele VBN 10_1101-2020_12_31_424971 147 21 at at IN 10_1101-2020_12_31_424971 147 22 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 147 23 proteins protein NNS 10_1101-2020_12_31_424971 147 24 bL9 bL9 NNP 10_1101-2020_12_31_424971 147 25 and and CC 10_1101-2020_12_31_424971 147 26 uL1 uL1 NNP 10_1101-2020_12_31_424971 147 27 , , , 10_1101-2020_12_31_424971 147 28 respectively respectively RB 10_1101-2020_12_31_424971 147 29 , , , 10_1101-2020_12_31_424971 147 30 to to TO 10_1101-2020_12_31_424971 147 31 report report VB 10_1101-2020_12_31_424971 147 32 on on IN 10_1101-2020_12_31_424971 147 33 360 360 CD 10_1101-2020_12_31_424971 147 34 ‘ ' `` 10_1101-2020_12_31_424971 147 35 opening opening NN 10_1101-2020_12_31_424971 147 36 ’ ' '' 10_1101-2020_12_31_424971 147 37 and and CC 10_1101-2020_12_31_424971 147 38 ‘ ' `` 10_1101-2020_12_31_424971 147 39 closing close VBG 10_1101-2020_12_31_424971 147 40 ’ ' '' 10_1101-2020_12_31_424971 147 41 of of IN 10_1101-2020_12_31_424971 147 42 the the DT 10_1101-2020_12_31_424971 147 43 L1 L1 NNP 10_1101-2020_12_31_424971 147 44 stalk stalk NN 10_1101-2020_12_31_424971 147 45 of of IN 10_1101-2020_12_31_424971 147 46 the the DT 10_1101-2020_12_31_424971 147 47 50S 50S NNP 10_1101-2020_12_31_424971 147 48 subunit subunit NN 10_1101-2020_12_31_424971 147 49 . . . 10_1101-2020_12_31_424971 148 1 Accordingly accordingly RB 10_1101-2020_12_31_424971 148 2 , , , 10_1101-2020_12_31_424971 148 3 individual individual JJ 10_1101-2020_12_31_424971 148 4 FRET FRET NNP 10_1101-2020_12_31_424971 148 5 efficiency efficiency NN 10_1101-2020_12_31_424971 148 6 361 361 CD 10_1101-2020_12_31_424971 148 7 ( ( -LRB- 10_1101-2020_12_31_424971 148 8 EFRET EFRET NNP 10_1101-2020_12_31_424971 148 9 ) ) -RRB- 10_1101-2020_12_31_424971 148 10 vs. vs. FW 10_1101-2020_12_31_424971 148 11 time time NN 10_1101-2020_12_31_424971 148 12 trajectories trajectory NNS 10_1101-2020_12_31_424971 148 13 recorded record VBN 10_1101-2020_12_31_424971 148 14 using use VBG 10_1101-2020_12_31_424971 148 15 this this DT 10_1101-2020_12_31_424971 148 16 signal signal NN 10_1101-2020_12_31_424971 148 17 exhibit exhibit NN 10_1101-2020_12_31_424971 148 18 transitions transition NNS 10_1101-2020_12_31_424971 148 19 between between IN 10_1101-2020_12_31_424971 148 20 two two CD 10_1101-2020_12_31_424971 148 21 FRET FRET NNP 10_1101-2020_12_31_424971 148 22 states state NNS 10_1101-2020_12_31_424971 148 23 362 362 CD 10_1101-2020_12_31_424971 148 24 corresponding correspond VBG 10_1101-2020_12_31_424971 148 25 to to IN 10_1101-2020_12_31_424971 148 26 the the DT 10_1101-2020_12_31_424971 148 27 ‘ ' `` 10_1101-2020_12_31_424971 148 28 open open JJ 10_1101-2020_12_31_424971 148 29 ’ ' '' 10_1101-2020_12_31_424971 148 30 ( ( -LRB- 10_1101-2020_12_31_424971 148 31 EFRET EFRET NNP 10_1101-2020_12_31_424971 148 32 = = SYM 10_1101-2020_12_31_424971 148 33 ~0.55 ~0.55 NFP 10_1101-2020_12_31_424971 148 34 ) ) -RRB- 10_1101-2020_12_31_424971 148 35 and and CC 10_1101-2020_12_31_424971 148 36 ‘ ' `` 10_1101-2020_12_31_424971 148 37 closed close VBN 10_1101-2020_12_31_424971 148 38 ’ ' '' 10_1101-2020_12_31_424971 148 39 ( ( -LRB- 10_1101-2020_12_31_424971 148 40 EFRET EFRET NNP 10_1101-2020_12_31_424971 148 41 = = NFP 10_1101-2020_12_31_424971 148 42 ~0.31 ~0.31 NFP 10_1101-2020_12_31_424971 148 43 ) ) -RRB- 10_1101-2020_12_31_424971 148 44 conformations conformation NNS 10_1101-2020_12_31_424971 148 45 of of IN 10_1101-2020_12_31_424971 148 46 the the DT 10_1101-2020_12_31_424971 148 47 L1 L1 NNP 10_1101-2020_12_31_424971 148 48 363 363 CD 10_1101-2020_12_31_424971 148 49 stalk stalk NN 10_1101-2020_12_31_424971 148 50 ( ( -LRB- 10_1101-2020_12_31_424971 148 51 Figure Figure NNP 10_1101-2020_12_31_424971 148 52 6d 6d NNP 10_1101-2020_12_31_424971 148 53 ) ) -RRB- 10_1101-2020_12_31_424971 148 54 . . . 10_1101-2020_12_31_424971 149 1 364 364 CD 10_1101-2020_12_31_424971 149 2 Previously previously RB 10_1101-2020_12_31_424971 149 3 , , , 10_1101-2020_12_31_424971 149 4 we -PRON- PRP 10_1101-2020_12_31_424971 149 5 have have VBP 10_1101-2020_12_31_424971 149 6 shown show VBN 10_1101-2020_12_31_424971 149 7 that that IN 10_1101-2020_12_31_424971 149 8 open→closed open→closed JJ 10_1101-2020_12_31_424971 149 9 and and CC 10_1101-2020_12_31_424971 149 10 closed→open closed→open NNP 10_1101-2020_12_31_424971 149 11 L1 L1 NNP 10_1101-2020_12_31_424971 149 12 stalk stalk NN 10_1101-2020_12_31_424971 149 13 transitions transition NNS 10_1101-2020_12_31_424971 149 14 correlate correlate VBP 10_1101-2020_12_31_424971 149 15 365 365 CD 10_1101-2020_12_31_424971 149 16 with with IN 10_1101-2020_12_31_424971 149 17 a a DT 10_1101-2020_12_31_424971 149 18 complex complex JJ 10_1101-2020_12_31_424971 149 19 series series NN 10_1101-2020_12_31_424971 149 20 of of IN 10_1101-2020_12_31_424971 149 21 conformational conformational JJ 10_1101-2020_12_31_424971 149 22 changes change NNS 10_1101-2020_12_31_424971 149 23 that that WDT 10_1101-2020_12_31_424971 149 24 take take VBP 10_1101-2020_12_31_424971 149 25 place place NN 10_1101-2020_12_31_424971 149 26 during during IN 10_1101-2020_12_31_424971 149 27 an an DT 10_1101-2020_12_31_424971 149 28 elongation elongation NN 10_1101-2020_12_31_424971 149 29 cycle35 cycle35 NNP 10_1101-2020_12_31_424971 149 30 - - SYM 10_1101-2020_12_31_424971 149 31 37 37 CD 10_1101-2020_12_31_424971 149 32 . . . 10_1101-2020_12_31_424971 150 1 366 366 CD 10_1101-2020_12_31_424971 150 2 The the DT 10_1101-2020_12_31_424971 150 3 L1 L1 NNP 10_1101-2020_12_31_424971 150 4 stalk stalk NN 10_1101-2020_12_31_424971 150 5 initially initially RB 10_1101-2020_12_31_424971 150 6 occupies occupy VBZ 10_1101-2020_12_31_424971 150 7 the the DT 10_1101-2020_12_31_424971 150 8 open open JJ 10_1101-2020_12_31_424971 150 9 conformation conformation NN 10_1101-2020_12_31_424971 150 10 as as IN 10_1101-2020_12_31_424971 150 11 an an DT 10_1101-2020_12_31_424971 150 12 aa aa NNP 10_1101-2020_12_31_424971 150 13 - - HYPH 10_1101-2020_12_31_424971 150 14 tRNA tRNA NNP 10_1101-2020_12_31_424971 150 15 is be VBZ 10_1101-2020_12_31_424971 150 16 delivered deliver VBN 10_1101-2020_12_31_424971 150 17 to to IN 10_1101-2020_12_31_424971 150 18 the the DT 10_1101-2020_12_31_424971 150 19 A A NNP 10_1101-2020_12_31_424971 150 20 site site NN 10_1101-2020_12_31_424971 150 21 of of IN 10_1101-2020_12_31_424971 150 22 367 367 CD 10_1101-2020_12_31_424971 150 23 a a DT 10_1101-2020_12_31_424971 150 24 70S 70S NNP 10_1101-2020_12_31_424971 150 25 IC IC NNP 10_1101-2020_12_31_424971 150 26 or or CC 10_1101-2020_12_31_424971 150 27 POST POST NNP 10_1101-2020_12_31_424971 150 28 complex complex JJ 10_1101-2020_12_31_424971 150 29 and and CC 10_1101-2020_12_31_424971 150 30 peptide peptide NN 10_1101-2020_12_31_424971 150 31 - - HYPH 10_1101-2020_12_31_424971 150 32 bond bond NN 10_1101-2020_12_31_424971 150 33 formation formation NN 10_1101-2020_12_31_424971 150 34 generates generate VBZ 10_1101-2020_12_31_424971 150 35 a a DT 10_1101-2020_12_31_424971 150 36 PRE PRE NNP 10_1101-2020_12_31_424971 150 37 complex complex NN 10_1101-2020_12_31_424971 150 38 that that WDT 10_1101-2020_12_31_424971 150 39 is be VBZ 10_1101-2020_12_31_424971 150 40 in in IN 10_1101-2020_12_31_424971 150 41 a a DT 10_1101-2020_12_31_424971 150 42 368 368 CD 10_1101-2020_12_31_424971 150 43 global global JJ 10_1101-2020_12_31_424971 150 44 conformation conformation NN 10_1101-2020_12_31_424971 150 45 we -PRON- PRP 10_1101-2020_12_31_424971 150 46 refer refer VBP 10_1101-2020_12_31_424971 150 47 to to IN 10_1101-2020_12_31_424971 150 48 as as IN 10_1101-2020_12_31_424971 150 49 global global JJ 10_1101-2020_12_31_424971 150 50 state state NN 10_1101-2020_12_31_424971 150 51 ( ( -LRB- 10_1101-2020_12_31_424971 150 52 GS GS NNP 10_1101-2020_12_31_424971 150 53 ) ) -RRB- 10_1101-2020_12_31_424971 150 54 1 1 CD 10_1101-2020_12_31_424971 150 55 . . . 10_1101-2020_12_31_424971 151 1 The the DT 10_1101-2020_12_31_424971 151 2 PRE PRE NNP 10_1101-2020_12_31_424971 151 3 complex complex NN 10_1101-2020_12_31_424971 151 4 then then RB 10_1101-2020_12_31_424971 151 5 undergoes undergo VBZ 10_1101-2020_12_31_424971 151 6 a a DT 10_1101-2020_12_31_424971 151 7 large-369 large-369 NNP 10_1101-2020_12_31_424971 151 8 scale scale NN 10_1101-2020_12_31_424971 151 9 structural structural JJ 10_1101-2020_12_31_424971 151 10 rearrangement rearrangement NN 10_1101-2020_12_31_424971 151 11 that that WDT 10_1101-2020_12_31_424971 151 12 includes include VBZ 10_1101-2020_12_31_424971 151 13 an an DT 10_1101-2020_12_31_424971 151 14 open→closed open→closed JJ 10_1101-2020_12_31_424971 151 15 transition transition NN 10_1101-2020_12_31_424971 151 16 of of IN 10_1101-2020_12_31_424971 151 17 the the DT 10_1101-2020_12_31_424971 151 18 L1 L1 NNP 10_1101-2020_12_31_424971 151 19 stalk stalk NN 10_1101-2020_12_31_424971 151 20 so so IN 10_1101-2020_12_31_424971 151 21 as as IN 10_1101-2020_12_31_424971 151 22 to to TO 10_1101-2020_12_31_424971 151 23 370 370 CD 10_1101-2020_12_31_424971 151 24 occupy occupy VB 10_1101-2020_12_31_424971 151 25 a a DT 10_1101-2020_12_31_424971 151 26 second second JJ 10_1101-2020_12_31_424971 151 27 global global JJ 10_1101-2020_12_31_424971 151 28 conformation conformation NN 10_1101-2020_12_31_424971 151 29 we -PRON- PRP 10_1101-2020_12_31_424971 151 30 refer refer VBP 10_1101-2020_12_31_424971 151 31 to to IN 10_1101-2020_12_31_424971 151 32 as as IN 10_1101-2020_12_31_424971 151 33 GS2 GS2 NNP 10_1101-2020_12_31_424971 151 34 ( ( -LRB- 10_1101-2020_12_31_424971 151 35 i.e. i.e. FW 10_1101-2020_12_31_424971 151 36 , , , 10_1101-2020_12_31_424971 151 37 the the DT 10_1101-2020_12_31_424971 151 38 0.55→0.31 0.55→0.31 CD 10_1101-2020_12_31_424971 151 39 EFRET EFRET NNP 10_1101-2020_12_31_424971 151 40 transition transition NN 10_1101-2020_12_31_424971 151 41 371 371 CD 10_1101-2020_12_31_424971 151 42 denoted denote VBN 10_1101-2020_12_31_424971 151 43 by by IN 10_1101-2020_12_31_424971 151 44 the the DT 10_1101-2020_12_31_424971 151 45 rate rate NN 10_1101-2020_12_31_424971 151 46 k70S k70s ADD 10_1101-2020_12_31_424971 151 47 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 151 48 in in IN 10_1101-2020_12_31_424971 151 49 Figures Figures NNP 10_1101-2020_12_31_424971 151 50 6d 6d NNP 10_1101-2020_12_31_424971 151 51 and and CC 10_1101-2020_12_31_424971 151 52 e e NNP 10_1101-2020_12_31_424971 151 53 , , , 10_1101-2020_12_31_424971 151 54 corresponding correspond VBG 10_1101-2020_12_31_424971 151 55 to to IN 10_1101-2020_12_31_424971 151 56 the the DT 10_1101-2020_12_31_424971 151 57 multi multi JJ 10_1101-2020_12_31_424971 151 58 - - JJ 10_1101-2020_12_31_424971 151 59 step step NN 10_1101-2020_12_31_424971 151 60 70S 70S NNP 10_1101-2020_12_31_424971 151 61 IC→GS2 ic→gs2 NN 10_1101-2020_12_31_424971 151 62 372 372 CD 10_1101-2020_12_31_424971 151 63 transition transition NN 10_1101-2020_12_31_424971 151 64 in in IN 10_1101-2020_12_31_424971 151 65 Figure Figure NNP 10_1101-2020_12_31_424971 151 66 6a 6a NNP 10_1101-2020_12_31_424971 151 67 ) ) -RRB- 10_1101-2020_12_31_424971 151 68 . . . 10_1101-2020_12_31_424971 152 1 Subsequently subsequently RB 10_1101-2020_12_31_424971 152 2 , , , 10_1101-2020_12_31_424971 152 3 in in IN 10_1101-2020_12_31_424971 152 4 the the DT 10_1101-2020_12_31_424971 152 5 absence absence NN 10_1101-2020_12_31_424971 152 6 of of IN 10_1101-2020_12_31_424971 152 7 EF EF NNP 10_1101-2020_12_31_424971 152 8 - - HYPH 10_1101-2020_12_31_424971 152 9 G G NNP 10_1101-2020_12_31_424971 152 10 , , , 10_1101-2020_12_31_424971 152 11 the the DT 10_1101-2020_12_31_424971 152 12 L1 L1 NNP 10_1101-2020_12_31_424971 152 13 stalk stalk NN 10_1101-2020_12_31_424971 152 14 goes go VBZ 10_1101-2020_12_31_424971 152 15 through through IN 10_1101-2020_12_31_424971 152 16 373 373 CD 10_1101-2020_12_31_424971 152 17 successive successive JJ 10_1101-2020_12_31_424971 152 18 closed→open closed→open NN 10_1101-2020_12_31_424971 152 19 and and CC 10_1101-2020_12_31_424971 152 20 open→closed open→closed JJ 10_1101-2020_12_31_424971 152 21 transitions transition NNS 10_1101-2020_12_31_424971 152 22 as as IN 10_1101-2020_12_31_424971 152 23 the the DT 10_1101-2020_12_31_424971 152 24 PRE PRE NNP 10_1101-2020_12_31_424971 152 25 complex complex JJ 10_1101-2020_12_31_424971 152 26 undergoes undergoe NNS 10_1101-2020_12_31_424971 152 27 multiple multiple NN 10_1101-2020_12_31_424971 152 28 374 374 CD 10_1101-2020_12_31_424971 152 29 GS2→GS1 GS2→GS1 NNP 10_1101-2020_12_31_424971 152 30 and and CC 10_1101-2020_12_31_424971 152 31 GS1→GS2 GS1→GS2 NNP 10_1101-2020_12_31_424971 152 32 transitions transition NNS 10_1101-2020_12_31_424971 152 33 that that WDT 10_1101-2020_12_31_424971 152 34 establish establish VBP 10_1101-2020_12_31_424971 152 35 a a DT 10_1101-2020_12_31_424971 152 36 GS1 GS1 NNP 10_1101-2020_12_31_424971 152 37 ⇄ ⇄ NNP 10_1101-2020_12_31_424971 152 38 GS2 GS2 NNP 10_1101-2020_12_31_424971 152 39 equilibrium equilibrium NN 10_1101-2020_12_31_424971 152 40 ( ( -LRB- 10_1101-2020_12_31_424971 152 41 i.e. i.e. FW 10_1101-2020_12_31_424971 152 42 , , , 10_1101-2020_12_31_424971 152 43 the the DT 10_1101-2020_12_31_424971 152 44 0.55 0.55 CD 10_1101-2020_12_31_424971 152 45 ⇄ ⇄ NNP 10_1101-2020_12_31_424971 152 46 0.31 0.31 CD 10_1101-2020_12_31_424971 152 47 375 375 CD 10_1101-2020_12_31_424971 152 48 EFRET efret NN 10_1101-2020_12_31_424971 152 49 transitions transition NNS 10_1101-2020_12_31_424971 152 50 denoted denote VBN 10_1101-2020_12_31_424971 152 51 by by IN 10_1101-2020_12_31_424971 152 52 the the DT 10_1101-2020_12_31_424971 152 53 rates rate NNS 10_1101-2020_12_31_424971 152 54 kGS1→GS2 kGS1→GS2 NNP 10_1101-2020_12_31_424971 152 55 and and CC 10_1101-2020_12_31_424971 152 56 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 152 57 and and CC 10_1101-2020_12_31_424971 152 58 the the DT 10_1101-2020_12_31_424971 152 59 equilibrium equilibrium NN 10_1101-2020_12_31_424971 152 60 constant constant JJ 10_1101-2020_12_31_424971 152 61 Keq Keq NNP 10_1101-2020_12_31_424971 152 62 = = SYM 10_1101-2020_12_31_424971 152 63 376 376 CD 10_1101-2020_12_31_424971 152 64 ( ( -LRB- 10_1101-2020_12_31_424971 152 65 kGS1→GS2)/(kGS2→GS1 kGS1→GS2)/(kGS2→GS1 NNP 10_1101-2020_12_31_424971 152 66 ) ) -RRB- 10_1101-2020_12_31_424971 152 67 in in IN 10_1101-2020_12_31_424971 152 68 Figure Figure NNP 10_1101-2020_12_31_424971 152 69 6d 6d NNP 10_1101-2020_12_31_424971 152 70 , , , 10_1101-2020_12_31_424971 152 71 corresponding correspond VBG 10_1101-2020_12_31_424971 152 72 to to IN 10_1101-2020_12_31_424971 152 73 the the DT 10_1101-2020_12_31_424971 152 74 GS1 GS1 NNP 10_1101-2020_12_31_424971 152 75 ⇄ ⇄ NNP 10_1101-2020_12_31_424971 152 76 GS2 GS2 NNP 10_1101-2020_12_31_424971 152 77 transitions transition NNS 10_1101-2020_12_31_424971 152 78 in in IN 10_1101-2020_12_31_424971 152 79 Figure Figure NNP 10_1101-2020_12_31_424971 152 80 6a 6a NNP 10_1101-2020_12_31_424971 152 81 ) ) -RRB- 10_1101-2020_12_31_424971 152 82 . . . 10_1101-2020_12_31_424971 153 1 In in IN 10_1101-2020_12_31_424971 153 2 the the DT 10_1101-2020_12_31_424971 153 3 377 377 CD 10_1101-2020_12_31_424971 153 4 presence presence NN 10_1101-2020_12_31_424971 153 5 of of IN 10_1101-2020_12_31_424971 153 6 EF EF NNP 10_1101-2020_12_31_424971 153 7 - - HYPH 10_1101-2020_12_31_424971 153 8 G G NNP 10_1101-2020_12_31_424971 153 9 , , , 10_1101-2020_12_31_424971 153 10 however however RB 10_1101-2020_12_31_424971 153 11 , , , 10_1101-2020_12_31_424971 153 12 a a DT 10_1101-2020_12_31_424971 153 13 single single JJ 10_1101-2020_12_31_424971 153 14 closed→open closed→open NNP 10_1101-2020_12_31_424971 153 15 L1 L1 NNP 10_1101-2020_12_31_424971 153 16 stalk stalk NN 10_1101-2020_12_31_424971 153 17 transition transition NN 10_1101-2020_12_31_424971 153 18 reports report VBZ 10_1101-2020_12_31_424971 153 19 on on IN 10_1101-2020_12_31_424971 153 20 conformational conformational JJ 10_1101-2020_12_31_424971 153 21 378 378 CD 10_1101-2020_12_31_424971 153 22 changes change NNS 10_1101-2020_12_31_424971 153 23 of of IN 10_1101-2020_12_31_424971 153 24 the the DT 10_1101-2020_12_31_424971 153 25 PRE PRE NNP 10_1101-2020_12_31_424971 153 26 complex complex NN 10_1101-2020_12_31_424971 153 27 as as IN 10_1101-2020_12_31_424971 153 28 it -PRON- PRP 10_1101-2020_12_31_424971 153 29 undergoes undergo VBZ 10_1101-2020_12_31_424971 153 30 EF EF NNP 10_1101-2020_12_31_424971 153 31 - - HYPH 10_1101-2020_12_31_424971 153 32 G g NN 10_1101-2020_12_31_424971 153 33 binding binding NN 10_1101-2020_12_31_424971 153 34 and and CC 10_1101-2020_12_31_424971 153 35 completes complete VBZ 10_1101-2020_12_31_424971 153 36 translocation translocation NN 10_1101-2020_12_31_424971 153 37 ( ( -LRB- 10_1101-2020_12_31_424971 153 38 i.e. i.e. FW 10_1101-2020_12_31_424971 153 39 , , , 10_1101-2020_12_31_424971 153 40 the the DT 10_1101-2020_12_31_424971 153 41 379 379 CD 10_1101-2020_12_31_424971 153 42 0.31→0.55 0.31→0.55 NNS 10_1101-2020_12_31_424971 153 43 EFRET EFRET NNP 10_1101-2020_12_31_424971 153 44 transition transition NN 10_1101-2020_12_31_424971 153 45 denoted denote VBN 10_1101-2020_12_31_424971 153 46 by by IN 10_1101-2020_12_31_424971 153 47 the the DT 10_1101-2020_12_31_424971 153 48 rate rate NN 10_1101-2020_12_31_424971 153 49 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 153 50 in in IN 10_1101-2020_12_31_424971 153 51 Figures Figures NNP 10_1101-2020_12_31_424971 153 52 6d 6d NNP 10_1101-2020_12_31_424971 153 53 and and CC 10_1101-2020_12_31_424971 153 54 e e NNP 10_1101-2020_12_31_424971 153 55 , , , 10_1101-2020_12_31_424971 153 56 corresponding correspond VBG 10_1101-2020_12_31_424971 153 57 to to IN 10_1101-2020_12_31_424971 153 58 380 380 CD 10_1101-2020_12_31_424971 153 59 the the DT 10_1101-2020_12_31_424971 153 60 multi multi JJ 10_1101-2020_12_31_424971 153 61 - - JJ 10_1101-2020_12_31_424971 153 62 step step JJ 10_1101-2020_12_31_424971 153 63 GS2→POST gs2→post NN 10_1101-2020_12_31_424971 153 64 transition transition NN 10_1101-2020_12_31_424971 153 65 that that WDT 10_1101-2020_12_31_424971 153 66 takes take VBZ 10_1101-2020_12_31_424971 153 67 place place NN 10_1101-2020_12_31_424971 153 68 in in IN 10_1101-2020_12_31_424971 153 69 the the DT 10_1101-2020_12_31_424971 153 70 presence presence NN 10_1101-2020_12_31_424971 153 71 of of IN 10_1101-2020_12_31_424971 153 72 EF EF NNP 10_1101-2020_12_31_424971 153 73 - - HYPH 10_1101-2020_12_31_424971 153 74 G G NNP 10_1101-2020_12_31_424971 153 75 and and CC 10_1101-2020_12_31_424971 153 76 bridges bridge NNS 10_1101-2020_12_31_424971 153 77 across across IN 10_1101-2020_12_31_424971 153 78 381 381 CD 10_1101-2020_12_31_424971 153 79 Figures Figures NNPS 10_1101-2020_12_31_424971 153 80 6a 6a CD 10_1101-2020_12_31_424971 153 81 and and CC 10_1101-2020_12_31_424971 153 82 b b NNP 10_1101-2020_12_31_424971 153 83 ) ) -RRB- 10_1101-2020_12_31_424971 153 84 . . . 10_1101-2020_12_31_424971 154 1 Using use VBG 10_1101-2020_12_31_424971 154 2 this this DT 10_1101-2020_12_31_424971 154 3 approach approach NN 10_1101-2020_12_31_424971 154 4 , , , 10_1101-2020_12_31_424971 154 5 we -PRON- PRP 10_1101-2020_12_31_424971 154 6 have have VBP 10_1101-2020_12_31_424971 154 7 successfully successfully RB 10_1101-2020_12_31_424971 154 8 monitored monitor VBN 10_1101-2020_12_31_424971 154 9 the the DT 10_1101-2020_12_31_424971 154 10 conformational conformational JJ 10_1101-2020_12_31_424971 154 11 382 382 CD 10_1101-2020_12_31_424971 154 12 .CC .CC , 10_1101-2020_12_31_424971 154 13 - - HYPH 10_1101-2020_12_31_424971 154 14 BY by IN 10_1101-2020_12_31_424971 154 15 - - HYPH 10_1101-2020_12_31_424971 154 16 NC NC NNP 10_1101-2020_12_31_424971 154 17 - - HYPH 10_1101-2020_12_31_424971 154 18 ND ND NNP 10_1101-2020_12_31_424971 154 19 4.0 4.0 CD 10_1101-2020_12_31_424971 154 20 International International NNP 10_1101-2020_12_31_424971 154 21 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 154 22 under under IN 10_1101-2020_12_31_424971 154 23 a a DT 10_1101-2020_12_31_424971 154 24 ( ( -LRB- 10_1101-2020_12_31_424971 154 25 which which WDT 10_1101-2020_12_31_424971 154 26 was be VBD 10_1101-2020_12_31_424971 154 27 not not RB 10_1101-2020_12_31_424971 154 28 certified certify VBN 10_1101-2020_12_31_424971 154 29 by by IN 10_1101-2020_12_31_424971 154 30 peer peer NN 10_1101-2020_12_31_424971 154 31 review review NN 10_1101-2020_12_31_424971 154 32 ) ) -RRB- 10_1101-2020_12_31_424971 154 33 is be VBZ 10_1101-2020_12_31_424971 154 34 the the DT 10_1101-2020_12_31_424971 154 35 author author NN 10_1101-2020_12_31_424971 154 36 / / SYM 10_1101-2020_12_31_424971 154 37 funder funder NN 10_1101-2020_12_31_424971 154 38 , , , 10_1101-2020_12_31_424971 154 39 who who WP 10_1101-2020_12_31_424971 154 40 has have VBZ 10_1101-2020_12_31_424971 154 41 granted grant VBN 10_1101-2020_12_31_424971 154 42 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 154 43 a a DT 10_1101-2020_12_31_424971 154 44 license license NN 10_1101-2020_12_31_424971 154 45 to to TO 10_1101-2020_12_31_424971 154 46 display display VB 10_1101-2020_12_31_424971 154 47 the the DT 10_1101-2020_12_31_424971 154 48 preprint preprint NN 10_1101-2020_12_31_424971 154 49 in in IN 10_1101-2020_12_31_424971 154 50 perpetuity perpetuity NN 10_1101-2020_12_31_424971 154 51 . . . 10_1101-2020_12_31_424971 155 1 It -PRON- PRP 10_1101-2020_12_31_424971 155 2 is be VBZ 10_1101-2020_12_31_424971 155 3 made make VBN 10_1101-2020_12_31_424971 155 4 The the DT 10_1101-2020_12_31_424971 155 5 copyright copyright NN 10_1101-2020_12_31_424971 155 6 holder holder NN 10_1101-2020_12_31_424971 155 7 for for IN 10_1101-2020_12_31_424971 155 8 this this DT 10_1101-2020_12_31_424971 155 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 155 10 version version NN 10_1101-2020_12_31_424971 155 11 posted post VBD 10_1101-2020_12_31_424971 155 12 January January NNP 10_1101-2020_12_31_424971 155 13 2 2 CD 10_1101-2020_12_31_424971 155 14 , , , 10_1101-2020_12_31_424971 155 15 2021 2021 CD 10_1101-2020_12_31_424971 155 16 . . . 10_1101-2020_12_31_424971 155 17 ; ; : 10_1101-2020_12_31_424971 155 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 155 19 : : : 10_1101-2020_12_31_424971 155 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 155 21 preprint preprint NN 10_1101-2020_12_31_424971 155 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 155 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 155 24 16 16 CD 10_1101-2020_12_31_424971 155 25 dynamics dynamic NNS 10_1101-2020_12_31_424971 155 26 of of IN 10_1101-2020_12_31_424971 155 27 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 155 28 complexes complex NNS 10_1101-2020_12_31_424971 155 29 during during IN 10_1101-2020_12_31_424971 155 30 individual individual JJ 10_1101-2020_12_31_424971 155 31 elongation elongation NN 10_1101-2020_12_31_424971 155 32 cycles36,38 cycles36,38 NNP 10_1101-2020_12_31_424971 155 33 - - HYPH 10_1101-2020_12_31_424971 155 34 41 41 CD 10_1101-2020_12_31_424971 155 35 , , , 10_1101-2020_12_31_424971 155 36 including include VBG 10_1101-2020_12_31_424971 155 37 in in IN 10_1101-2020_12_31_424971 155 38 a a DT 10_1101-2020_12_31_424971 155 39 study study NN 10_1101-2020_12_31_424971 155 40 383 383 CD 10_1101-2020_12_31_424971 155 41 of of IN 10_1101-2020_12_31_424971 155 42 – – : 10_1101-2020_12_31_424971 155 43 1 1 CD 10_1101-2020_12_31_424971 155 44 frameshifting41 frameshifting41 CD 10_1101-2020_12_31_424971 155 45 . . . 10_1101-2020_12_31_424971 156 1 384 384 CD 10_1101-2020_12_31_424971 156 2 385 385 CD 10_1101-2020_12_31_424971 156 3 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 156 4 interferes interfere VBZ 10_1101-2020_12_31_424971 156 5 with with IN 10_1101-2020_12_31_424971 156 6 elongation elongation NN 10_1101-2020_12_31_424971 156 7 complex complex JJ 10_1101-2020_12_31_424971 156 8 dynamics dynamic NNS 10_1101-2020_12_31_424971 156 9 during during IN 10_1101-2020_12_31_424971 156 10 late late JJ 10_1101-2020_12_31_424971 156 11 steps step NNS 10_1101-2020_12_31_424971 156 12 in in IN 10_1101-2020_12_31_424971 156 13 translocation translocation NN 10_1101-2020_12_31_424971 156 14 386 386 CD 10_1101-2020_12_31_424971 156 15 We -PRON- PRP 10_1101-2020_12_31_424971 156 16 began begin VBD 10_1101-2020_12_31_424971 156 17 by by IN 10_1101-2020_12_31_424971 156 18 asking ask VBG 10_1101-2020_12_31_424971 156 19 whether whether IN 10_1101-2020_12_31_424971 156 20 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 156 21 alters alter VBZ 10_1101-2020_12_31_424971 156 22 the the DT 10_1101-2020_12_31_424971 156 23 dynamics dynamic NNS 10_1101-2020_12_31_424971 156 24 of of IN 10_1101-2020_12_31_424971 156 25 elongation elongation NN 10_1101-2020_12_31_424971 156 26 complexes complex NNS 10_1101-2020_12_31_424971 156 27 during during IN 10_1101-2020_12_31_424971 156 28 the the DT 10_1101-2020_12_31_424971 156 29 387 387 CD 10_1101-2020_12_31_424971 156 30 earlier early JJR 10_1101-2020_12_31_424971 156 31 steps step NNS 10_1101-2020_12_31_424971 156 32 of of IN 10_1101-2020_12_31_424971 156 33 the the DT 10_1101-2020_12_31_424971 156 34 elongation elongation NN 10_1101-2020_12_31_424971 156 35 cycle cycle NN 10_1101-2020_12_31_424971 156 36 . . . 10_1101-2020_12_31_424971 157 1 We -PRON- PRP 10_1101-2020_12_31_424971 157 2 stopped stop VBD 10_1101-2020_12_31_424971 157 3 - - HYPH 10_1101-2020_12_31_424971 157 4 flow flow NN 10_1101-2020_12_31_424971 157 5 delivered deliver VBN 10_1101-2020_12_31_424971 157 6 SufB2- SufB2- NNP 10_1101-2020_12_31_424971 157 7 or or CC 10_1101-2020_12_31_424971 157 8 ProL ProL NNP 10_1101-2020_12_31_424971 157 9 - - HYPH 10_1101-2020_12_31_424971 157 10 TC TC NNP 10_1101-2020_12_31_424971 157 11 to to IN 10_1101-2020_12_31_424971 157 12 70S 70S NNP 10_1101-2020_12_31_424971 157 13 ICs ICs NNP 10_1101-2020_12_31_424971 157 14 388 388 CD 10_1101-2020_12_31_424971 157 15 and and CC 10_1101-2020_12_31_424971 157 16 recorded record VBD 10_1101-2020_12_31_424971 157 17 pre pre JJ 10_1101-2020_12_31_424971 157 18 - - JJ 10_1101-2020_12_31_424971 157 19 steady steady JJ 10_1101-2020_12_31_424971 157 20 - - HYPH 10_1101-2020_12_31_424971 157 21 state state NN 10_1101-2020_12_31_424971 157 22 movies movie NNS 10_1101-2020_12_31_424971 157 23 during during IN 10_1101-2020_12_31_424971 157 24 delivery delivery NN 10_1101-2020_12_31_424971 157 25 , , , 10_1101-2020_12_31_424971 157 26 and and CC 10_1101-2020_12_31_424971 157 27 steady steady JJ 10_1101-2020_12_31_424971 157 28 - - HYPH 10_1101-2020_12_31_424971 157 29 state state NN 10_1101-2020_12_31_424971 157 30 movies movie NNS 10_1101-2020_12_31_424971 157 31 1 1 CD 10_1101-2020_12_31_424971 157 32 min min NN 10_1101-2020_12_31_424971 157 33 post-389 post-389 NNP 10_1101-2020_12_31_424971 157 34 delivery delivery NN 10_1101-2020_12_31_424971 157 35 ( ( -LRB- 10_1101-2020_12_31_424971 157 36 Figures Figures NNP 10_1101-2020_12_31_424971 157 37 6a 6a NNP 10_1101-2020_12_31_424971 157 38 , , , 10_1101-2020_12_31_424971 157 39 d d NN 10_1101-2020_12_31_424971 157 40 , , , 10_1101-2020_12_31_424971 157 41 and and CC 10_1101-2020_12_31_424971 157 42 f f NNP 10_1101-2020_12_31_424971 157 43 , , , 10_1101-2020_12_31_424971 157 44 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 157 45 Figures Figures NNP 10_1101-2020_12_31_424971 157 46 3 3 CD 10_1101-2020_12_31_424971 157 47 , , , 10_1101-2020_12_31_424971 157 48 4a 4a CD 10_1101-2020_12_31_424971 157 49 , , , 10_1101-2020_12_31_424971 157 50 and and CC 10_1101-2020_12_31_424971 157 51 4b 4b NNP 10_1101-2020_12_31_424971 157 52 ) ) -RRB- 10_1101-2020_12_31_424971 157 53 . . . 10_1101-2020_12_31_424971 158 1 The the DT 10_1101-2020_12_31_424971 158 2 results result NNS 10_1101-2020_12_31_424971 158 3 showed show VBD 10_1101-2020_12_31_424971 158 4 that that IN 10_1101-2020_12_31_424971 158 5 k70S k70s RB 10_1101-2020_12_31_424971 158 6 390 390 CD 10_1101-2020_12_31_424971 158 7 IC→GS2 ic→gs2 NN 10_1101-2020_12_31_424971 158 8 , , , 10_1101-2020_12_31_424971 158 9 as as RB 10_1101-2020_12_31_424971 158 10 well well RB 10_1101-2020_12_31_424971 158 11 as as IN 10_1101-2020_12_31_424971 158 12 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 158 13 , , , 10_1101-2020_12_31_424971 158 14 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 158 15 , , , 10_1101-2020_12_31_424971 158 16 and and CC 10_1101-2020_12_31_424971 158 17 Keq Keq NNP 10_1101-2020_12_31_424971 158 18 at at IN 10_1101-2020_12_31_424971 158 19 1 1 CD 10_1101-2020_12_31_424971 158 20 min min NN 10_1101-2020_12_31_424971 158 21 post post JJ 10_1101-2020_12_31_424971 158 22 - - JJ 10_1101-2020_12_31_424971 158 23 delivery delivery NN 10_1101-2020_12_31_424971 158 24 , , , 10_1101-2020_12_31_424971 158 25 for for IN 10_1101-2020_12_31_424971 158 26 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 158 27 were be VBD 10_1101-2020_12_31_424971 158 28 each each DT 10_1101-2020_12_31_424971 158 29 less less JJR 10_1101-2020_12_31_424971 158 30 391 391 CD 10_1101-2020_12_31_424971 158 31 than than IN 10_1101-2020_12_31_424971 158 32 2-fold 2-fold CD 10_1101-2020_12_31_424971 158 33 different different JJ 10_1101-2020_12_31_424971 158 34 than than IN 10_1101-2020_12_31_424971 158 35 the the DT 10_1101-2020_12_31_424971 158 36 corresponding corresponding JJ 10_1101-2020_12_31_424971 158 37 value value NN 10_1101-2020_12_31_424971 158 38 for for IN 10_1101-2020_12_31_424971 158 39 ProL ProL NNP 10_1101-2020_12_31_424971 158 40 - - HYPH 10_1101-2020_12_31_424971 158 41 TC TC NNP 10_1101-2020_12_31_424971 158 42 ( ( -LRB- 10_1101-2020_12_31_424971 158 43 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 158 44 Table Table NNP 10_1101-2020_12_31_424971 158 45 4 4 CD 10_1101-2020_12_31_424971 158 46 ) ) -RRB- 10_1101-2020_12_31_424971 158 47 . . . 10_1101-2020_12_31_424971 159 1 The the DT 10_1101-2020_12_31_424971 159 2 392 392 CD 10_1101-2020_12_31_424971 159 3 close close JJ 10_1101-2020_12_31_424971 159 4 correspondence correspondence NN 10_1101-2020_12_31_424971 159 5 of of IN 10_1101-2020_12_31_424971 159 6 these these DT 10_1101-2020_12_31_424971 159 7 rates rate NNS 10_1101-2020_12_31_424971 159 8 indicates indicate VBZ 10_1101-2020_12_31_424971 159 9 that that IN 10_1101-2020_12_31_424971 159 10 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 159 11 is be VBZ 10_1101-2020_12_31_424971 159 12 delivered deliver VBN 10_1101-2020_12_31_424971 159 13 to to IN 10_1101-2020_12_31_424971 159 14 the the DT 10_1101-2020_12_31_424971 159 15 A A NNP 10_1101-2020_12_31_424971 159 16 site site NN 10_1101-2020_12_31_424971 159 17 , , , 10_1101-2020_12_31_424971 159 18 393 393 CD 10_1101-2020_12_31_424971 159 19 participates participate NNS 10_1101-2020_12_31_424971 159 20 in in IN 10_1101-2020_12_31_424971 159 21 peptide peptide NN 10_1101-2020_12_31_424971 159 22 - - HYPH 10_1101-2020_12_31_424971 159 23 bond bond NN 10_1101-2020_12_31_424971 159 24 formation formation NN 10_1101-2020_12_31_424971 159 25 , , , 10_1101-2020_12_31_424971 159 26 undergoes undergo VBZ 10_1101-2020_12_31_424971 159 27 GS2 gs2 NN 10_1101-2020_12_31_424971 159 28 formation formation NN 10_1101-2020_12_31_424971 159 29 , , , 10_1101-2020_12_31_424971 159 30 and and CC 10_1101-2020_12_31_424971 159 31 exhibits exhibit VBZ 10_1101-2020_12_31_424971 159 32 GS1→GS2 GS1→GS2 NNP 10_1101-2020_12_31_424971 159 33 and and CC 10_1101-2020_12_31_424971 159 34 394 394 CD 10_1101-2020_12_31_424971 159 35 GS2→GS1 gs2→gs1 NN 10_1101-2020_12_31_424971 159 36 transitions transition NNS 10_1101-2020_12_31_424971 159 37 within within IN 10_1101-2020_12_31_424971 159 38 the the DT 10_1101-2020_12_31_424971 159 39 GS1 GS1 NNP 10_1101-2020_12_31_424971 159 40 ⇄ ⇄ NNP 10_1101-2020_12_31_424971 159 41 GS2 GS2 NNP 10_1101-2020_12_31_424971 159 42 equilibrium equilibrium NN 10_1101-2020_12_31_424971 159 43 in in IN 10_1101-2020_12_31_424971 159 44 a a DT 10_1101-2020_12_31_424971 159 45 manner manner NN 10_1101-2020_12_31_424971 159 46 that that WDT 10_1101-2020_12_31_424971 159 47 is be VBZ 10_1101-2020_12_31_424971 159 48 similar similar JJ 10_1101-2020_12_31_424971 159 49 to to IN 10_1101-2020_12_31_424971 159 50 ProL ProL NNP 10_1101-2020_12_31_424971 159 51 - - HYPH 10_1101-2020_12_31_424971 159 52 TC TC NNP 10_1101-2020_12_31_424971 159 53 , , , 10_1101-2020_12_31_424971 159 54 395 395 CD 10_1101-2020_12_31_424971 159 55 consistent consistent JJ 10_1101-2020_12_31_424971 159 56 with with IN 10_1101-2020_12_31_424971 159 57 the the DT 10_1101-2020_12_31_424971 159 58 results result NNS 10_1101-2020_12_31_424971 159 59 of of IN 10_1101-2020_12_31_424971 159 60 ensemble ensemble JJ 10_1101-2020_12_31_424971 159 61 kinetic kinetic NN 10_1101-2020_12_31_424971 159 62 assays assay NNS 10_1101-2020_12_31_424971 159 63 ( ( -LRB- 10_1101-2020_12_31_424971 159 64 Figures Figures NNP 10_1101-2020_12_31_424971 159 65 3a 3a NNP 10_1101-2020_12_31_424971 159 66 - - HYPH 10_1101-2020_12_31_424971 159 67 c c NNP 10_1101-2020_12_31_424971 159 68 , , , 10_1101-2020_12_31_424971 159 69 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 159 70 Table Table NNP 10_1101-2020_12_31_424971 159 71 2 2 CD 10_1101-2020_12_31_424971 159 72 , , , 10_1101-2020_12_31_424971 159 73 and and CC 10_1101-2020_12_31_424971 159 74 396 396 CD 10_1101-2020_12_31_424971 159 75 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 159 76 Figure Figure NNP 10_1101-2020_12_31_424971 159 77 2a 2a NNP 10_1101-2020_12_31_424971 159 78 ) ) -RRB- 10_1101-2020_12_31_424971 159 79 and and CC 10_1101-2020_12_31_424971 159 80 thereby thereby RB 10_1101-2020_12_31_424971 159 81 strengthening strengthen VBG 10_1101-2020_12_31_424971 159 82 our -PRON- PRP$ 10_1101-2020_12_31_424971 159 83 interpretation interpretation NN 10_1101-2020_12_31_424971 159 84 that that IN 10_1101-2020_12_31_424971 159 85 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 159 86 uses use VBZ 10_1101-2020_12_31_424971 159 87 triplet triplet NN 10_1101-2020_12_31_424971 159 88 397 397 CD 10_1101-2020_12_31_424971 159 89 pairing pair VBG 10_1101-2020_12_31_424971 159 90 in in IN 10_1101-2020_12_31_424971 159 91 the the DT 10_1101-2020_12_31_424971 159 92 0-frame 0-frame CD 10_1101-2020_12_31_424971 159 93 at at IN 10_1101-2020_12_31_424971 159 94 the the DT 10_1101-2020_12_31_424971 159 95 A A NNP 10_1101-2020_12_31_424971 159 96 site site NN 10_1101-2020_12_31_424971 159 97 during during IN 10_1101-2020_12_31_424971 159 98 the the DT 10_1101-2020_12_31_424971 159 99 early early JJ 10_1101-2020_12_31_424971 159 100 stages stage NNS 10_1101-2020_12_31_424971 159 101 of of IN 10_1101-2020_12_31_424971 159 102 the the DT 10_1101-2020_12_31_424971 159 103 elongation elongation NN 10_1101-2020_12_31_424971 159 104 cycle cycle NN 10_1101-2020_12_31_424971 159 105 that that WDT 10_1101-2020_12_31_424971 159 106 precede precede VBP 10_1101-2020_12_31_424971 159 107 398 398 CD 10_1101-2020_12_31_424971 159 108 EF EF NNP 10_1101-2020_12_31_424971 159 109 - - HYPH 10_1101-2020_12_31_424971 159 110 G g NN 10_1101-2020_12_31_424971 159 111 binding binding NN 10_1101-2020_12_31_424971 159 112 and and CC 10_1101-2020_12_31_424971 159 113 EF EF NNP 10_1101-2020_12_31_424971 159 114 - - HYPH 10_1101-2020_12_31_424971 159 115 G G NNP 10_1101-2020_12_31_424971 159 116 - - HYPH 10_1101-2020_12_31_424971 159 117 catalyzed catalyze VBN 10_1101-2020_12_31_424971 159 118 translocation translocation NN 10_1101-2020_12_31_424971 159 119 . . . 10_1101-2020_12_31_424971 160 1 Although although IN 10_1101-2020_12_31_424971 160 2 we -PRON- PRP 10_1101-2020_12_31_424971 160 3 could could MD 10_1101-2020_12_31_424971 160 4 not not RB 10_1101-2020_12_31_424971 160 5 confidently confidently RB 10_1101-2020_12_31_424971 160 6 detect detect VB 10_1101-2020_12_31_424971 160 7 the the DT 10_1101-2020_12_31_424971 160 8 399 399 CD 10_1101-2020_12_31_424971 160 9 presence presence NN 10_1101-2020_12_31_424971 160 10 of of IN 10_1101-2020_12_31_424971 160 11 two two CD 10_1101-2020_12_31_424971 160 12 sub sub NN 10_1101-2020_12_31_424971 160 13 - - NNS 10_1101-2020_12_31_424971 160 14 populations population NNS 10_1101-2020_12_31_424971 160 15 of of IN 10_1101-2020_12_31_424971 160 16 A a DT 10_1101-2020_12_31_424971 160 17 site site NN 10_1101-2020_12_31_424971 160 18 - - HYPH 10_1101-2020_12_31_424971 160 19 bound bind VBN 10_1101-2020_12_31_424971 160 20 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 160 21 in in IN 10_1101-2020_12_31_424971 160 22 the the DT 10_1101-2020_12_31_424971 160 23 smFRET smFRET NNP 10_1101-2020_12_31_424971 160 24 data datum NNS 10_1101-2020_12_31_424971 160 25 that that WDT 10_1101-2020_12_31_424971 160 26 might may MD 10_1101-2020_12_31_424971 160 27 differ differ VB 10_1101-2020_12_31_424971 160 28 in in IN 10_1101-2020_12_31_424971 160 29 400 400 CD 10_1101-2020_12_31_424971 160 30 their -PRON- PRP$ 10_1101-2020_12_31_424971 160 31 propensity propensity NN 10_1101-2020_12_31_424971 160 32 of of IN 10_1101-2020_12_31_424971 160 33 undergoing undergoing JJ 10_1101-2020_12_31_424971 160 34 +1 +1 NNS 10_1101-2020_12_31_424971 160 35 frameshifting frameshifting NN 10_1101-2020_12_31_424971 160 36 , , , 10_1101-2020_12_31_424971 160 37 as as IN 10_1101-2020_12_31_424971 160 38 suggested suggest VBN 10_1101-2020_12_31_424971 160 39 by by IN 10_1101-2020_12_31_424971 160 40 the the DT 10_1101-2020_12_31_424971 160 41 results result NNS 10_1101-2020_12_31_424971 160 42 presented present VBN 10_1101-2020_12_31_424971 160 43 in in IN 10_1101-2020_12_31_424971 160 44 Figure Figure NNP 10_1101-2020_12_31_424971 160 45 401 401 CD 10_1101-2020_12_31_424971 160 46 5a 5a CD 10_1101-2020_12_31_424971 160 47 , , , 10_1101-2020_12_31_424971 160 48 it -PRON- PRP 10_1101-2020_12_31_424971 160 49 is be VBZ 10_1101-2020_12_31_424971 160 50 possible possible JJ 10_1101-2020_12_31_424971 160 51 that that IN 10_1101-2020_12_31_424971 160 52 the the DT 10_1101-2020_12_31_424971 160 53 distance distance NN 10_1101-2020_12_31_424971 160 54 between between IN 10_1101-2020_12_31_424971 160 55 our -PRON- PRP$ 10_1101-2020_12_31_424971 160 56 smFRET smFRET NNP 10_1101-2020_12_31_424971 160 57 probes probe NNS 10_1101-2020_12_31_424971 160 58 and/or and/or CC 10_1101-2020_12_31_424971 160 59 the the DT 10_1101-2020_12_31_424971 160 60 time time NN 10_1101-2020_12_31_424971 160 61 spent spend VBN 10_1101-2020_12_31_424971 160 62 in in IN 10_1101-2020_12_31_424971 160 63 one one CD 10_1101-2020_12_31_424971 160 64 of of IN 10_1101-2020_12_31_424971 160 65 402 402 CD 10_1101-2020_12_31_424971 160 66 the the DT 10_1101-2020_12_31_424971 160 67 observed observed JJ 10_1101-2020_12_31_424971 160 68 FRET FRET NNP 10_1101-2020_12_31_424971 160 69 states state NNS 10_1101-2020_12_31_424971 160 70 are be VBP 10_1101-2020_12_31_424971 160 71 not not RB 10_1101-2020_12_31_424971 160 72 sensitive sensitive JJ 10_1101-2020_12_31_424971 160 73 enough enough RB 10_1101-2020_12_31_424971 160 74 to to TO 10_1101-2020_12_31_424971 160 75 detect detect VB 10_1101-2020_12_31_424971 160 76 the the DT 10_1101-2020_12_31_424971 160 77 structural structural JJ 10_1101-2020_12_31_424971 160 78 and/or and/or CC 10_1101-2020_12_31_424971 160 79 energetic energetic JJ 10_1101-2020_12_31_424971 160 80 403 403 CD 10_1101-2020_12_31_424971 160 81 differences difference NNS 10_1101-2020_12_31_424971 160 82 between between IN 10_1101-2020_12_31_424971 160 83 these these DT 10_1101-2020_12_31_424971 160 84 sub sub NN 10_1101-2020_12_31_424971 160 85 - - NNS 10_1101-2020_12_31_424971 160 86 populations population NNS 10_1101-2020_12_31_424971 160 87 of of IN 10_1101-2020_12_31_424971 160 88 A a DT 10_1101-2020_12_31_424971 160 89 site site NN 10_1101-2020_12_31_424971 160 90 - - HYPH 10_1101-2020_12_31_424971 160 91 bound bind VBN 10_1101-2020_12_31_424971 160 92 SufB2 sufb2 NN 10_1101-2020_12_31_424971 160 93 . . . 10_1101-2020_12_31_424971 161 1 The the DT 10_1101-2020_12_31_424971 161 2 development development NN 10_1101-2020_12_31_424971 161 3 of of IN 10_1101-2020_12_31_424971 161 4 different different JJ 10_1101-2020_12_31_424971 161 5 404 404 CD 10_1101-2020_12_31_424971 161 6 smFRET smFRET NNP 10_1101-2020_12_31_424971 161 7 signals signal NNS 10_1101-2020_12_31_424971 161 8 and/or and/or CC 10_1101-2020_12_31_424971 161 9 the the DT 10_1101-2020_12_31_424971 161 10 use use NN 10_1101-2020_12_31_424971 161 11 of of IN 10_1101-2020_12_31_424971 161 12 variants variant NNS 10_1101-2020_12_31_424971 161 13 of of IN 10_1101-2020_12_31_424971 161 14 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 161 15 and/or and/or CC 10_1101-2020_12_31_424971 161 16 the the DT 10_1101-2020_12_31_424971 161 17 CCC CCC NNP 10_1101-2020_12_31_424971 161 18 - - HYPH 10_1101-2020_12_31_424971 161 19 C C NNP 10_1101-2020_12_31_424971 161 20 motif motif NN 10_1101-2020_12_31_424971 161 21 with with IN 10_1101-2020_12_31_424971 161 22 different different JJ 10_1101-2020_12_31_424971 161 23 405 405 CD 10_1101-2020_12_31_424971 161 24 propensities propensity NNS 10_1101-2020_12_31_424971 161 25 of of IN 10_1101-2020_12_31_424971 161 26 undergoing undergo VBG 10_1101-2020_12_31_424971 161 27 +1 +1 NNS 10_1101-2020_12_31_424971 161 28 frameshifting frameshifting NN 10_1101-2020_12_31_424971 161 29 may may MD 10_1101-2020_12_31_424971 161 30 allow allow VB 10_1101-2020_12_31_424971 161 31 future future JJ 10_1101-2020_12_31_424971 161 32 smFRET smFRET NNP 10_1101-2020_12_31_424971 161 33 investigations investigation NNS 10_1101-2020_12_31_424971 161 34 to to TO 10_1101-2020_12_31_424971 161 35 identify identify VB 10_1101-2020_12_31_424971 161 36 406 406 CD 10_1101-2020_12_31_424971 161 37 and and CC 10_1101-2020_12_31_424971 161 38 characterize characterize VB 10_1101-2020_12_31_424971 161 39 such such JJ 10_1101-2020_12_31_424971 161 40 sub sub NN 10_1101-2020_12_31_424971 161 41 - - NNS 10_1101-2020_12_31_424971 161 42 populations population NNS 10_1101-2020_12_31_424971 161 43 . . . 10_1101-2020_12_31_424971 162 1 407 407 CD 10_1101-2020_12_31_424971 162 2 .CC .CC : 10_1101-2020_12_31_424971 162 3 - - HYPH 10_1101-2020_12_31_424971 162 4 BY by IN 10_1101-2020_12_31_424971 162 5 - - HYPH 10_1101-2020_12_31_424971 162 6 NC NC NNP 10_1101-2020_12_31_424971 162 7 - - HYPH 10_1101-2020_12_31_424971 162 8 ND ND NNP 10_1101-2020_12_31_424971 162 9 4.0 4.0 CD 10_1101-2020_12_31_424971 162 10 International International NNP 10_1101-2020_12_31_424971 162 11 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 162 12 under under IN 10_1101-2020_12_31_424971 162 13 a a DT 10_1101-2020_12_31_424971 162 14 ( ( -LRB- 10_1101-2020_12_31_424971 162 15 which which WDT 10_1101-2020_12_31_424971 162 16 was be VBD 10_1101-2020_12_31_424971 162 17 not not RB 10_1101-2020_12_31_424971 162 18 certified certify VBN 10_1101-2020_12_31_424971 162 19 by by IN 10_1101-2020_12_31_424971 162 20 peer peer NN 10_1101-2020_12_31_424971 162 21 review review NN 10_1101-2020_12_31_424971 162 22 ) ) -RRB- 10_1101-2020_12_31_424971 162 23 is be VBZ 10_1101-2020_12_31_424971 162 24 the the DT 10_1101-2020_12_31_424971 162 25 author author NN 10_1101-2020_12_31_424971 162 26 / / SYM 10_1101-2020_12_31_424971 162 27 funder funder NN 10_1101-2020_12_31_424971 162 28 , , , 10_1101-2020_12_31_424971 162 29 who who WP 10_1101-2020_12_31_424971 162 30 has have VBZ 10_1101-2020_12_31_424971 162 31 granted grant VBN 10_1101-2020_12_31_424971 162 32 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 162 33 a a DT 10_1101-2020_12_31_424971 162 34 license license NN 10_1101-2020_12_31_424971 162 35 to to TO 10_1101-2020_12_31_424971 162 36 display display VB 10_1101-2020_12_31_424971 162 37 the the DT 10_1101-2020_12_31_424971 162 38 preprint preprint NN 10_1101-2020_12_31_424971 162 39 in in IN 10_1101-2020_12_31_424971 162 40 perpetuity perpetuity NN 10_1101-2020_12_31_424971 162 41 . . . 10_1101-2020_12_31_424971 163 1 It -PRON- PRP 10_1101-2020_12_31_424971 163 2 is be VBZ 10_1101-2020_12_31_424971 163 3 made make VBN 10_1101-2020_12_31_424971 163 4 The the DT 10_1101-2020_12_31_424971 163 5 copyright copyright NN 10_1101-2020_12_31_424971 163 6 holder holder NN 10_1101-2020_12_31_424971 163 7 for for IN 10_1101-2020_12_31_424971 163 8 this this DT 10_1101-2020_12_31_424971 163 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 163 10 version version NN 10_1101-2020_12_31_424971 163 11 posted post VBD 10_1101-2020_12_31_424971 163 12 January January NNP 10_1101-2020_12_31_424971 163 13 2 2 CD 10_1101-2020_12_31_424971 163 14 , , , 10_1101-2020_12_31_424971 163 15 2021 2021 CD 10_1101-2020_12_31_424971 163 16 . . . 10_1101-2020_12_31_424971 163 17 ; ; : 10_1101-2020_12_31_424971 163 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 163 19 : : : 10_1101-2020_12_31_424971 163 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 163 21 preprint preprint NN 10_1101-2020_12_31_424971 163 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 163 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 163 24 17 17 CD 10_1101-2020_12_31_424971 163 25 We -PRON- PRP 10_1101-2020_12_31_424971 163 26 then then RB 10_1101-2020_12_31_424971 163 27 investigated investigate VBD 10_1101-2020_12_31_424971 163 28 whether whether IN 10_1101-2020_12_31_424971 163 29 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 163 30 alters alter VBZ 10_1101-2020_12_31_424971 163 31 the the DT 10_1101-2020_12_31_424971 163 32 dynamics dynamic NNS 10_1101-2020_12_31_424971 163 33 of of IN 10_1101-2020_12_31_424971 163 34 elongation elongation NN 10_1101-2020_12_31_424971 163 35 complexes complex NNS 10_1101-2020_12_31_424971 163 36 during during IN 10_1101-2020_12_31_424971 163 37 the the DT 10_1101-2020_12_31_424971 163 38 408 408 CD 10_1101-2020_12_31_424971 163 39 later later JJ 10_1101-2020_12_31_424971 163 40 steps step NNS 10_1101-2020_12_31_424971 163 41 of of IN 10_1101-2020_12_31_424971 163 42 the the DT 10_1101-2020_12_31_424971 163 43 elongation elongation NN 10_1101-2020_12_31_424971 163 44 cycle cycle NN 10_1101-2020_12_31_424971 163 45 . . . 10_1101-2020_12_31_424971 164 1 We -PRON- PRP 10_1101-2020_12_31_424971 164 2 stopped stop VBD 10_1101-2020_12_31_424971 164 3 - - HYPH 10_1101-2020_12_31_424971 164 4 flow flow NN 10_1101-2020_12_31_424971 164 5 delivered deliver VBN 10_1101-2020_12_31_424971 164 6 SufB2- SufB2- NNP 10_1101-2020_12_31_424971 164 7 or or CC 10_1101-2020_12_31_424971 164 8 ProL ProL NNP 10_1101-2020_12_31_424971 164 9 - - HYPH 10_1101-2020_12_31_424971 164 10 TC TC NNP 10_1101-2020_12_31_424971 164 11 and and CC 10_1101-2020_12_31_424971 164 12 EF EF NNP 10_1101-2020_12_31_424971 164 13 - - HYPH 10_1101-2020_12_31_424971 164 14 G G NNP 10_1101-2020_12_31_424971 164 15 to to IN 10_1101-2020_12_31_424971 164 16 409 409 CD 10_1101-2020_12_31_424971 164 17 70S 70S NNP 10_1101-2020_12_31_424971 164 18 ICs ic NNS 10_1101-2020_12_31_424971 164 19 and and CC 10_1101-2020_12_31_424971 164 20 recorded record VBD 10_1101-2020_12_31_424971 164 21 pre pre JJ 10_1101-2020_12_31_424971 164 22 - - JJ 10_1101-2020_12_31_424971 164 23 steady steady JJ 10_1101-2020_12_31_424971 164 24 - - HYPH 10_1101-2020_12_31_424971 164 25 state state NN 10_1101-2020_12_31_424971 164 26 movies movie NNS 10_1101-2020_12_31_424971 164 27 during during IN 10_1101-2020_12_31_424971 164 28 delivery delivery NN 10_1101-2020_12_31_424971 164 29 , , , 10_1101-2020_12_31_424971 164 30 and and CC 10_1101-2020_12_31_424971 164 31 steady steady JJ 10_1101-2020_12_31_424971 164 32 - - HYPH 10_1101-2020_12_31_424971 164 33 state state NN 10_1101-2020_12_31_424971 164 34 movies movie NNS 10_1101-2020_12_31_424971 164 35 1 1 CD 10_1101-2020_12_31_424971 164 36 , , , 10_1101-2020_12_31_424971 164 37 3 3 CD 10_1101-2020_12_31_424971 164 38 , , , 10_1101-2020_12_31_424971 164 39 10 10 CD 10_1101-2020_12_31_424971 164 40 , , , 10_1101-2020_12_31_424971 164 41 410 410 CD 10_1101-2020_12_31_424971 164 42 and and CC 10_1101-2020_12_31_424971 164 43 20 20 CD 10_1101-2020_12_31_424971 164 44 min min NN 10_1101-2020_12_31_424971 164 45 post post JJ 10_1101-2020_12_31_424971 164 46 - - JJ 10_1101-2020_12_31_424971 164 47 delivery delivery JJ 10_1101-2020_12_31_424971 164 48 ( ( -LRB- 10_1101-2020_12_31_424971 164 49 Figures Figures NNPS 10_1101-2020_12_31_424971 164 50 6b 6b NNP 10_1101-2020_12_31_424971 164 51 , , , 10_1101-2020_12_31_424971 164 52 e e NN 10_1101-2020_12_31_424971 164 53 , , , 10_1101-2020_12_31_424971 164 54 and and CC 10_1101-2020_12_31_424971 164 55 g g NN 10_1101-2020_12_31_424971 164 56 , , , 10_1101-2020_12_31_424971 164 57 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 164 58 Figures Figures NNP 10_1101-2020_12_31_424971 164 59 4c 4c NNP 10_1101-2020_12_31_424971 164 60 , , , 10_1101-2020_12_31_424971 164 61 4d 4d CD 10_1101-2020_12_31_424971 164 62 , , , 10_1101-2020_12_31_424971 164 63 and and CC 10_1101-2020_12_31_424971 164 64 5 5 CD 10_1101-2020_12_31_424971 164 65 ) ) -RRB- 10_1101-2020_12_31_424971 164 66 . . . 10_1101-2020_12_31_424971 165 1 The the DT 10_1101-2020_12_31_424971 165 2 results result NNS 10_1101-2020_12_31_424971 165 3 411 411 CD 10_1101-2020_12_31_424971 165 4 showed show VBD 10_1101-2020_12_31_424971 165 5 that that IN 10_1101-2020_12_31_424971 165 6 k70S k70S NNP 10_1101-2020_12_31_424971 165 7 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 165 8 for for IN 10_1101-2020_12_31_424971 165 9 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 165 10 and and CC 10_1101-2020_12_31_424971 165 11 ProL ProL NNP 10_1101-2020_12_31_424971 165 12 - - HYPH 10_1101-2020_12_31_424971 165 13 TC TC NNP 10_1101-2020_12_31_424971 165 14 were be VBD 10_1101-2020_12_31_424971 165 15 within within IN 10_1101-2020_12_31_424971 165 16 error error NN 10_1101-2020_12_31_424971 165 17 of of IN 10_1101-2020_12_31_424971 165 18 each each DT 10_1101-2020_12_31_424971 165 19 other other JJ 10_1101-2020_12_31_424971 165 20 ( ( -LRB- 10_1101-2020_12_31_424971 165 21 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 165 22 412 412 CD 10_1101-2020_12_31_424971 165 23 Table table NN 10_1101-2020_12_31_424971 165 24 5 5 CD 10_1101-2020_12_31_424971 165 25 ) ) -RRB- 10_1101-2020_12_31_424971 165 26 , , , 10_1101-2020_12_31_424971 165 27 again again RB 10_1101-2020_12_31_424971 165 28 suggesting suggest VBG 10_1101-2020_12_31_424971 165 29 that that IN 10_1101-2020_12_31_424971 165 30 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 165 31 is be VBZ 10_1101-2020_12_31_424971 165 32 delivered deliver VBN 10_1101-2020_12_31_424971 165 33 to to IN 10_1101-2020_12_31_424971 165 34 the the DT 10_1101-2020_12_31_424971 165 35 A A NNP 10_1101-2020_12_31_424971 165 36 site site NN 10_1101-2020_12_31_424971 165 37 , , , 10_1101-2020_12_31_424971 165 38 participates participate VBZ 10_1101-2020_12_31_424971 165 39 in in IN 10_1101-2020_12_31_424971 165 40 peptide peptide NN 10_1101-2020_12_31_424971 165 41 - - HYPH 10_1101-2020_12_31_424971 165 42 bond bond NN 10_1101-2020_12_31_424971 165 43 413 413 CD 10_1101-2020_12_31_424971 165 44 formation formation NN 10_1101-2020_12_31_424971 165 45 , , , 10_1101-2020_12_31_424971 165 46 and and CC 10_1101-2020_12_31_424971 165 47 undergoes undergo VBZ 10_1101-2020_12_31_424971 165 48 GS2 gs2 NN 10_1101-2020_12_31_424971 165 49 formation formation NN 10_1101-2020_12_31_424971 165 50 in in IN 10_1101-2020_12_31_424971 165 51 a a DT 10_1101-2020_12_31_424971 165 52 manner manner NN 10_1101-2020_12_31_424971 165 53 that that WDT 10_1101-2020_12_31_424971 165 54 is be VBZ 10_1101-2020_12_31_424971 165 55 similar similar JJ 10_1101-2020_12_31_424971 165 56 to to IN 10_1101-2020_12_31_424971 165 57 ProL ProL NNP 10_1101-2020_12_31_424971 165 58 - - HYPH 10_1101-2020_12_31_424971 165 59 TC TC NNP 10_1101-2020_12_31_424971 165 60 . . . 10_1101-2020_12_31_424971 166 1 Notably notably RB 10_1101-2020_12_31_424971 166 2 , , , 10_1101-2020_12_31_424971 166 3 the the DT 10_1101-2020_12_31_424971 166 4 k70S k70S NNP 10_1101-2020_12_31_424971 166 5 414 414 CD 10_1101-2020_12_31_424971 166 6 IC→GS2s ic→gs2s CD 10_1101-2020_12_31_424971 166 7 obtained obtain VBN 10_1101-2020_12_31_424971 166 8 in in IN 10_1101-2020_12_31_424971 166 9 the the DT 10_1101-2020_12_31_424971 166 10 presence presence NN 10_1101-2020_12_31_424971 166 11 of of IN 10_1101-2020_12_31_424971 166 12 EF EF NNP 10_1101-2020_12_31_424971 166 13 - - HYPH 10_1101-2020_12_31_424971 166 14 G G NNP 10_1101-2020_12_31_424971 166 15 were be VBD 10_1101-2020_12_31_424971 166 16 within within IN 10_1101-2020_12_31_424971 166 17 error error NN 10_1101-2020_12_31_424971 166 18 of of IN 10_1101-2020_12_31_424971 166 19 the the DT 10_1101-2020_12_31_424971 166 20 ones one NNS 10_1101-2020_12_31_424971 166 21 obtained obtain VBN 10_1101-2020_12_31_424971 166 22 in in IN 10_1101-2020_12_31_424971 166 23 the the DT 10_1101-2020_12_31_424971 166 24 absence absence NN 10_1101-2020_12_31_424971 166 25 of of IN 10_1101-2020_12_31_424971 166 26 415 415 CD 10_1101-2020_12_31_424971 166 27 EF EF NNP 10_1101-2020_12_31_424971 166 28 - - HYPH 10_1101-2020_12_31_424971 166 29 G G NNP 10_1101-2020_12_31_424971 166 30 , , , 10_1101-2020_12_31_424971 166 31 consistent consistent JJ 10_1101-2020_12_31_424971 166 32 with with IN 10_1101-2020_12_31_424971 166 33 reports report NNS 10_1101-2020_12_31_424971 166 34 that that IN 10_1101-2020_12_31_424971 166 35 EF EF NNP 10_1101-2020_12_31_424971 166 36 - - HYPH 10_1101-2020_12_31_424971 166 37 G G NNP 10_1101-2020_12_31_424971 166 38 has have VBZ 10_1101-2020_12_31_424971 166 39 little little JJ 10_1101-2020_12_31_424971 166 40 to to IN 10_1101-2020_12_31_424971 166 41 no no DT 10_1101-2020_12_31_424971 166 42 effect effect NN 10_1101-2020_12_31_424971 166 43 on on IN 10_1101-2020_12_31_424971 166 44 the the DT 10_1101-2020_12_31_424971 166 45 rate rate NN 10_1101-2020_12_31_424971 166 46 with with IN 10_1101-2020_12_31_424971 166 47 which which WDT 10_1101-2020_12_31_424971 166 48 PRE PRE NNP 10_1101-2020_12_31_424971 166 49 416 416 CD 10_1101-2020_12_31_424971 166 50 complexes complex NNS 10_1101-2020_12_31_424971 166 51 undergo undergo VBP 10_1101-2020_12_31_424971 166 52 GS1→GS2 GS1→GS2 NNP 10_1101-2020_12_31_424971 166 53 transitions37,42 transitions37,42 NNP 10_1101-2020_12_31_424971 166 54 . . . 10_1101-2020_12_31_424971 167 1 417 417 CD 10_1101-2020_12_31_424971 167 2 Once once IN 10_1101-2020_12_31_424971 167 3 it -PRON- PRP 10_1101-2020_12_31_424971 167 4 transitions transition VBZ 10_1101-2020_12_31_424971 167 5 into into IN 10_1101-2020_12_31_424971 167 6 GS2 GS2 NNP 10_1101-2020_12_31_424971 167 7 , , , 10_1101-2020_12_31_424971 167 8 however however RB 10_1101-2020_12_31_424971 167 9 , , , 10_1101-2020_12_31_424971 167 10 the the DT 10_1101-2020_12_31_424971 167 11 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 167 12 PRE PRE NNP 10_1101-2020_12_31_424971 167 13 complex complex NN 10_1101-2020_12_31_424971 167 14 can can MD 10_1101-2020_12_31_424971 167 15 bind bind VB 10_1101-2020_12_31_424971 167 16 EF EF NNP 10_1101-2020_12_31_424971 167 17 - - HYPH 10_1101-2020_12_31_424971 167 18 G37,42 G37,42 NNP 10_1101-2020_12_31_424971 167 19 and and CC 10_1101-2020_12_31_424971 167 20 we -PRON- PRP 10_1101-2020_12_31_424971 167 21 find find VBP 10_1101-2020_12_31_424971 167 22 418 418 CD 10_1101-2020_12_31_424971 167 23 that that IN 10_1101-2020_12_31_424971 167 24 it -PRON- PRP 10_1101-2020_12_31_424971 167 25 becomes become VBZ 10_1101-2020_12_31_424971 167 26 arrested arrest VBN 10_1101-2020_12_31_424971 167 27 in in IN 10_1101-2020_12_31_424971 167 28 an an DT 10_1101-2020_12_31_424971 167 29 EF EF NNP 10_1101-2020_12_31_424971 167 30 - - HYPH 10_1101-2020_12_31_424971 167 31 G G NNP 10_1101-2020_12_31_424971 167 32 - - HYPH 10_1101-2020_12_31_424971 167 33 bound bind VBN 10_1101-2020_12_31_424971 167 34 GS2-like GS2-like NNP 10_1101-2020_12_31_424971 167 35 conformation conformation NN 10_1101-2020_12_31_424971 167 36 for for IN 10_1101-2020_12_31_424971 167 37 up up IN 10_1101-2020_12_31_424971 167 38 to to IN 10_1101-2020_12_31_424971 167 39 several several JJ 10_1101-2020_12_31_424971 167 40 minutes minute NNS 10_1101-2020_12_31_424971 167 41 , , , 10_1101-2020_12_31_424971 167 42 419 419 CD 10_1101-2020_12_31_424971 167 43 during during IN 10_1101-2020_12_31_424971 167 44 which which WDT 10_1101-2020_12_31_424971 167 45 it -PRON- PRP 10_1101-2020_12_31_424971 167 46 slowly slowly RB 10_1101-2020_12_31_424971 167 47 undergoes undergo VBZ 10_1101-2020_12_31_424971 167 48 a a DT 10_1101-2020_12_31_424971 167 49 GS2→POST gs2→post JJ 10_1101-2020_12_31_424971 167 50 transition transition NN 10_1101-2020_12_31_424971 167 51 ( ( -LRB- 10_1101-2020_12_31_424971 167 52 Figure figure NN 10_1101-2020_12_31_424971 167 53 6 6 CD 10_1101-2020_12_31_424971 167 54 g g NN 10_1101-2020_12_31_424971 167 55 , , , 10_1101-2020_12_31_424971 167 56 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 167 57 Figure Figure NNP 10_1101-2020_12_31_424971 167 58 5 5 CD 10_1101-2020_12_31_424971 167 59 ) ) -RRB- 10_1101-2020_12_31_424971 167 60 . . . 10_1101-2020_12_31_424971 168 1 420 420 CD 10_1101-2020_12_31_424971 168 2 While while IN 10_1101-2020_12_31_424971 168 3 the the DT 10_1101-2020_12_31_424971 168 4 limited limited JJ 10_1101-2020_12_31_424971 168 5 number number NN 10_1101-2020_12_31_424971 168 6 of of IN 10_1101-2020_12_31_424971 168 7 time time NN 10_1101-2020_12_31_424971 168 8 points point NNS 10_1101-2020_12_31_424971 168 9 did do VBD 10_1101-2020_12_31_424971 168 10 not not RB 10_1101-2020_12_31_424971 168 11 allow allow VB 10_1101-2020_12_31_424971 168 12 rigorous rigorous JJ 10_1101-2020_12_31_424971 168 13 determination determination NN 10_1101-2020_12_31_424971 168 14 of of IN 10_1101-2020_12_31_424971 168 15 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 168 16 for for IN 10_1101-2020_12_31_424971 168 17 the the DT 10_1101-2020_12_31_424971 168 18 421 421 CD 10_1101-2020_12_31_424971 168 19 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 168 20 PRE PRE NNP 10_1101-2020_12_31_424971 168 21 complex complex JJ 10_1101-2020_12_31_424971 168 22 , , , 10_1101-2020_12_31_424971 168 23 visual visual JJ 10_1101-2020_12_31_424971 168 24 inspection inspection NN 10_1101-2020_12_31_424971 168 25 ( ( -LRB- 10_1101-2020_12_31_424971 168 26 Figure figure NN 10_1101-2020_12_31_424971 168 27 6 6 CD 10_1101-2020_12_31_424971 168 28 g g NN 10_1101-2020_12_31_424971 168 29 ) ) -RRB- 10_1101-2020_12_31_424971 168 30 and and CC 10_1101-2020_12_31_424971 168 31 quantitative quantitative JJ 10_1101-2020_12_31_424971 168 32 analysis analysis NN 10_1101-2020_12_31_424971 168 33 ( ( -LRB- 10_1101-2020_12_31_424971 168 34 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 168 35 422 422 CD 10_1101-2020_12_31_424971 168 36 Tables Tables NNPS 10_1101-2020_12_31_424971 168 37 5 5 CD 10_1101-2020_12_31_424971 168 38 and and CC 10_1101-2020_12_31_424971 168 39 6 6 CD 10_1101-2020_12_31_424971 168 40 ) ) -RRB- 10_1101-2020_12_31_424971 168 41 showed show VBD 10_1101-2020_12_31_424971 168 42 that that IN 10_1101-2020_12_31_424971 168 43 the the DT 10_1101-2020_12_31_424971 168 44 GS2→POST gs2→post JJ 10_1101-2020_12_31_424971 168 45 reaction reaction NN 10_1101-2020_12_31_424971 168 46 was be VBD 10_1101-2020_12_31_424971 168 47 complete complete JJ 10_1101-2020_12_31_424971 168 48 between between IN 10_1101-2020_12_31_424971 168 49 3 3 CD 10_1101-2020_12_31_424971 168 50 and and CC 10_1101-2020_12_31_424971 168 51 10 10 CD 10_1101-2020_12_31_424971 168 52 min min NN 10_1101-2020_12_31_424971 168 53 post-423 post-423 NNP 10_1101-2020_12_31_424971 168 54 delivery delivery NN 10_1101-2020_12_31_424971 168 55 ( ( -LRB- 10_1101-2020_12_31_424971 168 56 i.e. i.e. FW 10_1101-2020_12_31_424971 168 57 , , , 10_1101-2020_12_31_424971 168 58 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 168 59 = = SYM 10_1101-2020_12_31_424971 168 60 ~0.0017–0.0060 ~0.0017–0.0060 NFP 10_1101-2020_12_31_424971 168 61 s–1 s–1 NNP 10_1101-2020_12_31_424971 168 62 ) ) -RRB- 10_1101-2020_12_31_424971 168 63 . . . 10_1101-2020_12_31_424971 169 1 Remarkably remarkably RB 10_1101-2020_12_31_424971 169 2 , , , 10_1101-2020_12_31_424971 169 3 this this DT 10_1101-2020_12_31_424971 169 4 range range NN 10_1101-2020_12_31_424971 169 5 of of IN 10_1101-2020_12_31_424971 169 6 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 169 7 is be VBZ 10_1101-2020_12_31_424971 169 8 up up IN 10_1101-2020_12_31_424971 169 9 to to TO 10_1101-2020_12_31_424971 169 10 2 2 CD 10_1101-2020_12_31_424971 169 11 - - SYM 10_1101-2020_12_31_424971 169 12 3 3 CD 10_1101-2020_12_31_424971 169 13 424 424 CD 10_1101-2020_12_31_424971 169 14 orders order NNS 10_1101-2020_12_31_424971 169 15 of of IN 10_1101-2020_12_31_424971 169 16 magnitude magnitude NN 10_1101-2020_12_31_424971 169 17 lower low JJR 10_1101-2020_12_31_424971 169 18 than than IN 10_1101-2020_12_31_424971 169 19 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 169 20 measured measure VBN 10_1101-2020_12_31_424971 169 21 for for IN 10_1101-2020_12_31_424971 169 22 the the DT 10_1101-2020_12_31_424971 169 23 ProL ProL NNP 10_1101-2020_12_31_424971 169 24 PRE PRE NNP 10_1101-2020_12_31_424971 169 25 complex complex NN 10_1101-2020_12_31_424971 169 26 ( ( -LRB- 10_1101-2020_12_31_424971 169 27 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 169 28 425 425 CD 10_1101-2020_12_31_424971 169 29 Table table NN 10_1101-2020_12_31_424971 169 30 5 5 CD 10_1101-2020_12_31_424971 169 31 ) ) -RRB- 10_1101-2020_12_31_424971 169 32 . . . 10_1101-2020_12_31_424971 170 1 It -PRON- PRP 10_1101-2020_12_31_424971 170 2 is be VBZ 10_1101-2020_12_31_424971 170 3 also also RB 10_1101-2020_12_31_424971 170 4 up up IN 10_1101-2020_12_31_424971 170 5 to to TO 10_1101-2020_12_31_424971 170 6 2 2 CD 10_1101-2020_12_31_424971 170 7 - - SYM 10_1101-2020_12_31_424971 170 8 3 3 CD 10_1101-2020_12_31_424971 170 9 orders order NNS 10_1101-2020_12_31_424971 170 10 of of IN 10_1101-2020_12_31_424971 170 11 magnitude magnitude NN 10_1101-2020_12_31_424971 170 12 lower low JJR 10_1101-2020_12_31_424971 170 13 than than IN 10_1101-2020_12_31_424971 170 14 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 170 15 for for IN 10_1101-2020_12_31_424971 170 16 a a DT 10_1101-2020_12_31_424971 170 17 different different JJ 10_1101-2020_12_31_424971 170 18 PRE PRE NNP 10_1101-2020_12_31_424971 170 19 complex complex JJ 10_1101-2020_12_31_424971 170 20 426 426 CD 10_1101-2020_12_31_424971 170 21 measured measure VBN 10_1101-2020_12_31_424971 170 22 using use VBG 10_1101-2020_12_31_424971 170 23 a a DT 10_1101-2020_12_31_424971 170 24 different different JJ 10_1101-2020_12_31_424971 170 25 smFRET smfret NN 10_1101-2020_12_31_424971 170 26 signal signal NN 10_1101-2020_12_31_424971 170 27 under under IN 10_1101-2020_12_31_424971 170 28 the the DT 10_1101-2020_12_31_424971 170 29 same same JJ 10_1101-2020_12_31_424971 170 30 conditions43 conditions43 NN 10_1101-2020_12_31_424971 170 31 and and CC 10_1101-2020_12_31_424971 170 32 the the DT 10_1101-2020_12_31_424971 170 33 rate rate NN 10_1101-2020_12_31_424971 170 34 of of IN 10_1101-2020_12_31_424971 170 35 427 427 CD 10_1101-2020_12_31_424971 170 36 translocation translocation NN 10_1101-2020_12_31_424971 170 37 measured measure VBN 10_1101-2020_12_31_424971 170 38 using use VBG 10_1101-2020_12_31_424971 170 39 ensemble ensemble JJ 10_1101-2020_12_31_424971 170 40 rapid rapid JJ 10_1101-2020_12_31_424971 170 41 kinetic kinetic JJ 10_1101-2020_12_31_424971 170 42 approaches approach NNS 10_1101-2020_12_31_424971 170 43 under under IN 10_1101-2020_12_31_424971 170 44 similar similar JJ 10_1101-2020_12_31_424971 170 45 conditions44,45 conditions44,45 NNP 10_1101-2020_12_31_424971 170 46 . . . 10_1101-2020_12_31_424971 171 1 428 428 CD 10_1101-2020_12_31_424971 171 2 This this DT 10_1101-2020_12_31_424971 171 3 observation observation NN 10_1101-2020_12_31_424971 171 4 suggests suggest VBZ 10_1101-2020_12_31_424971 171 5 that that IN 10_1101-2020_12_31_424971 171 6 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 171 7 adopts adopt VBZ 10_1101-2020_12_31_424971 171 8 a a DT 10_1101-2020_12_31_424971 171 9 conformation conformation NN 10_1101-2020_12_31_424971 171 10 within within IN 10_1101-2020_12_31_424971 171 11 the the DT 10_1101-2020_12_31_424971 171 12 EF EF NNP 10_1101-2020_12_31_424971 171 13 - - HYPH 10_1101-2020_12_31_424971 171 14 G G NNP 10_1101-2020_12_31_424971 171 15 - - HYPH 10_1101-2020_12_31_424971 171 16 bound bind VBN 10_1101-2020_12_31_424971 171 17 PRE PRE NNP 10_1101-2020_12_31_424971 171 18 complex complex NN 10_1101-2020_12_31_424971 171 19 429 429 CD 10_1101-2020_12_31_424971 171 20 that that WDT 10_1101-2020_12_31_424971 171 21 significantly significantly RB 10_1101-2020_12_31_424971 171 22 impedes impede VBZ 10_1101-2020_12_31_424971 171 23 conformational conformational JJ 10_1101-2020_12_31_424971 171 24 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 171 25 of of IN 10_1101-2020_12_31_424971 171 26 the the DT 10_1101-2020_12_31_424971 171 27 complex complex NN 10_1101-2020_12_31_424971 171 28 that that WDT 10_1101-2020_12_31_424971 171 29 are be VBP 10_1101-2020_12_31_424971 171 30 known know VBN 10_1101-2020_12_31_424971 171 31 to to TO 10_1101-2020_12_31_424971 171 32 take take VB 10_1101-2020_12_31_424971 171 33 430 430 CD 10_1101-2020_12_31_424971 171 34 place place NN 10_1101-2020_12_31_424971 171 35 during during IN 10_1101-2020_12_31_424971 171 36 late late JJ 10_1101-2020_12_31_424971 171 37 steps step NNS 10_1101-2020_12_31_424971 171 38 in in IN 10_1101-2020_12_31_424971 171 39 translocation translocation NN 10_1101-2020_12_31_424971 171 40 . . . 10_1101-2020_12_31_424971 172 1 These these DT 10_1101-2020_12_31_424971 172 2 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 172 3 include include VBP 10_1101-2020_12_31_424971 172 4 the the DT 10_1101-2020_12_31_424971 172 5 severing severing NN 10_1101-2020_12_31_424971 172 6 of of IN 10_1101-2020_12_31_424971 172 7 interactions interaction NNS 10_1101-2020_12_31_424971 172 8 431 431 CD 10_1101-2020_12_31_424971 172 9 .CC .CC : 10_1101-2020_12_31_424971 172 10 - - : 10_1101-2020_12_31_424971 172 11 BY by IN 10_1101-2020_12_31_424971 172 12 - - HYPH 10_1101-2020_12_31_424971 172 13 NC NC NNP 10_1101-2020_12_31_424971 172 14 - - HYPH 10_1101-2020_12_31_424971 172 15 ND ND NNP 10_1101-2020_12_31_424971 172 16 4.0 4.0 CD 10_1101-2020_12_31_424971 172 17 International International NNP 10_1101-2020_12_31_424971 172 18 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 172 19 under under IN 10_1101-2020_12_31_424971 172 20 a a DT 10_1101-2020_12_31_424971 172 21 ( ( -LRB- 10_1101-2020_12_31_424971 172 22 which which WDT 10_1101-2020_12_31_424971 172 23 was be VBD 10_1101-2020_12_31_424971 172 24 not not RB 10_1101-2020_12_31_424971 172 25 certified certify VBN 10_1101-2020_12_31_424971 172 26 by by IN 10_1101-2020_12_31_424971 172 27 peer peer NN 10_1101-2020_12_31_424971 172 28 review review NN 10_1101-2020_12_31_424971 172 29 ) ) -RRB- 10_1101-2020_12_31_424971 172 30 is be VBZ 10_1101-2020_12_31_424971 172 31 the the DT 10_1101-2020_12_31_424971 172 32 author author NN 10_1101-2020_12_31_424971 172 33 / / SYM 10_1101-2020_12_31_424971 172 34 funder funder NN 10_1101-2020_12_31_424971 172 35 , , , 10_1101-2020_12_31_424971 172 36 who who WP 10_1101-2020_12_31_424971 172 37 has have VBZ 10_1101-2020_12_31_424971 172 38 granted grant VBN 10_1101-2020_12_31_424971 172 39 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 172 40 a a DT 10_1101-2020_12_31_424971 172 41 license license NN 10_1101-2020_12_31_424971 172 42 to to TO 10_1101-2020_12_31_424971 172 43 display display VB 10_1101-2020_12_31_424971 172 44 the the DT 10_1101-2020_12_31_424971 172 45 preprint preprint NN 10_1101-2020_12_31_424971 172 46 in in IN 10_1101-2020_12_31_424971 172 47 perpetuity perpetuity NN 10_1101-2020_12_31_424971 172 48 . . . 10_1101-2020_12_31_424971 173 1 It -PRON- PRP 10_1101-2020_12_31_424971 173 2 is be VBZ 10_1101-2020_12_31_424971 173 3 made make VBN 10_1101-2020_12_31_424971 173 4 The the DT 10_1101-2020_12_31_424971 173 5 copyright copyright NN 10_1101-2020_12_31_424971 173 6 holder holder NN 10_1101-2020_12_31_424971 173 7 for for IN 10_1101-2020_12_31_424971 173 8 this this DT 10_1101-2020_12_31_424971 173 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 173 10 version version NN 10_1101-2020_12_31_424971 173 11 posted post VBD 10_1101-2020_12_31_424971 173 12 January January NNP 10_1101-2020_12_31_424971 173 13 2 2 CD 10_1101-2020_12_31_424971 173 14 , , , 10_1101-2020_12_31_424971 173 15 2021 2021 CD 10_1101-2020_12_31_424971 173 16 . . . 10_1101-2020_12_31_424971 173 17 ; ; : 10_1101-2020_12_31_424971 173 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 173 19 : : : 10_1101-2020_12_31_424971 173 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 173 21 preprint preprint NN 10_1101-2020_12_31_424971 173 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 173 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 173 24 18 18 CD 10_1101-2020_12_31_424971 173 25 between between IN 10_1101-2020_12_31_424971 173 26 the the DT 10_1101-2020_12_31_424971 173 27 decoding decode VBG 10_1101-2020_12_31_424971 173 28 center center NN 10_1101-2020_12_31_424971 173 29 of of IN 10_1101-2020_12_31_424971 173 30 the the DT 10_1101-2020_12_31_424971 173 31 30S 30s CD 10_1101-2020_12_31_424971 173 32 subunit subunit NN 10_1101-2020_12_31_424971 173 33 and and CC 10_1101-2020_12_31_424971 173 34 the the DT 10_1101-2020_12_31_424971 173 35 anticodon anticodon NN 10_1101-2020_12_31_424971 173 36 - - HYPH 10_1101-2020_12_31_424971 173 37 codon codon NNP 10_1101-2020_12_31_424971 173 38 duplex duplex NN 10_1101-2020_12_31_424971 173 39 in in IN 10_1101-2020_12_31_424971 173 40 the the DT 10_1101-2020_12_31_424971 173 41 A A NNP 10_1101-2020_12_31_424971 173 42 site22 site22 NNP 10_1101-2020_12_31_424971 173 43 - - SYM 10_1101-2020_12_31_424971 173 44 432 432 CD 10_1101-2020_12_31_424971 173 45 25 25 CD 10_1101-2020_12_31_424971 173 46 ; ; : 10_1101-2020_12_31_424971 173 47 forward forward RB 10_1101-2020_12_31_424971 173 48 and and CC 10_1101-2020_12_31_424971 173 49 reverse reverse VB 10_1101-2020_12_31_424971 173 50 swiveling swiveling NN 10_1101-2020_12_31_424971 173 51 of of IN 10_1101-2020_12_31_424971 173 52 the the DT 10_1101-2020_12_31_424971 173 53 ‘ ' `` 10_1101-2020_12_31_424971 173 54 head head NN 10_1101-2020_12_31_424971 173 55 ’ ' '' 10_1101-2020_12_31_424971 173 56 domain domain NN 10_1101-2020_12_31_424971 173 57 of of IN 10_1101-2020_12_31_424971 173 58 the the DT 10_1101-2020_12_31_424971 173 59 30S 30S NNP 10_1101-2020_12_31_424971 173 60 subunit27,28 subunit27,28 NNP 10_1101-2020_12_31_424971 173 61 associated associate VBD 10_1101-2020_12_31_424971 173 62 with with IN 10_1101-2020_12_31_424971 173 63 433 433 CD 10_1101-2020_12_31_424971 173 64 opening opening NN 10_1101-2020_12_31_424971 173 65 and and CC 10_1101-2020_12_31_424971 173 66 closing closing NN 10_1101-2020_12_31_424971 173 67 , , , 10_1101-2020_12_31_424971 173 68 respectively respectively RB 10_1101-2020_12_31_424971 173 69 , , , 10_1101-2020_12_31_424971 173 70 of of IN 10_1101-2020_12_31_424971 173 71 the the DT 10_1101-2020_12_31_424971 173 72 ‘ ' `` 10_1101-2020_12_31_424971 173 73 E e NN 10_1101-2020_12_31_424971 173 74 - - NN 10_1101-2020_12_31_424971 173 75 site site NN 10_1101-2020_12_31_424971 173 76 gate gate NN 10_1101-2020_12_31_424971 173 77 ’ ' '' 10_1101-2020_12_31_424971 173 78 of of IN 10_1101-2020_12_31_424971 173 79 the the DT 10_1101-2020_12_31_424971 173 80 30S 30s CD 10_1101-2020_12_31_424971 173 81 subunit26 subunit26 VBZ 10_1101-2020_12_31_424971 173 82 ; ; : 10_1101-2020_12_31_424971 173 83 reverse reverse JJ 10_1101-2020_12_31_424971 173 84 relative relative JJ 10_1101-2020_12_31_424971 173 85 rotation rotation NN 10_1101-2020_12_31_424971 173 86 434 434 CD 10_1101-2020_12_31_424971 173 87 of of IN 10_1101-2020_12_31_424971 173 88 the the DT 10_1101-2020_12_31_424971 173 89 ribosomal ribosomal NN 10_1101-2020_12_31_424971 173 90 subunits46,47 subunits46,47 NNP 10_1101-2020_12_31_424971 173 91 ; ; : 10_1101-2020_12_31_424971 173 92 and and CC 10_1101-2020_12_31_424971 173 93 opening opening NN 10_1101-2020_12_31_424971 173 94 of of IN 10_1101-2020_12_31_424971 173 95 the the DT 10_1101-2020_12_31_424971 173 96 L1 L1 NNP 10_1101-2020_12_31_424971 173 97 stalk35,37,48 stalk35,37,48 NNP 10_1101-2020_12_31_424971 173 98 . . . 10_1101-2020_12_31_424971 174 1 Collectively collectively RB 10_1101-2020_12_31_424971 174 2 , , , 10_1101-2020_12_31_424971 174 3 these these DT 10_1101-2020_12_31_424971 174 4 dynamics dynamic NNS 10_1101-2020_12_31_424971 174 5 435 435 CD 10_1101-2020_12_31_424971 174 6 facilitate facilitate JJ 10_1101-2020_12_31_424971 174 7 movement movement NN 10_1101-2020_12_31_424971 174 8 of of IN 10_1101-2020_12_31_424971 174 9 the the DT 10_1101-2020_12_31_424971 174 10 tRNA trna NN 10_1101-2020_12_31_424971 174 11 ASLs asl NNS 10_1101-2020_12_31_424971 174 12 and and CC 10_1101-2020_12_31_424971 174 13 their -PRON- PRP$ 10_1101-2020_12_31_424971 174 14 associated associated JJ 10_1101-2020_12_31_424971 174 15 codons codon NNS 10_1101-2020_12_31_424971 174 16 from from IN 10_1101-2020_12_31_424971 174 17 the the DT 10_1101-2020_12_31_424971 174 18 P p NN 10_1101-2020_12_31_424971 174 19 and and CC 10_1101-2020_12_31_424971 174 20 A a DT 10_1101-2020_12_31_424971 174 21 sites site NNS 10_1101-2020_12_31_424971 174 22 to to IN 10_1101-2020_12_31_424971 174 23 the the DT 10_1101-2020_12_31_424971 174 24 436 436 CD 10_1101-2020_12_31_424971 174 25 E E NNS 10_1101-2020_12_31_424971 174 26 and and CC 10_1101-2020_12_31_424971 174 27 P p NN 10_1101-2020_12_31_424971 174 28 sites site NNS 10_1101-2020_12_31_424971 174 29 of of IN 10_1101-2020_12_31_424971 174 30 the the DT 10_1101-2020_12_31_424971 174 31 30S 30s CD 10_1101-2020_12_31_424971 174 32 subunit subunit NN 10_1101-2020_12_31_424971 174 33 . . . 10_1101-2020_12_31_424971 175 1 437 437 CD 10_1101-2020_12_31_424971 175 2 We -PRON- PRP 10_1101-2020_12_31_424971 175 3 next next RB 10_1101-2020_12_31_424971 175 4 explored explore VBD 10_1101-2020_12_31_424971 175 5 whether whether IN 10_1101-2020_12_31_424971 175 6 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 175 7 alters alter VBZ 10_1101-2020_12_31_424971 175 8 the the DT 10_1101-2020_12_31_424971 175 9 dynamics dynamic NNS 10_1101-2020_12_31_424971 175 10 of of IN 10_1101-2020_12_31_424971 175 11 elongation elongation NN 10_1101-2020_12_31_424971 175 12 complexes complex NNS 10_1101-2020_12_31_424971 175 13 after after IN 10_1101-2020_12_31_424971 175 14 it -PRON- PRP 10_1101-2020_12_31_424971 175 15 is be VBZ 10_1101-2020_12_31_424971 175 16 438 438 CD 10_1101-2020_12_31_424971 175 17 translocated translocate VBN 10_1101-2020_12_31_424971 175 18 into into IN 10_1101-2020_12_31_424971 175 19 the the DT 10_1101-2020_12_31_424971 175 20 P p NN 10_1101-2020_12_31_424971 175 21 site site NN 10_1101-2020_12_31_424971 175 22 . . . 10_1101-2020_12_31_424971 176 1 We -PRON- PRP 10_1101-2020_12_31_424971 176 2 prepared prepare VBD 10_1101-2020_12_31_424971 176 3 PRE PRE NNP 10_1101-2020_12_31_424971 176 4 - - HYPH 10_1101-2020_12_31_424971 176 5 like like JJ 10_1101-2020_12_31_424971 176 6 complexes complex NNS 10_1101-2020_12_31_424971 176 7 carrying carry VBG 10_1101-2020_12_31_424971 176 8 deacylated deacylate VBN 10_1101-2020_12_31_424971 176 9 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 176 10 or or CC 10_1101-2020_12_31_424971 176 11 ProL ProL NNP 10_1101-2020_12_31_424971 176 12 439 439 CD 10_1101-2020_12_31_424971 176 13 in in IN 10_1101-2020_12_31_424971 176 14 the the DT 10_1101-2020_12_31_424971 176 15 P p NN 10_1101-2020_12_31_424971 176 16 site site NN 10_1101-2020_12_31_424971 176 17 and and CC 10_1101-2020_12_31_424971 176 18 a a DT 10_1101-2020_12_31_424971 176 19 vacant vacant JJ 10_1101-2020_12_31_424971 176 20 A a NN 10_1101-2020_12_31_424971 176 21 site site NN 10_1101-2020_12_31_424971 176 22 ( ( -LRB- 10_1101-2020_12_31_424971 176 23 denoted denote VBN 10_1101-2020_12_31_424971 176 24 PRE PRE NNP 10_1101-2020_12_31_424971 176 25 – – : 10_1101-2020_12_31_424971 176 26 A a DT 10_1101-2020_12_31_424971 176 27 complexes complex NNS 10_1101-2020_12_31_424971 176 28 ) ) -RRB- 10_1101-2020_12_31_424971 176 29 and and CC 10_1101-2020_12_31_424971 176 30 recorded record VBD 10_1101-2020_12_31_424971 176 31 steady steady JJ 10_1101-2020_12_31_424971 176 32 - - HYPH 10_1101-2020_12_31_424971 176 33 state state NN 10_1101-2020_12_31_424971 176 34 movies movie NNS 10_1101-2020_12_31_424971 176 35 440 440 CD 10_1101-2020_12_31_424971 176 36 for for IN 10_1101-2020_12_31_424971 176 37 the the DT 10_1101-2020_12_31_424971 176 38 resulting result VBG 10_1101-2020_12_31_424971 176 39 GS1 GS1 NNP 10_1101-2020_12_31_424971 176 40 ⇄ ⇄ NNP 10_1101-2020_12_31_424971 176 41 GS2 GS2 NNP 10_1101-2020_12_31_424971 176 42 equilibria equilibrium NNS 10_1101-2020_12_31_424971 176 43 ( ( -LRB- 10_1101-2020_12_31_424971 176 44 Figures figure NNS 10_1101-2020_12_31_424971 176 45 6c 6c NNPS 10_1101-2020_12_31_424971 176 46 and and CC 10_1101-2020_12_31_424971 176 47 h h NN 10_1101-2020_12_31_424971 176 48 , , , 10_1101-2020_12_31_424971 176 49 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 176 50 Figure Figure NNP 10_1101-2020_12_31_424971 176 51 6 6 CD 10_1101-2020_12_31_424971 176 52 ) ) -RRB- 10_1101-2020_12_31_424971 176 53 . . . 10_1101-2020_12_31_424971 177 1 The the DT 10_1101-2020_12_31_424971 177 2 results result NNS 10_1101-2020_12_31_424971 177 3 441 441 CD 10_1101-2020_12_31_424971 177 4 showed show VBD 10_1101-2020_12_31_424971 177 5 that that IN 10_1101-2020_12_31_424971 177 6 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 177 7 and and CC 10_1101-2020_12_31_424971 177 8 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 177 9 for for IN 10_1101-2020_12_31_424971 177 10 the the DT 10_1101-2020_12_31_424971 177 11 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 177 12 PRE PRE NNP 10_1101-2020_12_31_424971 177 13 – – : 10_1101-2020_12_31_424971 177 14 A a DT 10_1101-2020_12_31_424971 177 15 complex complex NN 10_1101-2020_12_31_424971 177 16 were be VBD 10_1101-2020_12_31_424971 177 17 45 45 CD 10_1101-2020_12_31_424971 177 18 % % NN 10_1101-2020_12_31_424971 177 19 lower low JJR 10_1101-2020_12_31_424971 177 20 and and CC 10_1101-2020_12_31_424971 177 21 36 36 CD 10_1101-2020_12_31_424971 177 22 % % NN 10_1101-2020_12_31_424971 177 23 higher high JJR 10_1101-2020_12_31_424971 177 24 , , , 10_1101-2020_12_31_424971 177 25 442 442 CD 10_1101-2020_12_31_424971 177 26 respectively respectively RB 10_1101-2020_12_31_424971 177 27 , , , 10_1101-2020_12_31_424971 177 28 than than IN 10_1101-2020_12_31_424971 177 29 for for IN 10_1101-2020_12_31_424971 177 30 the the DT 10_1101-2020_12_31_424971 177 31 ProL ProL NNP 10_1101-2020_12_31_424971 177 32 PRE PRE NNP 10_1101-2020_12_31_424971 177 33 – – : 10_1101-2020_12_31_424971 177 34 A a DT 10_1101-2020_12_31_424971 177 35 complex complex NN 10_1101-2020_12_31_424971 177 36 , , , 10_1101-2020_12_31_424971 177 37 driving drive VBG 10_1101-2020_12_31_424971 177 38 a a DT 10_1101-2020_12_31_424971 177 39 2.5-fold 2.5-fold CD 10_1101-2020_12_31_424971 177 40 shift shift NN 10_1101-2020_12_31_424971 177 41 towards towards IN 10_1101-2020_12_31_424971 177 42 GS1 GS1 NNP 10_1101-2020_12_31_424971 177 43 in in IN 10_1101-2020_12_31_424971 177 44 the the DT 10_1101-2020_12_31_424971 177 45 443 443 CD 10_1101-2020_12_31_424971 177 46 GS1 GS1 NNP 10_1101-2020_12_31_424971 177 47 ⇄ ⇄ NNP 10_1101-2020_12_31_424971 177 48 GS2 GS2 NNP 10_1101-2020_12_31_424971 177 49 equilibrium equilibrium NN 10_1101-2020_12_31_424971 177 50 ( ( -LRB- 10_1101-2020_12_31_424971 177 51 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 177 52 Table Table NNP 10_1101-2020_12_31_424971 177 53 7 7 CD 10_1101-2020_12_31_424971 177 54 ) ) -RRB- 10_1101-2020_12_31_424971 177 55 , , , 10_1101-2020_12_31_424971 177 56 suggesting suggest VBG 10_1101-2020_12_31_424971 177 57 that that IN 10_1101-2020_12_31_424971 177 58 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 177 59 adopts adopt VBZ 10_1101-2020_12_31_424971 177 60 a a DT 10_1101-2020_12_31_424971 177 61 conformation conformation NN 10_1101-2020_12_31_424971 177 62 at at IN 10_1101-2020_12_31_424971 177 63 444 444 CD 10_1101-2020_12_31_424971 177 64 the the DT 10_1101-2020_12_31_424971 177 65 P p NN 10_1101-2020_12_31_424971 177 66 site site NN 10_1101-2020_12_31_424971 177 67 that that WDT 10_1101-2020_12_31_424971 177 68 is be VBZ 10_1101-2020_12_31_424971 177 69 different different JJ 10_1101-2020_12_31_424971 177 70 from from IN 10_1101-2020_12_31_424971 177 71 that that DT 10_1101-2020_12_31_424971 177 72 of of IN 10_1101-2020_12_31_424971 177 73 ProL. prol. NN 10_1101-2020_12_31_424971 178 1 Consistent consistent JJ 10_1101-2020_12_31_424971 178 2 with with IN 10_1101-2020_12_31_424971 178 3 this this DT 10_1101-2020_12_31_424971 178 4 interpretation interpretation NN 10_1101-2020_12_31_424971 178 5 , , , 10_1101-2020_12_31_424971 178 6 a a DT 10_1101-2020_12_31_424971 178 7 recent recent JJ 10_1101-2020_12_31_424971 178 8 structural structural JJ 10_1101-2020_12_31_424971 178 9 445 445 CD 10_1101-2020_12_31_424971 178 10 study study NN 10_1101-2020_12_31_424971 178 11 has have VBZ 10_1101-2020_12_31_424971 178 12 shown show VBN 10_1101-2020_12_31_424971 178 13 that that IN 10_1101-2020_12_31_424971 178 14 the the DT 10_1101-2020_12_31_424971 178 15 conformation conformation NN 10_1101-2020_12_31_424971 178 16 of of IN 10_1101-2020_12_31_424971 178 17 P p NN 10_1101-2020_12_31_424971 178 18 site site NN 10_1101-2020_12_31_424971 178 19 - - HYPH 10_1101-2020_12_31_424971 178 20 bound bind VBN 10_1101-2020_12_31_424971 178 21 SufA6 SufA6 NNP 10_1101-2020_12_31_424971 178 22 , , , 10_1101-2020_12_31_424971 178 23 a a DT 10_1101-2020_12_31_424971 178 24 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 178 25 tRNA trna NN 10_1101-2020_12_31_424971 178 26 with with IN 10_1101-2020_12_31_424971 178 27 an an DT 10_1101-2020_12_31_424971 178 28 446 446 CD 10_1101-2020_12_31_424971 178 29 extra extra JJ 10_1101-2020_12_31_424971 178 30 nucleotide nucleotide RB 10_1101-2020_12_31_424971 178 31 in in IN 10_1101-2020_12_31_424971 178 32 the the DT 10_1101-2020_12_31_424971 178 33 anticodon anticodon NNP 10_1101-2020_12_31_424971 178 34 loop loop NN 10_1101-2020_12_31_424971 178 35 , , , 10_1101-2020_12_31_424971 178 36 is be VBZ 10_1101-2020_12_31_424971 178 37 significantly significantly RB 10_1101-2020_12_31_424971 178 38 distorted distort VBN 10_1101-2020_12_31_424971 178 39 relative relative JJ 10_1101-2020_12_31_424971 178 40 to to IN 10_1101-2020_12_31_424971 178 41 a a DT 10_1101-2020_12_31_424971 178 42 canonical canonical JJ 10_1101-2020_12_31_424971 178 43 tRNA49 trna49 NN 10_1101-2020_12_31_424971 178 44 . . . 10_1101-2020_12_31_424971 179 1 447 447 CD 10_1101-2020_12_31_424971 179 2 448 448 CD 10_1101-2020_12_31_424971 179 3 DISCUSSION discussion NN 10_1101-2020_12_31_424971 179 4 449 449 CD 10_1101-2020_12_31_424971 179 5 Here here RB 10_1101-2020_12_31_424971 179 6 we -PRON- PRP 10_1101-2020_12_31_424971 179 7 leverage leverage VBP 10_1101-2020_12_31_424971 179 8 the the DT 10_1101-2020_12_31_424971 179 9 high high JJ 10_1101-2020_12_31_424971 179 10 efficiency efficiency NN 10_1101-2020_12_31_424971 179 11 of of IN 10_1101-2020_12_31_424971 179 12 recoding recode VBG 10_1101-2020_12_31_424971 179 13 by by IN 10_1101-2020_12_31_424971 179 14 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 179 15 to to TO 10_1101-2020_12_31_424971 179 16 identify identify VB 10_1101-2020_12_31_424971 179 17 the the DT 10_1101-2020_12_31_424971 179 18 steps step NNS 10_1101-2020_12_31_424971 179 19 of of IN 10_1101-2020_12_31_424971 179 20 the the DT 10_1101-2020_12_31_424971 179 21 450 450 CD 10_1101-2020_12_31_424971 179 22 elongation elongation NN 10_1101-2020_12_31_424971 179 23 cycle cycle NN 10_1101-2020_12_31_424971 179 24 during during IN 10_1101-2020_12_31_424971 179 25 which which WDT 10_1101-2020_12_31_424971 179 26 it -PRON- PRP 10_1101-2020_12_31_424971 179 27 induces induce VBZ 10_1101-2020_12_31_424971 179 28 +1 +1 : 10_1101-2020_12_31_424971 179 29 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 179 30 at at IN 10_1101-2020_12_31_424971 179 31 a a DT 10_1101-2020_12_31_424971 179 32 quadruplet quadruplet NN 10_1101-2020_12_31_424971 179 33 codon codon NN 10_1101-2020_12_31_424971 179 34 , , , 10_1101-2020_12_31_424971 179 35 thus thus RB 10_1101-2020_12_31_424971 179 36 answering answer VBG 10_1101-2020_12_31_424971 179 37 451 451 CD 10_1101-2020_12_31_424971 179 38 the the DT 10_1101-2020_12_31_424971 179 39 key key JJ 10_1101-2020_12_31_424971 179 40 questions question NNS 10_1101-2020_12_31_424971 179 41 of of IN 10_1101-2020_12_31_424971 179 42 where where WRB 10_1101-2020_12_31_424971 179 43 , , , 10_1101-2020_12_31_424971 179 44 when when WRB 10_1101-2020_12_31_424971 179 45 , , , 10_1101-2020_12_31_424971 179 46 and and CC 10_1101-2020_12_31_424971 179 47 how how WRB 10_1101-2020_12_31_424971 179 48 +1 +1 NNS 10_1101-2020_12_31_424971 179 49 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 179 50 occurs occur NNS 10_1101-2020_12_31_424971 179 51 . . . 10_1101-2020_12_31_424971 180 1 We -PRON- PRP 10_1101-2020_12_31_424971 180 2 are be VBP 10_1101-2020_12_31_424971 180 3 not not RB 10_1101-2020_12_31_424971 180 4 aware aware JJ 10_1101-2020_12_31_424971 180 5 of of IN 10_1101-2020_12_31_424971 180 6 any any DT 10_1101-2020_12_31_424971 180 7 452 452 CD 10_1101-2020_12_31_424971 180 8 other other JJ 10_1101-2020_12_31_424971 180 9 studies study NNS 10_1101-2020_12_31_424971 180 10 of of IN 10_1101-2020_12_31_424971 180 11 +1 +1 NNS 10_1101-2020_12_31_424971 180 12 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 180 13 that that WDT 10_1101-2020_12_31_424971 180 14 have have VBP 10_1101-2020_12_31_424971 180 15 addressed address VBN 10_1101-2020_12_31_424971 180 16 these these DT 10_1101-2020_12_31_424971 180 17 questions question NNS 10_1101-2020_12_31_424971 180 18 as as RB 10_1101-2020_12_31_424971 180 19 precisely precisely RB 10_1101-2020_12_31_424971 180 20 . . . 10_1101-2020_12_31_424971 181 1 In in IN 10_1101-2020_12_31_424971 181 2 addition addition NN 10_1101-2020_12_31_424971 181 3 to to IN 10_1101-2020_12_31_424971 181 4 453 453 CD 10_1101-2020_12_31_424971 181 5 elucidating elucidate VBG 10_1101-2020_12_31_424971 181 6 the the DT 10_1101-2020_12_31_424971 181 7 determinants determinant NNS 10_1101-2020_12_31_424971 181 8 of of IN 10_1101-2020_12_31_424971 181 9 reading reading NN 10_1101-2020_12_31_424971 181 10 - - HYPH 10_1101-2020_12_31_424971 181 11 frame frame NN 10_1101-2020_12_31_424971 181 12 maintenance maintenance NN 10_1101-2020_12_31_424971 181 13 and and CC 10_1101-2020_12_31_424971 181 14 the the DT 10_1101-2020_12_31_424971 181 15 mechanisms mechanism NNS 10_1101-2020_12_31_424971 181 16 of of IN 10_1101-2020_12_31_424971 181 17 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 181 18 - - SYM 10_1101-2020_12_31_424971 181 19 454 454 CD 10_1101-2020_12_31_424971 181 20 induced induced JJ 10_1101-2020_12_31_424971 181 21 +1 +1 NNS 10_1101-2020_12_31_424971 181 22 frameshifting frameshifting NN 10_1101-2020_12_31_424971 181 23 , , , 10_1101-2020_12_31_424971 181 24 our -PRON- PRP$ 10_1101-2020_12_31_424971 181 25 findings finding NNS 10_1101-2020_12_31_424971 181 26 reveal reveal VBP 10_1101-2020_12_31_424971 181 27 new new JJ 10_1101-2020_12_31_424971 181 28 principles principle NNS 10_1101-2020_12_31_424971 181 29 that that WDT 10_1101-2020_12_31_424971 181 30 can can MD 10_1101-2020_12_31_424971 181 31 be be VB 10_1101-2020_12_31_424971 181 32 used use VBN 10_1101-2020_12_31_424971 181 33 to to TO 10_1101-2020_12_31_424971 181 34 engineer engineer VB 10_1101-2020_12_31_424971 181 35 genome genome NNP 10_1101-2020_12_31_424971 181 36 455 455 CD 10_1101-2020_12_31_424971 181 37 recoding recode VBG 10_1101-2020_12_31_424971 181 38 with with IN 10_1101-2020_12_31_424971 181 39 higher high JJR 10_1101-2020_12_31_424971 181 40 efficiencies efficiency NNS 10_1101-2020_12_31_424971 181 41 . . . 10_1101-2020_12_31_424971 182 1 456 456 CD 10_1101-2020_12_31_424971 182 2 .CC .CC NFP 10_1101-2020_12_31_424971 182 3 - - HYPH 10_1101-2020_12_31_424971 182 4 BY by IN 10_1101-2020_12_31_424971 182 5 - - HYPH 10_1101-2020_12_31_424971 182 6 NC NC NNP 10_1101-2020_12_31_424971 182 7 - - HYPH 10_1101-2020_12_31_424971 182 8 ND ND NNP 10_1101-2020_12_31_424971 182 9 4.0 4.0 CD 10_1101-2020_12_31_424971 182 10 International International NNP 10_1101-2020_12_31_424971 182 11 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 182 12 under under IN 10_1101-2020_12_31_424971 182 13 a a DT 10_1101-2020_12_31_424971 182 14 ( ( -LRB- 10_1101-2020_12_31_424971 182 15 which which WDT 10_1101-2020_12_31_424971 182 16 was be VBD 10_1101-2020_12_31_424971 182 17 not not RB 10_1101-2020_12_31_424971 182 18 certified certify VBN 10_1101-2020_12_31_424971 182 19 by by IN 10_1101-2020_12_31_424971 182 20 peer peer NN 10_1101-2020_12_31_424971 182 21 review review NN 10_1101-2020_12_31_424971 182 22 ) ) -RRB- 10_1101-2020_12_31_424971 182 23 is be VBZ 10_1101-2020_12_31_424971 182 24 the the DT 10_1101-2020_12_31_424971 182 25 author author NN 10_1101-2020_12_31_424971 182 26 / / SYM 10_1101-2020_12_31_424971 182 27 funder funder NN 10_1101-2020_12_31_424971 182 28 , , , 10_1101-2020_12_31_424971 182 29 who who WP 10_1101-2020_12_31_424971 182 30 has have VBZ 10_1101-2020_12_31_424971 182 31 granted grant VBN 10_1101-2020_12_31_424971 182 32 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 182 33 a a DT 10_1101-2020_12_31_424971 182 34 license license NN 10_1101-2020_12_31_424971 182 35 to to TO 10_1101-2020_12_31_424971 182 36 display display VB 10_1101-2020_12_31_424971 182 37 the the DT 10_1101-2020_12_31_424971 182 38 preprint preprint NN 10_1101-2020_12_31_424971 182 39 in in IN 10_1101-2020_12_31_424971 182 40 perpetuity perpetuity NN 10_1101-2020_12_31_424971 182 41 . . . 10_1101-2020_12_31_424971 183 1 It -PRON- PRP 10_1101-2020_12_31_424971 183 2 is be VBZ 10_1101-2020_12_31_424971 183 3 made make VBN 10_1101-2020_12_31_424971 183 4 The the DT 10_1101-2020_12_31_424971 183 5 copyright copyright NN 10_1101-2020_12_31_424971 183 6 holder holder NN 10_1101-2020_12_31_424971 183 7 for for IN 10_1101-2020_12_31_424971 183 8 this this DT 10_1101-2020_12_31_424971 183 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 183 10 version version NN 10_1101-2020_12_31_424971 183 11 posted post VBD 10_1101-2020_12_31_424971 183 12 January January NNP 10_1101-2020_12_31_424971 183 13 2 2 CD 10_1101-2020_12_31_424971 183 14 , , , 10_1101-2020_12_31_424971 183 15 2021 2021 CD 10_1101-2020_12_31_424971 183 16 . . . 10_1101-2020_12_31_424971 183 17 ; ; : 10_1101-2020_12_31_424971 183 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 183 19 : : : 10_1101-2020_12_31_424971 183 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 183 21 preprint preprint NN 10_1101-2020_12_31_424971 183 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 183 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 183 24 19 19 CD 10_1101-2020_12_31_424971 183 25 Integrating integrate VBG 10_1101-2020_12_31_424971 183 26 our -PRON- PRP$ 10_1101-2020_12_31_424971 183 27 results result NNS 10_1101-2020_12_31_424971 183 28 with with IN 10_1101-2020_12_31_424971 183 29 the the DT 10_1101-2020_12_31_424971 183 30 available available JJ 10_1101-2020_12_31_424971 183 31 structural structural JJ 10_1101-2020_12_31_424971 183 32 , , , 10_1101-2020_12_31_424971 183 33 biophysical biophysical JJ 10_1101-2020_12_31_424971 183 34 , , , 10_1101-2020_12_31_424971 183 35 and and CC 10_1101-2020_12_31_424971 183 36 biochemical biochemical JJ 10_1101-2020_12_31_424971 183 37 data datum NNS 10_1101-2020_12_31_424971 183 38 on on IN 10_1101-2020_12_31_424971 183 39 the the DT 10_1101-2020_12_31_424971 183 40 457 457 CD 10_1101-2020_12_31_424971 183 41 mechanism mechanism NN 10_1101-2020_12_31_424971 183 42 of of IN 10_1101-2020_12_31_424971 183 43 translation translation NN 10_1101-2020_12_31_424971 183 44 elongation elongation NN 10_1101-2020_12_31_424971 183 45 results result NNS 10_1101-2020_12_31_424971 183 46 in in IN 10_1101-2020_12_31_424971 183 47 the the DT 10_1101-2020_12_31_424971 183 48 structure structure NN 10_1101-2020_12_31_424971 183 49 - - HYPH 10_1101-2020_12_31_424971 183 50 based base VBN 10_1101-2020_12_31_424971 183 51 model model NN 10_1101-2020_12_31_424971 183 52 for for IN 10_1101-2020_12_31_424971 183 53 SufB2-induced SufB2-induced NNP 10_1101-2020_12_31_424971 183 54 +1 +1 NNP 10_1101-2020_12_31_424971 183 55 458 458 CD 10_1101-2020_12_31_424971 183 56 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 183 57 that that WDT 10_1101-2020_12_31_424971 183 58 we -PRON- PRP 10_1101-2020_12_31_424971 183 59 present present VBP 10_1101-2020_12_31_424971 183 60 in in IN 10_1101-2020_12_31_424971 183 61 Figure Figure NNP 10_1101-2020_12_31_424971 183 62 7 7 CD 10_1101-2020_12_31_424971 183 63 . . . 10_1101-2020_12_31_424971 184 1 In in IN 10_1101-2020_12_31_424971 184 2 this this DT 10_1101-2020_12_31_424971 184 3 model model NN 10_1101-2020_12_31_424971 184 4 , , , 10_1101-2020_12_31_424971 184 5 POST POST NNP 10_1101-2020_12_31_424971 184 6 complexes complex NNS 10_1101-2020_12_31_424971 184 7 to to TO 10_1101-2020_12_31_424971 184 8 which which WDT 10_1101-2020_12_31_424971 184 9 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 184 10 or or CC 10_1101-2020_12_31_424971 184 11 ProL ProL NNP 10_1101-2020_12_31_424971 184 12 459 459 CD 10_1101-2020_12_31_424971 184 13 are be VBP 10_1101-2020_12_31_424971 184 14 delivered deliver VBN 10_1101-2020_12_31_424971 184 15 exhibit exhibit NN 10_1101-2020_12_31_424971 184 16 virtually virtually RB 10_1101-2020_12_31_424971 184 17 indistinguishable indistinguishable JJ 10_1101-2020_12_31_424971 184 18 conformational conformational JJ 10_1101-2020_12_31_424971 184 19 dynamics dynamic NNS 10_1101-2020_12_31_424971 184 20 in in IN 10_1101-2020_12_31_424971 184 21 the the DT 10_1101-2020_12_31_424971 184 22 early early JJ 10_1101-2020_12_31_424971 184 23 steps step NNS 10_1101-2020_12_31_424971 184 24 of of IN 10_1101-2020_12_31_424971 184 25 the the DT 10_1101-2020_12_31_424971 184 26 460 460 CD 10_1101-2020_12_31_424971 184 27 elongation elongation NN 10_1101-2020_12_31_424971 184 28 cycle cycle NN 10_1101-2020_12_31_424971 184 29 , , , 10_1101-2020_12_31_424971 184 30 up up IN 10_1101-2020_12_31_424971 184 31 to to IN 10_1101-2020_12_31_424971 184 32 and and CC 10_1101-2020_12_31_424971 184 33 including include VBG 10_1101-2020_12_31_424971 184 34 the the DT 10_1101-2020_12_31_424971 184 35 initial initial JJ 10_1101-2020_12_31_424971 184 36 GS1→GS2 GS1→GS2 NNP 10_1101-2020_12_31_424971 184 37 transition transition NN 10_1101-2020_12_31_424971 184 38 . . . 10_1101-2020_12_31_424971 185 1 However however RB 10_1101-2020_12_31_424971 185 2 , , , 10_1101-2020_12_31_424971 185 3 POST POST NNP 10_1101-2020_12_31_424971 185 4 complexes complex NNS 10_1101-2020_12_31_424971 185 5 461 461 CD 10_1101-2020_12_31_424971 185 6 to to TO 10_1101-2020_12_31_424971 185 7 which which WDT 10_1101-2020_12_31_424971 185 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 185 9 is be VBZ 10_1101-2020_12_31_424971 185 10 delivered deliver VBN 10_1101-2020_12_31_424971 185 11 exhibit exhibit NN 10_1101-2020_12_31_424971 185 12 a a DT 10_1101-2020_12_31_424971 185 13 kGS2→POST kgs2→post NN 10_1101-2020_12_31_424971 185 14 that that WDT 10_1101-2020_12_31_424971 185 15 is be VBZ 10_1101-2020_12_31_424971 185 16 more more JJR 10_1101-2020_12_31_424971 185 17 than than IN 10_1101-2020_12_31_424971 185 18 an an DT 10_1101-2020_12_31_424971 185 19 order order NN 10_1101-2020_12_31_424971 185 20 - - HYPH 10_1101-2020_12_31_424971 185 21 of of IN 10_1101-2020_12_31_424971 185 22 - - HYPH 10_1101-2020_12_31_424971 185 23 magnitude magnitude NN 10_1101-2020_12_31_424971 185 24 slower slow JJR 10_1101-2020_12_31_424971 185 25 462 462 CD 10_1101-2020_12_31_424971 185 26 than than IN 10_1101-2020_12_31_424971 185 27 those those DT 10_1101-2020_12_31_424971 185 28 to to TO 10_1101-2020_12_31_424971 185 29 which which WDT 10_1101-2020_12_31_424971 185 30 ProL ProL NNP 10_1101-2020_12_31_424971 185 31 is be VBZ 10_1101-2020_12_31_424971 185 32 delivered deliver VBN 10_1101-2020_12_31_424971 185 33 . . . 10_1101-2020_12_31_424971 186 1 Notably notably RB 10_1101-2020_12_31_424971 186 2 , , , 10_1101-2020_12_31_424971 186 3 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 186 4 comprises comprise VBZ 10_1101-2020_12_31_424971 186 5 a a DT 10_1101-2020_12_31_424971 186 6 series series NN 10_1101-2020_12_31_424971 186 7 of of IN 10_1101-2020_12_31_424971 186 8 conformational conformational JJ 10_1101-2020_12_31_424971 186 9 463 463 CD 10_1101-2020_12_31_424971 186 10 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 186 11 of of IN 10_1101-2020_12_31_424971 186 12 the the DT 10_1101-2020_12_31_424971 186 13 EF EF NNP 10_1101-2020_12_31_424971 186 14 - - HYPH 10_1101-2020_12_31_424971 186 15 G G NNP 10_1101-2020_12_31_424971 186 16 - - HYPH 10_1101-2020_12_31_424971 186 17 bound bind VBN 10_1101-2020_12_31_424971 186 18 PRE PRE NNP 10_1101-2020_12_31_424971 186 19 complex complex NN 10_1101-2020_12_31_424971 186 20 that that WDT 10_1101-2020_12_31_424971 186 21 facilitate facilitate VBP 10_1101-2020_12_31_424971 186 22 translocation translocation NN 10_1101-2020_12_31_424971 186 23 of of IN 10_1101-2020_12_31_424971 186 24 the the DT 10_1101-2020_12_31_424971 186 25 tRNA trna NN 10_1101-2020_12_31_424971 186 26 ASLs ASLs NNP 10_1101-2020_12_31_424971 186 27 464 464 CD 10_1101-2020_12_31_424971 186 28 and and CC 10_1101-2020_12_31_424971 186 29 associated associated JJ 10_1101-2020_12_31_424971 186 30 codons codon NNS 10_1101-2020_12_31_424971 186 31 within within IN 10_1101-2020_12_31_424971 186 32 the the DT 10_1101-2020_12_31_424971 186 33 30S 30s CD 10_1101-2020_12_31_424971 186 34 subunit subunit NN 10_1101-2020_12_31_424971 186 35 . . . 10_1101-2020_12_31_424971 187 1 These these DT 10_1101-2020_12_31_424971 187 2 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 187 3 encompass encompass VBP 10_1101-2020_12_31_424971 187 4 the the DT 10_1101-2020_12_31_424971 187 5 severing sever VBG 10_1101-2020_12_31_424971 187 6 465 465 CD 10_1101-2020_12_31_424971 187 7 of of IN 10_1101-2020_12_31_424971 187 8 decoding decode VBG 10_1101-2020_12_31_424971 187 9 center center NN 10_1101-2020_12_31_424971 187 10 interactions interaction NNS 10_1101-2020_12_31_424971 187 11 with with IN 10_1101-2020_12_31_424971 187 12 the the DT 10_1101-2020_12_31_424971 187 13 anticodon anticodon NN 10_1101-2020_12_31_424971 187 14 - - HYPH 10_1101-2020_12_31_424971 187 15 codon codon NNP 10_1101-2020_12_31_424971 187 16 duplex duplex NN 10_1101-2020_12_31_424971 187 17 in in IN 10_1101-2020_12_31_424971 187 18 the the DT 10_1101-2020_12_31_424971 187 19 A A NNP 10_1101-2020_12_31_424971 187 20 site22 site22 NNP 10_1101-2020_12_31_424971 187 21 - - SYM 10_1101-2020_12_31_424971 187 22 25 25 CD 10_1101-2020_12_31_424971 187 23 ; ; : 10_1101-2020_12_31_424971 187 24 forward forward RB 10_1101-2020_12_31_424971 187 25 and and CC 10_1101-2020_12_31_424971 187 26 466 466 CD 10_1101-2020_12_31_424971 187 27 reverse reverse JJ 10_1101-2020_12_31_424971 187 28 head head NN 10_1101-2020_12_31_424971 187 29 swiveling27,28,50 swiveling27,28,50 RB 10_1101-2020_12_31_424971 187 30 and and CC 10_1101-2020_12_31_424971 187 31 associated associate VBD 10_1101-2020_12_31_424971 187 32 opening opening NN 10_1101-2020_12_31_424971 187 33 and and CC 10_1101-2020_12_31_424971 187 34 closing closing NN 10_1101-2020_12_31_424971 187 35 , , , 10_1101-2020_12_31_424971 187 36 respectively respectively RB 10_1101-2020_12_31_424971 187 37 , , , 10_1101-2020_12_31_424971 187 38 of of IN 10_1101-2020_12_31_424971 187 39 the the DT 10_1101-2020_12_31_424971 187 40 E e NN 10_1101-2020_12_31_424971 187 41 - - NN 10_1101-2020_12_31_424971 187 42 site site NN 10_1101-2020_12_31_424971 187 43 467 467 CD 10_1101-2020_12_31_424971 187 44 gate26 gate26 NN 10_1101-2020_12_31_424971 187 45 ; ; : 10_1101-2020_12_31_424971 187 46 reverse reverse JJ 10_1101-2020_12_31_424971 187 47 relative relative JJ 10_1101-2020_12_31_424971 187 48 rotation rotation NN 10_1101-2020_12_31_424971 187 49 of of IN 10_1101-2020_12_31_424971 187 50 the the DT 10_1101-2020_12_31_424971 187 51 subunits46,47 subunits46,47 NNP 10_1101-2020_12_31_424971 187 52 ; ; : 10_1101-2020_12_31_424971 187 53 and and CC 10_1101-2020_12_31_424971 187 54 opening opening NN 10_1101-2020_12_31_424971 187 55 of of IN 10_1101-2020_12_31_424971 187 56 the the DT 10_1101-2020_12_31_424971 187 57 L1 L1 NNP 10_1101-2020_12_31_424971 187 58 stalk35,37,48 stalk35,37,48 NN 10_1101-2020_12_31_424971 187 59 ( ( -LRB- 10_1101-2020_12_31_424971 187 60 steps step NNS 10_1101-2020_12_31_424971 187 61 PRE-468 pre-468 UH 10_1101-2020_12_31_424971 187 62 G2 g2 NN 10_1101-2020_12_31_424971 187 63 to to IN 10_1101-2020_12_31_424971 187 64 PRE PRE NNP 10_1101-2020_12_31_424971 187 65 - - HYPH 10_1101-2020_12_31_424971 187 66 G4 G4 NNP 10_1101-2020_12_31_424971 187 67 , , , 10_1101-2020_12_31_424971 187 68 denoted denote VBN 10_1101-2020_12_31_424971 187 69 with with IN 10_1101-2020_12_31_424971 187 70 red red JJ 10_1101-2020_12_31_424971 187 71 arrows arrow NNS 10_1101-2020_12_31_424971 187 72 , , , 10_1101-2020_12_31_424971 187 73 in in IN 10_1101-2020_12_31_424971 187 74 Figure Figure NNP 10_1101-2020_12_31_424971 187 75 7 7 CD 10_1101-2020_12_31_424971 187 76 ) ) -RRB- 10_1101-2020_12_31_424971 187 77 . . . 10_1101-2020_12_31_424971 188 1 Given give VBN 10_1101-2020_12_31_424971 188 2 the the DT 10_1101-2020_12_31_424971 188 3 importance importance NN 10_1101-2020_12_31_424971 188 4 of of IN 10_1101-2020_12_31_424971 188 5 these these DT 10_1101-2020_12_31_424971 188 6 469 469 CD 10_1101-2020_12_31_424971 188 7 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 188 8 in in IN 10_1101-2020_12_31_424971 188 9 translocation translocation NN 10_1101-2020_12_31_424971 188 10 of of IN 10_1101-2020_12_31_424971 188 11 the the DT 10_1101-2020_12_31_424971 188 12 tRNA trna NN 10_1101-2020_12_31_424971 188 13 ASLs asl NNS 10_1101-2020_12_31_424971 188 14 and and CC 10_1101-2020_12_31_424971 188 15 their -PRON- PRP$ 10_1101-2020_12_31_424971 188 16 associated associated JJ 10_1101-2020_12_31_424971 188 17 codons codon NNS 10_1101-2020_12_31_424971 188 18 within within IN 10_1101-2020_12_31_424971 188 19 the the DT 10_1101-2020_12_31_424971 188 20 30S 30S NNP 10_1101-2020_12_31_424971 188 21 470 470 CD 10_1101-2020_12_31_424971 188 22 subunit subunit NN 10_1101-2020_12_31_424971 188 23 , , , 10_1101-2020_12_31_424971 188 24 we -PRON- PRP 10_1101-2020_12_31_424971 188 25 propose propose VBP 10_1101-2020_12_31_424971 188 26 that that IN 10_1101-2020_12_31_424971 188 27 SufB2-mediated sufb2-mediated JJ 10_1101-2020_12_31_424971 188 28 perturbation perturbation NN 10_1101-2020_12_31_424971 188 29 of of IN 10_1101-2020_12_31_424971 188 30 these these DT 10_1101-2020_12_31_424971 188 31 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 188 32 underlies underlie NNS 10_1101-2020_12_31_424971 188 33 +1 +1 : 10_1101-2020_12_31_424971 188 34 471 471 CD 10_1101-2020_12_31_424971 188 35 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 188 36 . . . 10_1101-2020_12_31_424971 189 1 More more RBR 10_1101-2020_12_31_424971 189 2 specifically specifically RB 10_1101-2020_12_31_424971 189 3 , , , 10_1101-2020_12_31_424971 189 4 because because IN 10_1101-2020_12_31_424971 189 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 189 6 does do VBZ 10_1101-2020_12_31_424971 189 7 not not RB 10_1101-2020_12_31_424971 189 8 seem seem VB 10_1101-2020_12_31_424971 189 9 to to TO 10_1101-2020_12_31_424971 189 10 impede impede VB 10_1101-2020_12_31_424971 189 11 the the DT 10_1101-2020_12_31_424971 189 12 reverse reverse JJ 10_1101-2020_12_31_424971 189 13 relative relative JJ 10_1101-2020_12_31_424971 189 14 472 472 CD 10_1101-2020_12_31_424971 189 15 rotation rotation NN 10_1101-2020_12_31_424971 189 16 of of IN 10_1101-2020_12_31_424971 189 17 the the DT 10_1101-2020_12_31_424971 189 18 subunits subunit NNS 10_1101-2020_12_31_424971 189 19 or or CC 10_1101-2020_12_31_424971 189 20 opening opening NN 10_1101-2020_12_31_424971 189 21 of of IN 10_1101-2020_12_31_424971 189 22 the the DT 10_1101-2020_12_31_424971 189 23 L1 L1 NNP 10_1101-2020_12_31_424971 189 24 stalk stalk NN 10_1101-2020_12_31_424971 189 25 during during IN 10_1101-2020_12_31_424971 189 26 the the DT 10_1101-2020_12_31_424971 189 27 GS2→GS1 GS2→GS1 NNP 10_1101-2020_12_31_424971 189 28 transitions transition NNS 10_1101-2020_12_31_424971 189 29 within within IN 10_1101-2020_12_31_424971 189 30 the the DT 10_1101-2020_12_31_424971 189 31 473 473 CD 10_1101-2020_12_31_424971 189 32 GS1 GS1 NNP 10_1101-2020_12_31_424971 189 33 ⇄ ⇄ NNP 10_1101-2020_12_31_424971 189 34 GS2 GS2 NNP 10_1101-2020_12_31_424971 189 35 equilibrium equilibrium NN 10_1101-2020_12_31_424971 189 36 in in IN 10_1101-2020_12_31_424971 189 37 the the DT 10_1101-2020_12_31_424971 189 38 absence absence NN 10_1101-2020_12_31_424971 189 39 of of IN 10_1101-2020_12_31_424971 189 40 EF EF NNP 10_1101-2020_12_31_424971 189 41 - - HYPH 10_1101-2020_12_31_424971 189 42 G G NNP 10_1101-2020_12_31_424971 189 43 ( ( -LRB- 10_1101-2020_12_31_424971 189 44 compare compare VB 10_1101-2020_12_31_424971 189 45 kGS2→GS1 kGS2→GS1 NNP 10_1101-2020_12_31_424971 189 46 for for IN 10_1101-2020_12_31_424971 189 47 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 189 48 vs. vs. IN 10_1101-2020_12_31_424971 189 49 ProL ProL NNP 10_1101-2020_12_31_424971 189 50 - - HYPH 10_1101-2020_12_31_424971 189 51 TC TC NNP 10_1101-2020_12_31_424971 189 52 in in IN 10_1101-2020_12_31_424971 189 53 474 474 CD 10_1101-2020_12_31_424971 189 54 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 189 55 Table Table NNP 10_1101-2020_12_31_424971 189 56 4 4 CD 10_1101-2020_12_31_424971 189 57 ) ) -RRB- 10_1101-2020_12_31_424971 189 58 , , , 10_1101-2020_12_31_424971 189 59 it -PRON- PRP 10_1101-2020_12_31_424971 189 60 most most RBS 10_1101-2020_12_31_424971 189 61 likely likely RB 10_1101-2020_12_31_424971 189 62 interferes interfere VBZ 10_1101-2020_12_31_424971 189 63 with with IN 10_1101-2020_12_31_424971 189 64 the the DT 10_1101-2020_12_31_424971 189 65 severing severing NN 10_1101-2020_12_31_424971 189 66 of of IN 10_1101-2020_12_31_424971 189 67 decoding decode VBG 10_1101-2020_12_31_424971 189 68 center center NN 10_1101-2020_12_31_424971 189 69 interactions interaction NNS 10_1101-2020_12_31_424971 189 70 475 475 CD 10_1101-2020_12_31_424971 189 71 with with IN 10_1101-2020_12_31_424971 189 72 the the DT 10_1101-2020_12_31_424971 189 73 anticodon anticodon NN 10_1101-2020_12_31_424971 189 74 - - HYPH 10_1101-2020_12_31_424971 189 75 codon codon NNP 10_1101-2020_12_31_424971 189 76 duplex duplex NN 10_1101-2020_12_31_424971 189 77 in in IN 10_1101-2020_12_31_424971 189 78 the the DT 10_1101-2020_12_31_424971 189 79 A A NNP 10_1101-2020_12_31_424971 189 80 site site NN 10_1101-2020_12_31_424971 189 81 and/or and/or CC 10_1101-2020_12_31_424971 189 82 forward forward RB 10_1101-2020_12_31_424971 189 83 and/or and/or CC 10_1101-2020_12_31_424971 189 84 reverse reverse VB 10_1101-2020_12_31_424971 189 85 head head NN 10_1101-2020_12_31_424971 189 86 swiveling swivel VBG 10_1101-2020_12_31_424971 189 87 and and CC 10_1101-2020_12_31_424971 189 88 476 476 CD 10_1101-2020_12_31_424971 189 89 associated associate VBN 10_1101-2020_12_31_424971 189 90 opening open VBG 10_1101-2020_12_31_424971 189 91 and/or and/or CC 10_1101-2020_12_31_424971 189 92 closing closing NN 10_1101-2020_12_31_424971 189 93 , , , 10_1101-2020_12_31_424971 189 94 respectively respectively RB 10_1101-2020_12_31_424971 189 95 , , , 10_1101-2020_12_31_424971 189 96 of of IN 10_1101-2020_12_31_424971 189 97 the the DT 10_1101-2020_12_31_424971 189 98 E e NN 10_1101-2020_12_31_424971 189 99 - - NN 10_1101-2020_12_31_424971 189 100 site site NN 10_1101-2020_12_31_424971 189 101 gate gate NN 10_1101-2020_12_31_424971 189 102 . . . 10_1101-2020_12_31_424971 190 1 The the DT 10_1101-2020_12_31_424971 190 2 latter latter JJ 10_1101-2020_12_31_424971 190 3 rearrangement rearrangement NN 10_1101-2020_12_31_424971 190 4 is be VBZ 10_1101-2020_12_31_424971 190 5 477 477 CD 10_1101-2020_12_31_424971 190 6 particularly particularly RB 10_1101-2020_12_31_424971 190 7 important important JJ 10_1101-2020_12_31_424971 190 8 for for IN 10_1101-2020_12_31_424971 190 9 movement movement NN 10_1101-2020_12_31_424971 190 10 of of IN 10_1101-2020_12_31_424971 190 11 the the DT 10_1101-2020_12_31_424971 190 12 tRNA trna NN 10_1101-2020_12_31_424971 190 13 ASLs asl NNS 10_1101-2020_12_31_424971 190 14 and and CC 10_1101-2020_12_31_424971 190 15 their -PRON- PRP$ 10_1101-2020_12_31_424971 190 16 associated associated JJ 10_1101-2020_12_31_424971 190 17 codons codon NNS 10_1101-2020_12_31_424971 190 18 within within IN 10_1101-2020_12_31_424971 190 19 the the DT 10_1101-2020_12_31_424971 190 20 30S 30S NNP 10_1101-2020_12_31_424971 190 21 478 478 CD 10_1101-2020_12_31_424971 190 22 subunit26 subunit26 NNP 10_1101-2020_12_31_424971 190 23 - - SYM 10_1101-2020_12_31_424971 190 24 28,50 28,50 CD 10_1101-2020_12_31_424971 190 25 , , , 10_1101-2020_12_31_424971 190 26 suggesting suggest VBG 10_1101-2020_12_31_424971 190 27 that that IN 10_1101-2020_12_31_424971 190 28 SufB2-mediated sufb2-mediated JJ 10_1101-2020_12_31_424971 190 29 perturbation perturbation NN 10_1101-2020_12_31_424971 190 30 of of IN 10_1101-2020_12_31_424971 190 31 head head NN 10_1101-2020_12_31_424971 190 32 swiveling swiveling NN 10_1101-2020_12_31_424971 190 33 may may MD 10_1101-2020_12_31_424971 190 34 make make VB 10_1101-2020_12_31_424971 190 35 the the DT 10_1101-2020_12_31_424971 190 36 most most JJS 10_1101-2020_12_31_424971 190 37 479 479 CD 10_1101-2020_12_31_424971 190 38 important important JJ 10_1101-2020_12_31_424971 190 39 contribution contribution NN 10_1101-2020_12_31_424971 190 40 to to IN 10_1101-2020_12_31_424971 190 41 +1 +1 NNS 10_1101-2020_12_31_424971 190 42 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 190 43 . . . 10_1101-2020_12_31_424971 191 1 Consistent consistent JJ 10_1101-2020_12_31_424971 191 2 with with IN 10_1101-2020_12_31_424971 191 3 this this DT 10_1101-2020_12_31_424971 191 4 , , , 10_1101-2020_12_31_424971 191 5 a a DT 10_1101-2020_12_31_424971 191 6 recent recent JJ 10_1101-2020_12_31_424971 191 7 structural structural JJ 10_1101-2020_12_31_424971 191 8 study study NN 10_1101-2020_12_31_424971 191 9 showed show VBD 10_1101-2020_12_31_424971 191 10 480 480 CD 10_1101-2020_12_31_424971 191 11 that that IN 10_1101-2020_12_31_424971 191 12 upon upon IN 10_1101-2020_12_31_424971 191 13 forward forward JJ 10_1101-2020_12_31_424971 191 14 head head NN 10_1101-2020_12_31_424971 191 15 swiveling swivel VBG 10_1101-2020_12_31_424971 191 16 , , , 10_1101-2020_12_31_424971 191 17 the the DT 10_1101-2020_12_31_424971 191 18 ASLs asl NNS 10_1101-2020_12_31_424971 191 19 of of IN 10_1101-2020_12_31_424971 191 20 the the DT 10_1101-2020_12_31_424971 191 21 P- P- NNP 10_1101-2020_12_31_424971 191 22 and and CC 10_1101-2020_12_31_424971 191 23 A A NNP 10_1101-2020_12_31_424971 191 24 - - HYPH 10_1101-2020_12_31_424971 191 25 site site NN 10_1101-2020_12_31_424971 191 26 tRNAs tRNAs '' 10_1101-2020_12_31_424971 191 27 can can MD 10_1101-2020_12_31_424971 191 28 disengage disengage VB 10_1101-2020_12_31_424971 191 29 from from IN 10_1101-2020_12_31_424971 191 30 their -PRON- PRP$ 10_1101-2020_12_31_424971 191 31 481 481 CD 10_1101-2020_12_31_424971 191 32 .CC .CC NFP 10_1101-2020_12_31_424971 191 33 - - HYPH 10_1101-2020_12_31_424971 191 34 BY by IN 10_1101-2020_12_31_424971 191 35 - - HYPH 10_1101-2020_12_31_424971 191 36 NC NC NNP 10_1101-2020_12_31_424971 191 37 - - HYPH 10_1101-2020_12_31_424971 191 38 ND ND NNP 10_1101-2020_12_31_424971 191 39 4.0 4.0 CD 10_1101-2020_12_31_424971 191 40 International International NNP 10_1101-2020_12_31_424971 191 41 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 191 42 under under IN 10_1101-2020_12_31_424971 191 43 a a DT 10_1101-2020_12_31_424971 191 44 ( ( -LRB- 10_1101-2020_12_31_424971 191 45 which which WDT 10_1101-2020_12_31_424971 191 46 was be VBD 10_1101-2020_12_31_424971 191 47 not not RB 10_1101-2020_12_31_424971 191 48 certified certify VBN 10_1101-2020_12_31_424971 191 49 by by IN 10_1101-2020_12_31_424971 191 50 peer peer NN 10_1101-2020_12_31_424971 191 51 review review NN 10_1101-2020_12_31_424971 191 52 ) ) -RRB- 10_1101-2020_12_31_424971 191 53 is be VBZ 10_1101-2020_12_31_424971 191 54 the the DT 10_1101-2020_12_31_424971 191 55 author author NN 10_1101-2020_12_31_424971 191 56 / / SYM 10_1101-2020_12_31_424971 191 57 funder funder NN 10_1101-2020_12_31_424971 191 58 , , , 10_1101-2020_12_31_424971 191 59 who who WP 10_1101-2020_12_31_424971 191 60 has have VBZ 10_1101-2020_12_31_424971 191 61 granted grant VBN 10_1101-2020_12_31_424971 191 62 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 191 63 a a DT 10_1101-2020_12_31_424971 191 64 license license NN 10_1101-2020_12_31_424971 191 65 to to TO 10_1101-2020_12_31_424971 191 66 display display VB 10_1101-2020_12_31_424971 191 67 the the DT 10_1101-2020_12_31_424971 191 68 preprint preprint NN 10_1101-2020_12_31_424971 191 69 in in IN 10_1101-2020_12_31_424971 191 70 perpetuity perpetuity NN 10_1101-2020_12_31_424971 191 71 . . . 10_1101-2020_12_31_424971 192 1 It -PRON- PRP 10_1101-2020_12_31_424971 192 2 is be VBZ 10_1101-2020_12_31_424971 192 3 made make VBN 10_1101-2020_12_31_424971 192 4 The the DT 10_1101-2020_12_31_424971 192 5 copyright copyright NN 10_1101-2020_12_31_424971 192 6 holder holder NN 10_1101-2020_12_31_424971 192 7 for for IN 10_1101-2020_12_31_424971 192 8 this this DT 10_1101-2020_12_31_424971 192 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 192 10 version version NN 10_1101-2020_12_31_424971 192 11 posted post VBD 10_1101-2020_12_31_424971 192 12 January January NNP 10_1101-2020_12_31_424971 192 13 2 2 CD 10_1101-2020_12_31_424971 192 14 , , , 10_1101-2020_12_31_424971 192 15 2021 2021 CD 10_1101-2020_12_31_424971 192 16 . . . 10_1101-2020_12_31_424971 192 17 ; ; : 10_1101-2020_12_31_424971 192 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 192 19 : : : 10_1101-2020_12_31_424971 192 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 192 21 preprint preprint NN 10_1101-2020_12_31_424971 192 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 192 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 192 24 20 20 CD 10_1101-2020_12_31_424971 192 25 associated associate VBN 10_1101-2020_12_31_424971 192 26 codons codon NNS 10_1101-2020_12_31_424971 192 27 and and CC 10_1101-2020_12_31_424971 192 28 occupy occupy NN 10_1101-2020_12_31_424971 192 29 positions position NNS 10_1101-2020_12_31_424971 192 30 similar similar JJ 10_1101-2020_12_31_424971 192 31 to to IN 10_1101-2020_12_31_424971 192 32 a a DT 10_1101-2020_12_31_424971 192 33 partial partial JJ 10_1101-2020_12_31_424971 192 34 +1 +1 NNS 10_1101-2020_12_31_424971 192 35 frameshift frameshift NN 10_1101-2020_12_31_424971 192 36 , , , 10_1101-2020_12_31_424971 192 37 even even RB 10_1101-2020_12_31_424971 192 38 in in IN 10_1101-2020_12_31_424971 192 39 the the DT 10_1101-2020_12_31_424971 192 40 presence presence NN 10_1101-2020_12_31_424971 192 41 482 482 CD 10_1101-2020_12_31_424971 192 42 of of IN 10_1101-2020_12_31_424971 192 43 a a DT 10_1101-2020_12_31_424971 192 44 non non JJ 10_1101-2020_12_31_424971 192 45 - - JJ 10_1101-2020_12_31_424971 192 46 frameshift frameshift JJ 10_1101-2020_12_31_424971 192 47 suppressor suppressor NN 10_1101-2020_12_31_424971 192 48 tRNA trna NN 10_1101-2020_12_31_424971 192 49 in in IN 10_1101-2020_12_31_424971 192 50 the the DT 10_1101-2020_12_31_424971 192 51 A A NNP 10_1101-2020_12_31_424971 192 52 site site NN 10_1101-2020_12_31_424971 192 53 and and CC 10_1101-2020_12_31_424971 192 54 the the DT 10_1101-2020_12_31_424971 192 55 absence absence NN 10_1101-2020_12_31_424971 192 56 of of IN 10_1101-2020_12_31_424971 192 57 EF EF NNP 10_1101-2020_12_31_424971 192 58 - - HYPH 10_1101-2020_12_31_424971 192 59 G51 G51 NNP 10_1101-2020_12_31_424971 192 60 . . . 10_1101-2020_12_31_424971 193 1 483 483 CD 10_1101-2020_12_31_424971 193 2 While while IN 10_1101-2020_12_31_424971 193 3 previous previous JJ 10_1101-2020_12_31_424971 193 4 structural structural JJ 10_1101-2020_12_31_424971 193 5 studies study NNS 10_1101-2020_12_31_424971 193 6 have have VBP 10_1101-2020_12_31_424971 193 7 demonstrated demonstrate VBN 10_1101-2020_12_31_424971 193 8 that that IN 10_1101-2020_12_31_424971 193 9 +1 +1 NNS 10_1101-2020_12_31_424971 193 10 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 193 11 tRNAs tRNAs '' 10_1101-2020_12_31_424971 193 12 bind bind NN 10_1101-2020_12_31_424971 193 13 to to IN 10_1101-2020_12_31_424971 193 14 the the DT 10_1101-2020_12_31_424971 193 15 484 484 CD 10_1101-2020_12_31_424971 193 16 A a DT 10_1101-2020_12_31_424971 193 17 site site NN 10_1101-2020_12_31_424971 193 18 in in IN 10_1101-2020_12_31_424971 193 19 the the DT 10_1101-2020_12_31_424971 193 20 0-frame16,17,49 0-frame16,17,49 NN 10_1101-2020_12_31_424971 193 21 and and CC 10_1101-2020_12_31_424971 193 22 to to IN 10_1101-2020_12_31_424971 193 23 the the DT 10_1101-2020_12_31_424971 193 24 P p NN 10_1101-2020_12_31_424971 193 25 site site NN 10_1101-2020_12_31_424971 193 26 in in IN 10_1101-2020_12_31_424971 193 27 the the DT 10_1101-2020_12_31_424971 193 28 +1-frame19 +1-frame19 NN 10_1101-2020_12_31_424971 193 29 , , , 10_1101-2020_12_31_424971 193 30 these these DT 10_1101-2020_12_31_424971 193 31 studies study NNS 10_1101-2020_12_31_424971 193 32 lacked lack VBD 10_1101-2020_12_31_424971 193 33 EF EF NNP 10_1101-2020_12_31_424971 193 34 - - HYPH 10_1101-2020_12_31_424971 193 35 G G NNP 10_1101-2020_12_31_424971 193 36 and and CC 10_1101-2020_12_31_424971 193 37 the the DT 10_1101-2020_12_31_424971 193 38 485 485 CD 10_1101-2020_12_31_424971 193 39 observed observe VBN 10_1101-2020_12_31_424971 193 40 structures structure NNS 10_1101-2020_12_31_424971 193 41 were be VBD 10_1101-2020_12_31_424971 193 42 obtained obtain VBN 10_1101-2020_12_31_424971 193 43 by by IN 10_1101-2020_12_31_424971 193 44 directly directly RB 10_1101-2020_12_31_424971 193 45 binding bind VBG 10_1101-2020_12_31_424971 193 46 a a DT 10_1101-2020_12_31_424971 193 47 deacylated deacylated JJ 10_1101-2020_12_31_424971 193 48 +1 +1 NNS 10_1101-2020_12_31_424971 193 49 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 193 50 tRNA trna NN 10_1101-2020_12_31_424971 193 51 to to IN 10_1101-2020_12_31_424971 193 52 the the DT 10_1101-2020_12_31_424971 193 53 486 486 CD 10_1101-2020_12_31_424971 193 54 P p NN 10_1101-2020_12_31_424971 193 55 site site NN 10_1101-2020_12_31_424971 193 56 . . . 10_1101-2020_12_31_424971 194 1 Specifically specifically RB 10_1101-2020_12_31_424971 194 2 , , , 10_1101-2020_12_31_424971 194 3 a a DT 10_1101-2020_12_31_424971 194 4 +1 +1 NNS 10_1101-2020_12_31_424971 194 5 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 194 6 peptidyl peptidyl NN 10_1101-2020_12_31_424971 194 7 - - HYPH 10_1101-2020_12_31_424971 194 8 tRNA trna NN 10_1101-2020_12_31_424971 194 9 was be VBD 10_1101-2020_12_31_424971 194 10 not not RB 10_1101-2020_12_31_424971 194 11 translocated translocate VBN 10_1101-2020_12_31_424971 194 12 from from IN 10_1101-2020_12_31_424971 194 13 the the DT 10_1101-2020_12_31_424971 194 14 A a NN 10_1101-2020_12_31_424971 194 15 to to IN 10_1101-2020_12_31_424971 194 16 P p NN 10_1101-2020_12_31_424971 194 17 sites site NNS 10_1101-2020_12_31_424971 194 18 , , , 10_1101-2020_12_31_424971 194 19 487 487 CD 10_1101-2020_12_31_424971 194 20 as as IN 10_1101-2020_12_31_424971 194 21 would would MD 10_1101-2020_12_31_424971 194 22 be be VB 10_1101-2020_12_31_424971 194 23 the the DT 10_1101-2020_12_31_424971 194 24 case case NN 10_1101-2020_12_31_424971 194 25 during during IN 10_1101-2020_12_31_424971 194 26 an an DT 10_1101-2020_12_31_424971 194 27 authentic authentic JJ 10_1101-2020_12_31_424971 194 28 translocation translocation NN 10_1101-2020_12_31_424971 194 29 event event NN 10_1101-2020_12_31_424971 194 30 . . . 10_1101-2020_12_31_424971 195 1 In in IN 10_1101-2020_12_31_424971 195 2 contrast contrast NN 10_1101-2020_12_31_424971 195 3 , , , 10_1101-2020_12_31_424971 195 4 our -PRON- PRP$ 10_1101-2020_12_31_424971 195 5 elucidation elucidation NN 10_1101-2020_12_31_424971 195 6 of of IN 10_1101-2020_12_31_424971 195 7 the the DT 10_1101-2020_12_31_424971 195 8 488 488 CD 10_1101-2020_12_31_424971 195 9 +1-frameshifting +1-frameshifting NN 10_1101-2020_12_31_424971 195 10 mechanism mechanism NN 10_1101-2020_12_31_424971 195 11 was be VBD 10_1101-2020_12_31_424971 195 12 executed execute VBN 10_1101-2020_12_31_424971 195 13 in in IN 10_1101-2020_12_31_424971 195 14 the the DT 10_1101-2020_12_31_424971 195 15 presence presence NN 10_1101-2020_12_31_424971 195 16 of of IN 10_1101-2020_12_31_424971 195 17 EF EF NNP 10_1101-2020_12_31_424971 195 18 - - HYPH 10_1101-2020_12_31_424971 195 19 G G NNP 10_1101-2020_12_31_424971 195 20 and and CC 10_1101-2020_12_31_424971 195 21 is be VBZ 10_1101-2020_12_31_424971 195 22 based base VBN 10_1101-2020_12_31_424971 195 23 on on IN 10_1101-2020_12_31_424971 195 24 extensive extensive JJ 10_1101-2020_12_31_424971 195 25 489 489 CD 10_1101-2020_12_31_424971 195 26 comparison comparison NN 10_1101-2020_12_31_424971 195 27 of of IN 10_1101-2020_12_31_424971 195 28 the the DT 10_1101-2020_12_31_424971 195 29 kinetics kinetic NNS 10_1101-2020_12_31_424971 195 30 with with IN 10_1101-2020_12_31_424971 195 31 which which WDT 10_1101-2020_12_31_424971 195 32 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 195 33 and and CC 10_1101-2020_12_31_424971 195 34 ProL ProL NNP 10_1101-2020_12_31_424971 195 35 undergo undergo VBP 10_1101-2020_12_31_424971 195 36 individual individual JJ 10_1101-2020_12_31_424971 195 37 reactions reaction NNS 10_1101-2020_12_31_424971 195 38 of of IN 10_1101-2020_12_31_424971 195 39 the the DT 10_1101-2020_12_31_424971 195 40 490 490 CD 10_1101-2020_12_31_424971 195 41 elongation elongation NN 10_1101-2020_12_31_424971 195 42 cycle cycle NN 10_1101-2020_12_31_424971 195 43 ( ( -LRB- 10_1101-2020_12_31_424971 195 44 i.e. i.e. FW 10_1101-2020_12_31_424971 195 45 , , , 10_1101-2020_12_31_424971 195 46 aa aa NNP 10_1101-2020_12_31_424971 195 47 - - HYPH 10_1101-2020_12_31_424971 195 48 tRNA tRNA NNP 10_1101-2020_12_31_424971 195 49 selection selection NN 10_1101-2020_12_31_424971 195 50 , , , 10_1101-2020_12_31_424971 195 51 peptide peptide NN 10_1101-2020_12_31_424971 195 52 - - HYPH 10_1101-2020_12_31_424971 195 53 bond bond NN 10_1101-2020_12_31_424971 195 54 formation formation NN 10_1101-2020_12_31_424971 195 55 , , , 10_1101-2020_12_31_424971 195 56 and and CC 10_1101-2020_12_31_424971 195 57 translocation translocation NN 10_1101-2020_12_31_424971 195 58 ) ) -RRB- 10_1101-2020_12_31_424971 195 59 and and CC 10_1101-2020_12_31_424971 195 60 the the DT 10_1101-2020_12_31_424971 195 61 491 491 CD 10_1101-2020_12_31_424971 195 62 associated associate VBN 10_1101-2020_12_31_424971 195 63 conformational conformational JJ 10_1101-2020_12_31_424971 195 64 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 195 65 of of IN 10_1101-2020_12_31_424971 195 66 the the DT 10_1101-2020_12_31_424971 195 67 elongation elongation NN 10_1101-2020_12_31_424971 195 68 complex complex NN 10_1101-2020_12_31_424971 195 69 . . . 10_1101-2020_12_31_424971 196 1 Additionally additionally RB 10_1101-2020_12_31_424971 196 2 , , , 10_1101-2020_12_31_424971 196 3 all all DT 10_1101-2020_12_31_424971 196 4 of of IN 10_1101-2020_12_31_424971 196 5 our -PRON- PRP$ 10_1101-2020_12_31_424971 196 6 in in IN 10_1101-2020_12_31_424971 196 7 492 492 CD 10_1101-2020_12_31_424971 196 8 vitro vitro FW 10_1101-2020_12_31_424971 196 9 biochemical biochemical NNP 10_1101-2020_12_31_424971 196 10 assays assay NNS 10_1101-2020_12_31_424971 196 11 , , , 10_1101-2020_12_31_424971 196 12 and and CC 10_1101-2020_12_31_424971 196 13 most most JJS 10_1101-2020_12_31_424971 196 14 of of IN 10_1101-2020_12_31_424971 196 15 our -PRON- PRP$ 10_1101-2020_12_31_424971 196 16 ensemble ensemble JJ 10_1101-2020_12_31_424971 196 17 rapid rapid JJ 10_1101-2020_12_31_424971 196 18 kinetics kinetic NNS 10_1101-2020_12_31_424971 196 19 assays assays RB 10_1101-2020_12_31_424971 196 20 were be VBD 10_1101-2020_12_31_424971 196 21 performed perform VBN 10_1101-2020_12_31_424971 196 22 under under IN 10_1101-2020_12_31_424971 196 23 493 493 CD 10_1101-2020_12_31_424971 196 24 the the DT 10_1101-2020_12_31_424971 196 25 conditions condition NNS 10_1101-2020_12_31_424971 196 26 in in IN 10_1101-2020_12_31_424971 196 27 which which WDT 10_1101-2020_12_31_424971 196 28 the the DT 10_1101-2020_12_31_424971 196 29 A A NNP 10_1101-2020_12_31_424971 196 30 site site NN 10_1101-2020_12_31_424971 196 31 is be VBZ 10_1101-2020_12_31_424971 196 32 always always RB 10_1101-2020_12_31_424971 196 33 occupied occupy VBN 10_1101-2020_12_31_424971 196 34 by by IN 10_1101-2020_12_31_424971 196 35 an an DT 10_1101-2020_12_31_424971 196 36 aa- aa- NN 10_1101-2020_12_31_424971 196 37 or or CC 10_1101-2020_12_31_424971 196 38 peptidyl peptidyl NN 10_1101-2020_12_31_424971 196 39 - - HYPH 10_1101-2020_12_31_424971 196 40 tRNA tRNA NNP 10_1101-2020_12_31_424971 196 41 , , , 10_1101-2020_12_31_424971 196 42 leaving leave VBG 10_1101-2020_12_31_424971 196 43 no no DT 10_1101-2020_12_31_424971 196 44 494 494 CD 10_1101-2020_12_31_424971 196 45 chance chance NN 10_1101-2020_12_31_424971 196 46 of of IN 10_1101-2020_12_31_424971 196 47 a a DT 10_1101-2020_12_31_424971 196 48 vacant vacant JJ 10_1101-2020_12_31_424971 196 49 A a NN 10_1101-2020_12_31_424971 196 50 site site NN 10_1101-2020_12_31_424971 196 51 . . . 10_1101-2020_12_31_424971 197 1 Therefore therefore RB 10_1101-2020_12_31_424971 197 2 , , , 10_1101-2020_12_31_424971 197 3 the the DT 10_1101-2020_12_31_424971 197 4 +1 +1 NNS 10_1101-2020_12_31_424971 197 5 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 197 6 mechanism mechanism NN 10_1101-2020_12_31_424971 197 7 we -PRON- PRP 10_1101-2020_12_31_424971 197 8 present present VBP 10_1101-2020_12_31_424971 197 9 here here RB 10_1101-2020_12_31_424971 197 10 is be VBZ 10_1101-2020_12_31_424971 197 11 distinct distinct JJ 10_1101-2020_12_31_424971 197 12 495 495 CD 10_1101-2020_12_31_424971 197 13 from from IN 10_1101-2020_12_31_424971 197 14 that that DT 10_1101-2020_12_31_424971 197 15 presented present VBN 10_1101-2020_12_31_424971 197 16 by by IN 10_1101-2020_12_31_424971 197 17 Farabaugh Farabaugh NNP 10_1101-2020_12_31_424971 197 18 and and CC 10_1101-2020_12_31_424971 197 19 co co NNP 10_1101-2020_12_31_424971 197 20 - - NN 10_1101-2020_12_31_424971 197 21 workers13 workers13 NNP 10_1101-2020_12_31_424971 197 22 , , , 10_1101-2020_12_31_424971 197 23 in in IN 10_1101-2020_12_31_424971 197 24 which which WDT 10_1101-2020_12_31_424971 197 25 the the DT 10_1101-2020_12_31_424971 197 26 ribosome ribosome NN 10_1101-2020_12_31_424971 197 27 is be VBZ 10_1101-2020_12_31_424971 197 28 stalled stall VBN 10_1101-2020_12_31_424971 197 29 due due JJ 10_1101-2020_12_31_424971 197 30 to to IN 10_1101-2020_12_31_424971 197 31 a a DT 10_1101-2020_12_31_424971 197 32 496 496 CD 10_1101-2020_12_31_424971 197 33 vacant vacant JJ 10_1101-2020_12_31_424971 197 34 A a DT 10_1101-2020_12_31_424971 197 35 site site NN 10_1101-2020_12_31_424971 197 36 , , , 10_1101-2020_12_31_424971 197 37 thus thus RB 10_1101-2020_12_31_424971 197 38 giving give VBG 10_1101-2020_12_31_424971 197 39 the the DT 10_1101-2020_12_31_424971 197 40 +1-frameshifting +1-frameshifting NN 10_1101-2020_12_31_424971 197 41 - - HYPH 10_1101-2020_12_31_424971 197 42 inducing induce VBG 10_1101-2020_12_31_424971 197 43 tRNA trna NN 10_1101-2020_12_31_424971 197 44 at at IN 10_1101-2020_12_31_424971 197 45 the the DT 10_1101-2020_12_31_424971 197 46 P p NN 10_1101-2020_12_31_424971 197 47 site site NN 10_1101-2020_12_31_424971 197 48 an an DT 10_1101-2020_12_31_424971 197 49 opportunity opportunity NN 10_1101-2020_12_31_424971 197 50 to to TO 10_1101-2020_12_31_424971 197 51 497 497 CD 10_1101-2020_12_31_424971 197 52 rearrange rearrange NN 10_1101-2020_12_31_424971 197 53 into into IN 10_1101-2020_12_31_424971 197 54 the the DT 10_1101-2020_12_31_424971 197 55 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 197 56 . . . 10_1101-2020_12_31_424971 198 1 The the DT 10_1101-2020_12_31_424971 198 2 fact fact NN 10_1101-2020_12_31_424971 198 3 that that IN 10_1101-2020_12_31_424971 198 4 all all DT 10_1101-2020_12_31_424971 198 5 well well RB 10_1101-2020_12_31_424971 198 6 - - HYPH 10_1101-2020_12_31_424971 198 7 characterized characterize VBN 10_1101-2020_12_31_424971 198 8 +1-frameshifting +1-frameshifting PRP$ 10_1101-2020_12_31_424971 198 9 tRNAs trnas CD 10_1101-2020_12_31_424971 198 10 contain contain NN 10_1101-2020_12_31_424971 198 11 498 498 CD 10_1101-2020_12_31_424971 198 12 an an DT 10_1101-2020_12_31_424971 198 13 extra extra JJ 10_1101-2020_12_31_424971 198 14 nucleotide nucleotide NN 10_1101-2020_12_31_424971 198 15 in in IN 10_1101-2020_12_31_424971 198 16 the the DT 10_1101-2020_12_31_424971 198 17 anticodon anticodon NNP 10_1101-2020_12_31_424971 198 18 loop loop NN 10_1101-2020_12_31_424971 198 19 , , , 10_1101-2020_12_31_424971 198 20 despite despite IN 10_1101-2020_12_31_424971 198 21 differences difference NNS 10_1101-2020_12_31_424971 198 22 in in IN 10_1101-2020_12_31_424971 198 23 their -PRON- PRP$ 10_1101-2020_12_31_424971 198 24 primary primary JJ 10_1101-2020_12_31_424971 198 25 sequences sequence NNS 10_1101-2020_12_31_424971 198 26 , , , 10_1101-2020_12_31_424971 198 27 the the DT 10_1101-2020_12_31_424971 198 28 499 499 CD 10_1101-2020_12_31_424971 198 29 amino amino JJ 10_1101-2020_12_31_424971 198 30 acids acid NNS 10_1101-2020_12_31_424971 198 31 they -PRON- PRP 10_1101-2020_12_31_424971 198 32 carry carry VBP 10_1101-2020_12_31_424971 198 33 , , , 10_1101-2020_12_31_424971 198 34 and and CC 10_1101-2020_12_31_424971 198 35 whether whether IN 10_1101-2020_12_31_424971 198 36 the the DT 10_1101-2020_12_31_424971 198 37 extra extra JJ 10_1101-2020_12_31_424971 198 38 nucleotide nucleotide NN 10_1101-2020_12_31_424971 198 39 is be VBZ 10_1101-2020_12_31_424971 198 40 inserted insert VBN 10_1101-2020_12_31_424971 198 41 at at IN 10_1101-2020_12_31_424971 198 42 the the DT 10_1101-2020_12_31_424971 198 43 3'- 3'- NNP 10_1101-2020_12_31_424971 198 44 or or CC 10_1101-2020_12_31_424971 198 45 5'-sides 5'-sides NNPS 10_1101-2020_12_31_424971 198 46 of of IN 10_1101-2020_12_31_424971 198 47 the the DT 10_1101-2020_12_31_424971 198 48 500 500 CD 10_1101-2020_12_31_424971 198 49 anticodon anticodon NN 10_1101-2020_12_31_424971 198 50 , , , 10_1101-2020_12_31_424971 198 51 suggests suggest VBZ 10_1101-2020_12_31_424971 198 52 that that IN 10_1101-2020_12_31_424971 198 53 the the DT 10_1101-2020_12_31_424971 198 54 results result NNS 10_1101-2020_12_31_424971 198 55 we -PRON- PRP 10_1101-2020_12_31_424971 198 56 report report VBP 10_1101-2020_12_31_424971 198 57 here here RB 10_1101-2020_12_31_424971 198 58 for for IN 10_1101-2020_12_31_424971 198 59 SufB2 sufb2 NN 10_1101-2020_12_31_424971 198 60 are be VBP 10_1101-2020_12_31_424971 198 61 likely likely RB 10_1101-2020_12_31_424971 198 62 applicable applicable JJ 10_1101-2020_12_31_424971 198 63 to to IN 10_1101-2020_12_31_424971 198 64 other other JJ 10_1101-2020_12_31_424971 198 65 +1 +1 : 10_1101-2020_12_31_424971 198 66 - - : 10_1101-2020_12_31_424971 198 67 501 501 CD 10_1101-2020_12_31_424971 198 68 frameshifting frameshifting NN 10_1101-2020_12_31_424971 198 69 tRNAs trnas CD 10_1101-2020_12_31_424971 198 70 with with IN 10_1101-2020_12_31_424971 198 71 an an DT 10_1101-2020_12_31_424971 198 72 expanded expand VBN 10_1101-2020_12_31_424971 198 73 anticodon anticodon NNP 10_1101-2020_12_31_424971 198 74 loop loop NN 10_1101-2020_12_31_424971 198 75 . . . 10_1101-2020_12_31_424971 199 1 502 502 CD 10_1101-2020_12_31_424971 199 2 While while IN 10_1101-2020_12_31_424971 199 3 an an DT 10_1101-2020_12_31_424971 199 4 expanded expand VBN 10_1101-2020_12_31_424971 199 5 anticodon anticodon NNP 10_1101-2020_12_31_424971 199 6 loop loop NN 10_1101-2020_12_31_424971 199 7 is be VBZ 10_1101-2020_12_31_424971 199 8 a a DT 10_1101-2020_12_31_424971 199 9 strong strong JJ 10_1101-2020_12_31_424971 199 10 feature feature NN 10_1101-2020_12_31_424971 199 11 associated associate VBN 10_1101-2020_12_31_424971 199 12 with with IN 10_1101-2020_12_31_424971 199 13 +1 +1 NNS 10_1101-2020_12_31_424971 199 14 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 199 15 , , , 10_1101-2020_12_31_424971 199 16 it -PRON- PRP 10_1101-2020_12_31_424971 199 17 is be VBZ 10_1101-2020_12_31_424971 199 18 503 503 CD 10_1101-2020_12_31_424971 199 19 not not RB 10_1101-2020_12_31_424971 199 20 associated associate VBN 10_1101-2020_12_31_424971 199 21 with with IN 10_1101-2020_12_31_424971 199 22 – – : 10_1101-2020_12_31_424971 199 23 1 1 CD 10_1101-2020_12_31_424971 199 24 frameshifting frameshifting NN 10_1101-2020_12_31_424971 199 25 , , , 10_1101-2020_12_31_424971 199 26 which which WDT 10_1101-2020_12_31_424971 199 27 instead instead RB 10_1101-2020_12_31_424971 199 28 is be VBZ 10_1101-2020_12_31_424971 199 29 typically typically RB 10_1101-2020_12_31_424971 199 30 induced induce VBN 10_1101-2020_12_31_424971 199 31 by by IN 10_1101-2020_12_31_424971 199 32 structural structural JJ 10_1101-2020_12_31_424971 199 33 barriers barrier NNS 10_1101-2020_12_31_424971 199 34 in in IN 10_1101-2020_12_31_424971 199 35 504 504 CD 10_1101-2020_12_31_424971 199 36 the the DT 10_1101-2020_12_31_424971 199 37 mRNA mrna NN 10_1101-2020_12_31_424971 199 38 that that WDT 10_1101-2020_12_31_424971 199 39 stall stall VBP 10_1101-2020_12_31_424971 199 40 a a DT 10_1101-2020_12_31_424971 199 41 translating translating NN 10_1101-2020_12_31_424971 199 42 ribosome ribosome NN 10_1101-2020_12_31_424971 199 43 from from IN 10_1101-2020_12_31_424971 199 44 moving move VBG 10_1101-2020_12_31_424971 199 45 forward forward RB 10_1101-2020_12_31_424971 199 46 , , , 10_1101-2020_12_31_424971 199 47 thus thus RB 10_1101-2020_12_31_424971 199 48 providing provide VBG 10_1101-2020_12_31_424971 199 49 the the DT 10_1101-2020_12_31_424971 199 50 ribosome ribosome NN 10_1101-2020_12_31_424971 199 51 with with IN 10_1101-2020_12_31_424971 199 52 505 505 CD 10_1101-2020_12_31_424971 199 53 an an DT 10_1101-2020_12_31_424971 199 54 opportunity opportunity NN 10_1101-2020_12_31_424971 199 55 to to TO 10_1101-2020_12_31_424971 199 56 shift shift VB 10_1101-2020_12_31_424971 199 57 backwards backwards RB 10_1101-2020_12_31_424971 199 58 in in IN 10_1101-2020_12_31_424971 199 59 the the DT 10_1101-2020_12_31_424971 199 60 – – : 10_1101-2020_12_31_424971 199 61 1 1 CD 10_1101-2020_12_31_424971 199 62 direction10,52 direction10,52 RB 10_1101-2020_12_31_424971 199 63 . . . 10_1101-2020_12_31_424971 200 1 Given give VBN 10_1101-2020_12_31_424971 200 2 the the DT 10_1101-2020_12_31_424971 200 3 unique unique JJ 10_1101-2020_12_31_424971 200 4 role role NN 10_1101-2020_12_31_424971 200 5 of of IN 10_1101-2020_12_31_424971 200 6 the the DT 10_1101-2020_12_31_424971 200 7 expanded expand VBN 10_1101-2020_12_31_424971 200 8 506 506 CD 10_1101-2020_12_31_424971 200 9 anticodon anticodon NNP 10_1101-2020_12_31_424971 200 10 loop loop NN 10_1101-2020_12_31_424971 200 11 in in IN 10_1101-2020_12_31_424971 200 12 +1 +1 NNS 10_1101-2020_12_31_424971 200 13 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 200 14 , , , 10_1101-2020_12_31_424971 200 15 here here RB 10_1101-2020_12_31_424971 200 16 we -PRON- PRP 10_1101-2020_12_31_424971 200 17 have have VBP 10_1101-2020_12_31_424971 200 18 identified identify VBN 10_1101-2020_12_31_424971 200 19 the the DT 10_1101-2020_12_31_424971 200 20 determinants determinant NNS 10_1101-2020_12_31_424971 200 21 that that WDT 10_1101-2020_12_31_424971 200 22 drive drive VBP 10_1101-2020_12_31_424971 200 23 the the DT 10_1101-2020_12_31_424971 200 24 507 507 CD 10_1101-2020_12_31_424971 200 25 .CC .CC NFP 10_1101-2020_12_31_424971 200 26 - - : 10_1101-2020_12_31_424971 200 27 BY by IN 10_1101-2020_12_31_424971 200 28 - - HYPH 10_1101-2020_12_31_424971 200 29 NC NC NNP 10_1101-2020_12_31_424971 200 30 - - HYPH 10_1101-2020_12_31_424971 200 31 ND ND NNP 10_1101-2020_12_31_424971 200 32 4.0 4.0 CD 10_1101-2020_12_31_424971 200 33 International International NNP 10_1101-2020_12_31_424971 200 34 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 200 35 under under IN 10_1101-2020_12_31_424971 200 36 a a DT 10_1101-2020_12_31_424971 200 37 ( ( -LRB- 10_1101-2020_12_31_424971 200 38 which which WDT 10_1101-2020_12_31_424971 200 39 was be VBD 10_1101-2020_12_31_424971 200 40 not not RB 10_1101-2020_12_31_424971 200 41 certified certify VBN 10_1101-2020_12_31_424971 200 42 by by IN 10_1101-2020_12_31_424971 200 43 peer peer NN 10_1101-2020_12_31_424971 200 44 review review NN 10_1101-2020_12_31_424971 200 45 ) ) -RRB- 10_1101-2020_12_31_424971 200 46 is be VBZ 10_1101-2020_12_31_424971 200 47 the the DT 10_1101-2020_12_31_424971 200 48 author author NN 10_1101-2020_12_31_424971 200 49 / / SYM 10_1101-2020_12_31_424971 200 50 funder funder NN 10_1101-2020_12_31_424971 200 51 , , , 10_1101-2020_12_31_424971 200 52 who who WP 10_1101-2020_12_31_424971 200 53 has have VBZ 10_1101-2020_12_31_424971 200 54 granted grant VBN 10_1101-2020_12_31_424971 200 55 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 200 56 a a DT 10_1101-2020_12_31_424971 200 57 license license NN 10_1101-2020_12_31_424971 200 58 to to TO 10_1101-2020_12_31_424971 200 59 display display VB 10_1101-2020_12_31_424971 200 60 the the DT 10_1101-2020_12_31_424971 200 61 preprint preprint NN 10_1101-2020_12_31_424971 200 62 in in IN 10_1101-2020_12_31_424971 200 63 perpetuity perpetuity NN 10_1101-2020_12_31_424971 200 64 . . . 10_1101-2020_12_31_424971 201 1 It -PRON- PRP 10_1101-2020_12_31_424971 201 2 is be VBZ 10_1101-2020_12_31_424971 201 3 made make VBN 10_1101-2020_12_31_424971 201 4 The the DT 10_1101-2020_12_31_424971 201 5 copyright copyright NN 10_1101-2020_12_31_424971 201 6 holder holder NN 10_1101-2020_12_31_424971 201 7 for for IN 10_1101-2020_12_31_424971 201 8 this this DT 10_1101-2020_12_31_424971 201 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 201 10 version version NN 10_1101-2020_12_31_424971 201 11 posted post VBD 10_1101-2020_12_31_424971 201 12 January January NNP 10_1101-2020_12_31_424971 201 13 2 2 CD 10_1101-2020_12_31_424971 201 14 , , , 10_1101-2020_12_31_424971 201 15 2021 2021 CD 10_1101-2020_12_31_424971 201 16 . . . 10_1101-2020_12_31_424971 201 17 ; ; : 10_1101-2020_12_31_424971 201 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 201 19 : : : 10_1101-2020_12_31_424971 201 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 201 21 preprint preprint NN 10_1101-2020_12_31_424971 201 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 201 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 201 24 21 21 CD 10_1101-2020_12_31_424971 201 25 ribosome ribosome NN 10_1101-2020_12_31_424971 201 26 to to TO 10_1101-2020_12_31_424971 201 27 shift shift VB 10_1101-2020_12_31_424971 201 28 in in IN 10_1101-2020_12_31_424971 201 29 the the DT 10_1101-2020_12_31_424971 201 30 +1 +1 NNS 10_1101-2020_12_31_424971 201 31 direction direction NN 10_1101-2020_12_31_424971 201 32 . . . 10_1101-2020_12_31_424971 202 1 We -PRON- PRP 10_1101-2020_12_31_424971 202 2 show show VBP 10_1101-2020_12_31_424971 202 3 that that IN 10_1101-2020_12_31_424971 202 4 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 202 5 exclusively exclusively RB 10_1101-2020_12_31_424971 202 6 uses use VBZ 10_1101-2020_12_31_424971 202 7 the the DT 10_1101-2020_12_31_424971 202 8 triplet triplet NN 10_1101-2020_12_31_424971 202 9 - - HYPH 10_1101-2020_12_31_424971 202 10 slippage slippage NN 10_1101-2020_12_31_424971 202 11 508 508 CD 10_1101-2020_12_31_424971 202 12 mechanism mechanism NN 10_1101-2020_12_31_424971 202 13 of of IN 10_1101-2020_12_31_424971 202 14 +1 +1 NNS 10_1101-2020_12_31_424971 202 15 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 202 16 in in IN 10_1101-2020_12_31_424971 202 17 the the DT 10_1101-2020_12_31_424971 202 18 m1G37 m1g37 NN 10_1101-2020_12_31_424971 202 19 + + CC 10_1101-2020_12_31_424971 202 20 condition condition NN 10_1101-2020_12_31_424971 202 21 , , , 10_1101-2020_12_31_424971 202 22 but but CC 10_1101-2020_12_31_424971 202 23 that that IN 10_1101-2020_12_31_424971 202 24 it -PRON- PRP 10_1101-2020_12_31_424971 202 25 explores explore VBZ 10_1101-2020_12_31_424971 202 26 other other JJ 10_1101-2020_12_31_424971 202 27 mechanisms mechanism NNS 10_1101-2020_12_31_424971 202 28 509 509 CD 10_1101-2020_12_31_424971 202 29 ( ( -LRB- 10_1101-2020_12_31_424971 202 30 e.g. e.g. RB 10_1101-2020_12_31_424971 202 31 , , , 10_1101-2020_12_31_424971 202 32 quadruplet quadruplet NN 10_1101-2020_12_31_424971 202 33 - - HYPH 10_1101-2020_12_31_424971 202 34 pairing pairing NN 10_1101-2020_12_31_424971 202 35 ) ) -RRB- 10_1101-2020_12_31_424971 202 36 in in IN 10_1101-2020_12_31_424971 202 37 the the DT 10_1101-2020_12_31_424971 202 38 m1G37 m1g37 NN 10_1101-2020_12_31_424971 202 39 – – : 10_1101-2020_12_31_424971 202 40 condition condition NN 10_1101-2020_12_31_424971 202 41 during during IN 10_1101-2020_12_31_424971 202 42 translocation translocation NN 10_1101-2020_12_31_424971 202 43 from from IN 10_1101-2020_12_31_424971 202 44 the the DT 10_1101-2020_12_31_424971 202 45 A A NNP 10_1101-2020_12_31_424971 202 46 site site NN 10_1101-2020_12_31_424971 202 47 to to IN 10_1101-2020_12_31_424971 202 48 the the DT 10_1101-2020_12_31_424971 202 49 P p NN 10_1101-2020_12_31_424971 202 50 510 510 CD 10_1101-2020_12_31_424971 202 51 site site NN 10_1101-2020_12_31_424971 202 52 . . . 10_1101-2020_12_31_424971 203 1 Under under IN 10_1101-2020_12_31_424971 203 2 conditions condition NNS 10_1101-2020_12_31_424971 203 3 that that WDT 10_1101-2020_12_31_424971 203 4 only only RB 10_1101-2020_12_31_424971 203 5 permit permit VBP 10_1101-2020_12_31_424971 203 6 the the DT 10_1101-2020_12_31_424971 203 7 triplet triplet NN 10_1101-2020_12_31_424971 203 8 - - HYPH 10_1101-2020_12_31_424971 203 9 slippage slippage NN 10_1101-2020_12_31_424971 203 10 mechanism mechanism NN 10_1101-2020_12_31_424971 203 11 ( ( -LRB- 10_1101-2020_12_31_424971 203 12 e.g. e.g. RB 10_1101-2020_12_31_424971 203 13 , , , 10_1101-2020_12_31_424971 203 14 in in IN 10_1101-2020_12_31_424971 203 15 the the DT 10_1101-2020_12_31_424971 203 16 presence presence NN 10_1101-2020_12_31_424971 203 17 of of IN 10_1101-2020_12_31_424971 203 18 511 511 CD 10_1101-2020_12_31_424971 203 19 m1G37 m1g37 NN 10_1101-2020_12_31_424971 203 20 ) ) -RRB- 10_1101-2020_12_31_424971 203 21 , , , 10_1101-2020_12_31_424971 203 22 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 203 23 exhibits exhibit VBZ 10_1101-2020_12_31_424971 203 24 a a DT 10_1101-2020_12_31_424971 203 25 relatively relatively RB 10_1101-2020_12_31_424971 203 26 low low JJ 10_1101-2020_12_31_424971 203 27 +1-frameshifting +1-frameshifting JJ 10_1101-2020_12_31_424971 203 28 efficiency efficiency NN 10_1101-2020_12_31_424971 203 29 of of IN 10_1101-2020_12_31_424971 203 30 ~30 ~30 $ 10_1101-2020_12_31_424971 203 31 % % NN 10_1101-2020_12_31_424971 203 32 , , , 10_1101-2020_12_31_424971 203 33 whereas whereas IN 10_1101-2020_12_31_424971 203 34 under under IN 10_1101-2020_12_31_424971 203 35 512 512 CD 10_1101-2020_12_31_424971 203 36 conditions condition NNS 10_1101-2020_12_31_424971 203 37 that that WDT 10_1101-2020_12_31_424971 203 38 permit permit VBP 10_1101-2020_12_31_424971 203 39 quadruplet quadruplet NN 10_1101-2020_12_31_424971 203 40 - - HYPH 10_1101-2020_12_31_424971 203 41 pairing pair VBG 10_1101-2020_12_31_424971 203 42 during during IN 10_1101-2020_12_31_424971 203 43 translocation translocation NN 10_1101-2020_12_31_424971 203 44 ( ( -LRB- 10_1101-2020_12_31_424971 203 45 e.g. e.g. RB 10_1101-2020_12_31_424971 203 46 , , , 10_1101-2020_12_31_424971 203 47 in in IN 10_1101-2020_12_31_424971 203 48 the the DT 10_1101-2020_12_31_424971 203 49 absence absence NN 10_1101-2020_12_31_424971 203 50 of of IN 10_1101-2020_12_31_424971 203 51 m1G37 m1g37 NN 10_1101-2020_12_31_424971 203 52 ) ) -RRB- 10_1101-2020_12_31_424971 203 53 , , , 10_1101-2020_12_31_424971 203 54 it -PRON- PRP 10_1101-2020_12_31_424971 203 55 513 513 CD 10_1101-2020_12_31_424971 203 56 exhibits exhibit VBZ 10_1101-2020_12_31_424971 203 57 a a DT 10_1101-2020_12_31_424971 203 58 relatively relatively RB 10_1101-2020_12_31_424971 203 59 high high JJ 10_1101-2020_12_31_424971 203 60 +1-frameshifting +1-frameshifting JJ 10_1101-2020_12_31_424971 203 61 efficiency efficiency NN 10_1101-2020_12_31_424971 203 62 of of IN 10_1101-2020_12_31_424971 203 63 ~90 ~90 NFP 10_1101-2020_12_31_424971 203 64 % % NN 10_1101-2020_12_31_424971 203 65 ( ( -LRB- 10_1101-2020_12_31_424971 203 66 Figures Figures NNP 10_1101-2020_12_31_424971 203 67 4c 4c NNP 10_1101-2020_12_31_424971 203 68 - - HYPH 10_1101-2020_12_31_424971 203 69 f f NN 10_1101-2020_12_31_424971 203 70 , , , 10_1101-2020_12_31_424971 203 71 5a 5a NNP 10_1101-2020_12_31_424971 203 72 ) ) -RRB- 10_1101-2020_12_31_424971 203 73 . . . 10_1101-2020_12_31_424971 204 1 This this DT 10_1101-2020_12_31_424971 204 2 feature feature NN 10_1101-2020_12_31_424971 204 3 is be VBZ 10_1101-2020_12_31_424971 204 4 514 514 CD 10_1101-2020_12_31_424971 204 5 observed observe VBN 10_1101-2020_12_31_424971 204 6 in in IN 10_1101-2020_12_31_424971 204 7 various various JJ 10_1101-2020_12_31_424971 204 8 sequence sequence NN 10_1101-2020_12_31_424971 204 9 contexts contexts NN 10_1101-2020_12_31_424971 204 10 . . . 10_1101-2020_12_31_424971 205 1 One one CD 10_1101-2020_12_31_424971 205 2 advantage advantage NN 10_1101-2020_12_31_424971 205 3 of of IN 10_1101-2020_12_31_424971 205 4 a a DT 10_1101-2020_12_31_424971 205 5 quadruplet quadruplet NN 10_1101-2020_12_31_424971 205 6 - - HYPH 10_1101-2020_12_31_424971 205 7 pairing pair VBG 10_1101-2020_12_31_424971 205 8 mechanism mechanism NN 10_1101-2020_12_31_424971 205 9 515 515 CD 10_1101-2020_12_31_424971 205 10 during during IN 10_1101-2020_12_31_424971 205 11 translocation translocation NN 10_1101-2020_12_31_424971 205 12 is be VBZ 10_1101-2020_12_31_424971 205 13 that that IN 10_1101-2020_12_31_424971 205 14 it -PRON- PRP 10_1101-2020_12_31_424971 205 15 would would MD 10_1101-2020_12_31_424971 205 16 enhance enhance VB 10_1101-2020_12_31_424971 205 17 the the DT 10_1101-2020_12_31_424971 205 18 thermodynamic thermodynamic JJ 10_1101-2020_12_31_424971 205 19 stability stability NN 10_1101-2020_12_31_424971 205 20 of of IN 10_1101-2020_12_31_424971 205 21 anticodon anticodon NNP 10_1101-2020_12_31_424971 205 22 - - HYPH 10_1101-2020_12_31_424971 205 23 codon codon NNP 10_1101-2020_12_31_424971 205 24 516 516 CD 10_1101-2020_12_31_424971 205 25 pairing pairing NN 10_1101-2020_12_31_424971 205 26 during during IN 10_1101-2020_12_31_424971 205 27 the the DT 10_1101-2020_12_31_424971 205 28 large large JJ 10_1101-2020_12_31_424971 205 29 EF EF NNP 10_1101-2020_12_31_424971 205 30 - - HYPH 10_1101-2020_12_31_424971 205 31 G G NNP 10_1101-2020_12_31_424971 205 32 - - HYPH 10_1101-2020_12_31_424971 205 33 catalyzed catalyzed JJ 10_1101-2020_12_31_424971 205 34 conformational conformational JJ 10_1101-2020_12_31_424971 205 35 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 205 36 that that IN 10_1101-2020_12_31_424971 205 37 PRE PRE NNP 10_1101-2020_12_31_424971 205 38 complexes complex NNS 10_1101-2020_12_31_424971 205 39 517 517 CD 10_1101-2020_12_31_424971 205 40 undergo undergo VBP 10_1101-2020_12_31_424971 205 41 during during IN 10_1101-2020_12_31_424971 205 42 translocation translocation NN 10_1101-2020_12_31_424971 205 43 to to TO 10_1101-2020_12_31_424971 205 44 form form VB 10_1101-2020_12_31_424971 205 45 POST POST NNP 10_1101-2020_12_31_424971 205 46 complexes complex NNS 10_1101-2020_12_31_424971 205 47 . . . 10_1101-2020_12_31_424971 206 1 Nonetheless nonetheless RB 10_1101-2020_12_31_424971 206 2 , , , 10_1101-2020_12_31_424971 206 3 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 206 4 is be VBZ 10_1101-2020_12_31_424971 206 5 naturally naturally RB 10_1101-2020_12_31_424971 206 6 518 518 CD 10_1101-2020_12_31_424971 206 7 methylated methylate VBN 10_1101-2020_12_31_424971 206 8 with with IN 10_1101-2020_12_31_424971 206 9 m1G37 m1g37 NN 10_1101-2020_12_31_424971 206 10 ( ( -LRB- 10_1101-2020_12_31_424971 206 11 Figure Figure NNP 10_1101-2020_12_31_424971 206 12 1c 1c NN 10_1101-2020_12_31_424971 206 13 ) ) -RRB- 10_1101-2020_12_31_424971 206 14 , , , 10_1101-2020_12_31_424971 206 15 indicating indicate VBG 10_1101-2020_12_31_424971 206 16 that that IN 10_1101-2020_12_31_424971 206 17 it -PRON- PRP 10_1101-2020_12_31_424971 206 18 makes make VBZ 10_1101-2020_12_31_424971 206 19 exclusive exclusive JJ 10_1101-2020_12_31_424971 206 20 use use NN 10_1101-2020_12_31_424971 206 21 of of IN 10_1101-2020_12_31_424971 206 22 the the DT 10_1101-2020_12_31_424971 206 23 triplet triplet NN 10_1101-2020_12_31_424971 206 24 - - HYPH 10_1101-2020_12_31_424971 206 25 slippage slippage NN 10_1101-2020_12_31_424971 206 26 519 519 CD 10_1101-2020_12_31_424971 206 27 mechanism mechanism NN 10_1101-2020_12_31_424971 206 28 in in IN 10_1101-2020_12_31_424971 206 29 vivo vivo NNP 10_1101-2020_12_31_424971 206 30 . . . 10_1101-2020_12_31_424971 207 1 This this DT 10_1101-2020_12_31_424971 207 2 mechanism mechanism NN 10_1101-2020_12_31_424971 207 3 is be VBZ 10_1101-2020_12_31_424971 207 4 likely likely JJ 10_1101-2020_12_31_424971 207 5 also also RB 10_1101-2020_12_31_424971 207 6 exclusively exclusively RB 10_1101-2020_12_31_424971 207 7 used use VBN 10_1101-2020_12_31_424971 207 8 in in IN 10_1101-2020_12_31_424971 207 9 vivo vivo NNP 10_1101-2020_12_31_424971 207 10 by by IN 10_1101-2020_12_31_424971 207 11 all all DT 10_1101-2020_12_31_424971 207 12 other other JJ 10_1101-2020_12_31_424971 207 13 +1 +1 : 10_1101-2020_12_31_424971 207 14 - - : 10_1101-2020_12_31_424971 207 15 520 520 CD 10_1101-2020_12_31_424971 207 16 frameshifting frameshifting NN 10_1101-2020_12_31_424971 207 17 tRNAs trnas CD 10_1101-2020_12_31_424971 207 18 that that WDT 10_1101-2020_12_31_424971 207 19 have have VBP 10_1101-2020_12_31_424971 207 20 evolved evolve VBN 10_1101-2020_12_31_424971 207 21 from from IN 10_1101-2020_12_31_424971 207 22 canonical canonical JJ 10_1101-2020_12_31_424971 207 23 tRNAs trnas CD 10_1101-2020_12_31_424971 207 24 to to TO 10_1101-2020_12_31_424971 207 25 retain retain VB 10_1101-2020_12_31_424971 207 26 a a DT 10_1101-2020_12_31_424971 207 27 purine purine NN 10_1101-2020_12_31_424971 207 28 at at IN 10_1101-2020_12_31_424971 207 29 position position NN 10_1101-2020_12_31_424971 207 30 37 37 CD 10_1101-2020_12_31_424971 207 31 , , , 10_1101-2020_12_31_424971 207 32 521 521 CD 10_1101-2020_12_31_424971 207 33 which which WDT 10_1101-2020_12_31_424971 207 34 is be VBZ 10_1101-2020_12_31_424971 207 35 almost almost RB 10_1101-2020_12_31_424971 207 36 universally universally RB 10_1101-2020_12_31_424971 207 37 post post JJ 10_1101-2020_12_31_424971 207 38 - - JJ 10_1101-2020_12_31_424971 207 39 transcriptionally transcriptionally RB 10_1101-2020_12_31_424971 207 40 modified modify VBN 10_1101-2020_12_31_424971 207 41 to to TO 10_1101-2020_12_31_424971 207 42 block block VB 10_1101-2020_12_31_424971 207 43 quadruplet quadruplet NN 10_1101-2020_12_31_424971 207 44 - - HYPH 10_1101-2020_12_31_424971 207 45 pairing pair VBG 10_1101-2020_12_31_424971 207 46 522 522 CD 10_1101-2020_12_31_424971 207 47 mechanisms mechanism NNS 10_1101-2020_12_31_424971 207 48 . . . 10_1101-2020_12_31_424971 208 1 523 523 CD 10_1101-2020_12_31_424971 208 2 The the DT 10_1101-2020_12_31_424971 208 3 key key JJ 10_1101-2020_12_31_424971 208 4 insight insight NN 10_1101-2020_12_31_424971 208 5 from from IN 10_1101-2020_12_31_424971 208 6 this this DT 10_1101-2020_12_31_424971 208 7 work work NN 10_1101-2020_12_31_424971 208 8 suggests suggest VBZ 10_1101-2020_12_31_424971 208 9 an an DT 10_1101-2020_12_31_424971 208 10 entirely entirely RB 10_1101-2020_12_31_424971 208 11 novel novel JJ 10_1101-2020_12_31_424971 208 12 pathway pathway NN 10_1101-2020_12_31_424971 208 13 to to TO 10_1101-2020_12_31_424971 208 14 increase increase VB 10_1101-2020_12_31_424971 208 15 the the DT 10_1101-2020_12_31_424971 208 16 efficiency efficiency NN 10_1101-2020_12_31_424971 208 17 524 524 CD 10_1101-2020_12_31_424971 208 18 of of IN 10_1101-2020_12_31_424971 208 19 genome genome NN 10_1101-2020_12_31_424971 208 20 recoding recode VBG 10_1101-2020_12_31_424971 208 21 at at IN 10_1101-2020_12_31_424971 208 22 quadruplet quadruplet NN 10_1101-2020_12_31_424971 208 23 codons codon NNS 10_1101-2020_12_31_424971 208 24 . . . 10_1101-2020_12_31_424971 209 1 While while IN 10_1101-2020_12_31_424971 209 2 initial initial JJ 10_1101-2020_12_31_424971 209 3 success success NN 10_1101-2020_12_31_424971 209 4 in in IN 10_1101-2020_12_31_424971 209 5 genome genome JJ 10_1101-2020_12_31_424971 209 6 recoding recoding NN 10_1101-2020_12_31_424971 209 7 has have VBZ 10_1101-2020_12_31_424971 209 8 been be VBN 10_1101-2020_12_31_424971 209 9 525 525 CD 10_1101-2020_12_31_424971 209 10 achieved achieve VBN 10_1101-2020_12_31_424971 209 11 by by IN 10_1101-2020_12_31_424971 209 12 engineering engineer VBG 10_1101-2020_12_31_424971 209 13 the the DT 10_1101-2020_12_31_424971 209 14 anticodon anticodon NN 10_1101-2020_12_31_424971 209 15 - - HYPH 10_1101-2020_12_31_424971 209 16 codon codon NN 10_1101-2020_12_31_424971 209 17 interactions interaction NNS 10_1101-2020_12_31_424971 209 18 of of IN 10_1101-2020_12_31_424971 209 19 a a DT 10_1101-2020_12_31_424971 209 20 +1-frameshifting +1-frameshifting NN 10_1101-2020_12_31_424971 209 21 - - HYPH 10_1101-2020_12_31_424971 209 22 inducing induce VBG 10_1101-2020_12_31_424971 209 23 tRNA trna NN 10_1101-2020_12_31_424971 209 24 at at IN 10_1101-2020_12_31_424971 209 25 526 526 CD 10_1101-2020_12_31_424971 209 26 the the DT 10_1101-2020_12_31_424971 209 27 A A NNP 10_1101-2020_12_31_424971 209 28 site6,53 site6,53 NNP 10_1101-2020_12_31_424971 209 29 , , , 10_1101-2020_12_31_424971 209 30 or or CC 10_1101-2020_12_31_424971 209 31 by by IN 10_1101-2020_12_31_424971 209 32 engineering engineer VBG 10_1101-2020_12_31_424971 209 33 a a DT 10_1101-2020_12_31_424971 209 34 new new JJ 10_1101-2020_12_31_424971 209 35 bacterial bacterial JJ 10_1101-2020_12_31_424971 209 36 genome genome NN 10_1101-2020_12_31_424971 209 37 with with IN 10_1101-2020_12_31_424971 209 38 a a DT 10_1101-2020_12_31_424971 209 39 minimal minimal JJ 10_1101-2020_12_31_424971 209 40 set set NN 10_1101-2020_12_31_424971 209 41 of of IN 10_1101-2020_12_31_424971 209 42 codons codon NNS 10_1101-2020_12_31_424971 209 43 for for IN 10_1101-2020_12_31_424971 209 44 all all DT 10_1101-2020_12_31_424971 209 45 amino amino NN 10_1101-2020_12_31_424971 209 46 527 527 CD 10_1101-2020_12_31_424971 209 47 acids54 acids54 NNPS 10_1101-2020_12_31_424971 209 48 , , , 10_1101-2020_12_31_424971 209 49 we -PRON- PRP 10_1101-2020_12_31_424971 209 50 suggest suggest VBP 10_1101-2020_12_31_424971 209 51 that that IN 10_1101-2020_12_31_424971 209 52 efforts effort NNS 10_1101-2020_12_31_424971 209 53 to to TO 10_1101-2020_12_31_424971 209 54 engineer engineer VB 10_1101-2020_12_31_424971 209 55 the the DT 10_1101-2020_12_31_424971 209 56 ‘ ' `` 10_1101-2020_12_31_424971 209 57 neck neck NN 10_1101-2020_12_31_424971 209 58 ’ ' '' 10_1101-2020_12_31_424971 209 59 structural structural JJ 10_1101-2020_12_31_424971 209 60 element element NN 10_1101-2020_12_31_424971 209 61 of of IN 10_1101-2020_12_31_424971 209 62 the the DT 10_1101-2020_12_31_424971 209 63 30S 30s CD 10_1101-2020_12_31_424971 209 64 subunit subunit NN 10_1101-2020_12_31_424971 209 65 that that IN 10_1101-2020_12_31_424971 209 66 528 528 CD 10_1101-2020_12_31_424971 209 67 regulates regulate NNS 10_1101-2020_12_31_424971 209 68 head head NN 10_1101-2020_12_31_424971 209 69 swiveling swiveling NN 10_1101-2020_12_31_424971 209 70 would would MD 10_1101-2020_12_31_424971 209 71 be be VB 10_1101-2020_12_31_424971 209 72 as as RB 10_1101-2020_12_31_424971 209 73 , , , 10_1101-2020_12_31_424971 209 74 or or CC 10_1101-2020_12_31_424971 209 75 even even RB 10_1101-2020_12_31_424971 209 76 more more JJR 10_1101-2020_12_31_424971 209 77 , , , 10_1101-2020_12_31_424971 209 78 effective effective JJ 10_1101-2020_12_31_424971 209 79 . . . 10_1101-2020_12_31_424971 210 1 This this DT 10_1101-2020_12_31_424971 210 2 can can MD 10_1101-2020_12_31_424971 210 3 be be VB 10_1101-2020_12_31_424971 210 4 achieved achieve VBN 10_1101-2020_12_31_424971 210 5 by by IN 10_1101-2020_12_31_424971 210 6 screening screen VBG 10_1101-2020_12_31_424971 210 7 529 529 CD 10_1101-2020_12_31_424971 210 8 for for IN 10_1101-2020_12_31_424971 210 9 30S 30s CD 10_1101-2020_12_31_424971 210 10 subunit subunit NN 10_1101-2020_12_31_424971 210 11 variants variant NNS 10_1101-2020_12_31_424971 210 12 that that WDT 10_1101-2020_12_31_424971 210 13 exhibit exhibit VBP 10_1101-2020_12_31_424971 210 14 high high JJ 10_1101-2020_12_31_424971 210 15 +1-frameshifting +1-frameshifting JJ 10_1101-2020_12_31_424971 210 16 efficiencies efficiency NNS 10_1101-2020_12_31_424971 210 17 mediated mediate VBN 10_1101-2020_12_31_424971 210 18 by by IN 10_1101-2020_12_31_424971 210 19 +1 +1 NNPS 10_1101-2020_12_31_424971 210 20 - - : 10_1101-2020_12_31_424971 210 21 530 530 CD 10_1101-2020_12_31_424971 210 22 frameshifting frameshifting NN 10_1101-2020_12_31_424971 210 23 tRNAs trnas CD 10_1101-2020_12_31_424971 210 24 at at IN 10_1101-2020_12_31_424971 210 25 quadruplet quadruplet NN 10_1101-2020_12_31_424971 210 26 codons codon NNS 10_1101-2020_12_31_424971 210 27 while while IN 10_1101-2020_12_31_424971 210 28 preserving preserve VBG 10_1101-2020_12_31_424971 210 29 0-frame 0-frame CD 10_1101-2020_12_31_424971 210 30 translation translation NN 10_1101-2020_12_31_424971 210 31 by by IN 10_1101-2020_12_31_424971 210 32 canonical canonical JJ 10_1101-2020_12_31_424971 210 33 531 531 CD 10_1101-2020_12_31_424971 210 34 tRNAs trnas CD 10_1101-2020_12_31_424971 210 35 at at IN 10_1101-2020_12_31_424971 210 36 triplet triplet NN 10_1101-2020_12_31_424971 210 37 codons codon NNS 10_1101-2020_12_31_424971 210 38 . . . 10_1101-2020_12_31_424971 211 1 Specifically specifically RB 10_1101-2020_12_31_424971 211 2 , , , 10_1101-2020_12_31_424971 211 3 head head NN 10_1101-2020_12_31_424971 211 4 swiveling swiveling NN 10_1101-2020_12_31_424971 211 5 is be VBZ 10_1101-2020_12_31_424971 211 6 driven drive VBN 10_1101-2020_12_31_424971 211 7 by by IN 10_1101-2020_12_31_424971 211 8 the the DT 10_1101-2020_12_31_424971 211 9 synergistic synergistic JJ 10_1101-2020_12_31_424971 211 10 action action NN 10_1101-2020_12_31_424971 211 11 of of IN 10_1101-2020_12_31_424971 211 12 two two CD 10_1101-2020_12_31_424971 211 13 532 532 CD 10_1101-2020_12_31_424971 211 14 hinges hinge NNS 10_1101-2020_12_31_424971 211 15 within within IN 10_1101-2020_12_31_424971 211 16 the the DT 10_1101-2020_12_31_424971 211 17 16S 16s CD 10_1101-2020_12_31_424971 211 18 ribosomal ribosomal NN 10_1101-2020_12_31_424971 211 19 RNA RNA NNP 10_1101-2020_12_31_424971 211 20 elements element NNS 10_1101-2020_12_31_424971 211 21 that that WDT 10_1101-2020_12_31_424971 211 22 comprise comprise VBP 10_1101-2020_12_31_424971 211 23 the the DT 10_1101-2020_12_31_424971 211 24 30S 30s CD 10_1101-2020_12_31_424971 211 25 subunit subunit NN 10_1101-2020_12_31_424971 211 26 neck55 neck55 NN 10_1101-2020_12_31_424971 211 27 . . . 10_1101-2020_12_31_424971 212 1 Hinge hinge NN 10_1101-2020_12_31_424971 212 2 1 1 CD 10_1101-2020_12_31_424971 212 3 is be VBZ 10_1101-2020_12_31_424971 212 4 533 533 CD 10_1101-2020_12_31_424971 212 5 .CC .CC : 10_1101-2020_12_31_424971 212 6 - - HYPH 10_1101-2020_12_31_424971 212 7 BY by IN 10_1101-2020_12_31_424971 212 8 - - HYPH 10_1101-2020_12_31_424971 212 9 NC NC NNP 10_1101-2020_12_31_424971 212 10 - - HYPH 10_1101-2020_12_31_424971 212 11 ND ND NNP 10_1101-2020_12_31_424971 212 12 4.0 4.0 CD 10_1101-2020_12_31_424971 212 13 International International NNP 10_1101-2020_12_31_424971 212 14 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 212 15 under under IN 10_1101-2020_12_31_424971 212 16 a a DT 10_1101-2020_12_31_424971 212 17 ( ( -LRB- 10_1101-2020_12_31_424971 212 18 which which WDT 10_1101-2020_12_31_424971 212 19 was be VBD 10_1101-2020_12_31_424971 212 20 not not RB 10_1101-2020_12_31_424971 212 21 certified certify VBN 10_1101-2020_12_31_424971 212 22 by by IN 10_1101-2020_12_31_424971 212 23 peer peer NN 10_1101-2020_12_31_424971 212 24 review review NN 10_1101-2020_12_31_424971 212 25 ) ) -RRB- 10_1101-2020_12_31_424971 212 26 is be VBZ 10_1101-2020_12_31_424971 212 27 the the DT 10_1101-2020_12_31_424971 212 28 author author NN 10_1101-2020_12_31_424971 212 29 / / SYM 10_1101-2020_12_31_424971 212 30 funder funder NN 10_1101-2020_12_31_424971 212 31 , , , 10_1101-2020_12_31_424971 212 32 who who WP 10_1101-2020_12_31_424971 212 33 has have VBZ 10_1101-2020_12_31_424971 212 34 granted grant VBN 10_1101-2020_12_31_424971 212 35 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 212 36 a a DT 10_1101-2020_12_31_424971 212 37 license license NN 10_1101-2020_12_31_424971 212 38 to to TO 10_1101-2020_12_31_424971 212 39 display display VB 10_1101-2020_12_31_424971 212 40 the the DT 10_1101-2020_12_31_424971 212 41 preprint preprint NN 10_1101-2020_12_31_424971 212 42 in in IN 10_1101-2020_12_31_424971 212 43 perpetuity perpetuity NN 10_1101-2020_12_31_424971 212 44 . . . 10_1101-2020_12_31_424971 213 1 It -PRON- PRP 10_1101-2020_12_31_424971 213 2 is be VBZ 10_1101-2020_12_31_424971 213 3 made make VBN 10_1101-2020_12_31_424971 213 4 The the DT 10_1101-2020_12_31_424971 213 5 copyright copyright NN 10_1101-2020_12_31_424971 213 6 holder holder NN 10_1101-2020_12_31_424971 213 7 for for IN 10_1101-2020_12_31_424971 213 8 this this DT 10_1101-2020_12_31_424971 213 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 213 10 version version NN 10_1101-2020_12_31_424971 213 11 posted post VBD 10_1101-2020_12_31_424971 213 12 January January NNP 10_1101-2020_12_31_424971 213 13 2 2 CD 10_1101-2020_12_31_424971 213 14 , , , 10_1101-2020_12_31_424971 213 15 2021 2021 CD 10_1101-2020_12_31_424971 213 16 . . . 10_1101-2020_12_31_424971 213 17 ; ; : 10_1101-2020_12_31_424971 213 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 213 19 : : : 10_1101-2020_12_31_424971 213 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 213 21 preprint preprint NN 10_1101-2020_12_31_424971 213 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 213 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 213 24 22 22 CD 10_1101-2020_12_31_424971 213 25 composed compose VBN 10_1101-2020_12_31_424971 213 26 of of IN 10_1101-2020_12_31_424971 213 27 two two CD 10_1101-2020_12_31_424971 213 28 G G NNP 10_1101-2020_12_31_424971 213 29 - - HYPH 10_1101-2020_12_31_424971 213 30 U u NN 10_1101-2020_12_31_424971 213 31 wobble wobble JJ 10_1101-2020_12_31_424971 213 32 base base NN 10_1101-2020_12_31_424971 213 33 pairs pair NNS 10_1101-2020_12_31_424971 213 34 that that WDT 10_1101-2020_12_31_424971 213 35 are be VBP 10_1101-2020_12_31_424971 213 36 separated separate VBN 10_1101-2020_12_31_424971 213 37 by by IN 10_1101-2020_12_31_424971 213 38 a a DT 10_1101-2020_12_31_424971 213 39 bulged bulge VBN 10_1101-2020_12_31_424971 213 40 G g NN 10_1101-2020_12_31_424971 213 41 within within IN 10_1101-2020_12_31_424971 213 42 helix helix NN 10_1101-2020_12_31_424971 213 43 28 28 CD 10_1101-2020_12_31_424971 213 44 ( ( -LRB- 10_1101-2020_12_31_424971 213 45 h28 h28 NN 10_1101-2020_12_31_424971 213 46 ) ) -RRB- 10_1101-2020_12_31_424971 213 47 , , , 10_1101-2020_12_31_424971 213 48 534 534 CD 10_1101-2020_12_31_424971 213 49 while while IN 10_1101-2020_12_31_424971 213 50 hinge hinge NN 10_1101-2020_12_31_424971 213 51 2 2 CD 10_1101-2020_12_31_424971 213 52 is be VBZ 10_1101-2020_12_31_424971 213 53 composed compose VBN 10_1101-2020_12_31_424971 213 54 of of IN 10_1101-2020_12_31_424971 213 55 a a DT 10_1101-2020_12_31_424971 213 56 GACU GACU NNP 10_1101-2020_12_31_424971 213 57 linker linker NN 10_1101-2020_12_31_424971 213 58 between between IN 10_1101-2020_12_31_424971 213 59 h34 h34 NN 10_1101-2020_12_31_424971 213 60 and and CC 10_1101-2020_12_31_424971 213 61 h35/36 h35/36 ADD 10_1101-2020_12_31_424971 213 62 within within IN 10_1101-2020_12_31_424971 213 63 a a DT 10_1101-2020_12_31_424971 213 64 three three CD 10_1101-2020_12_31_424971 213 65 - - HYPH 10_1101-2020_12_31_424971 213 66 helical helical JJ 10_1101-2020_12_31_424971 213 67 535 535 CD 10_1101-2020_12_31_424971 213 68 junction junction NN 10_1101-2020_12_31_424971 213 69 with with IN 10_1101-2020_12_31_424971 213 70 h38 h38 NN 10_1101-2020_12_31_424971 213 71 . . . 10_1101-2020_12_31_424971 214 1 Co Co NNS 10_1101-2020_12_31_424971 214 2 - - VB 10_1101-2020_12_31_424971 214 3 engineering engineer VBG 10_1101-2020_12_31_424971 214 4 these these DT 10_1101-2020_12_31_424971 214 5 two two CD 10_1101-2020_12_31_424971 214 6 hinges hinge NNS 10_1101-2020_12_31_424971 214 7 by by IN 10_1101-2020_12_31_424971 214 8 directed direct VBN 10_1101-2020_12_31_424971 214 9 evolution evolution NN 10_1101-2020_12_31_424971 214 10 should should MD 10_1101-2020_12_31_424971 214 11 identify identify VB 10_1101-2020_12_31_424971 214 12 such such JJ 10_1101-2020_12_31_424971 214 13 30S 30S NNP 10_1101-2020_12_31_424971 214 14 536 536 CD 10_1101-2020_12_31_424971 214 15 subunit subunit NN 10_1101-2020_12_31_424971 214 16 variants variant NNS 10_1101-2020_12_31_424971 214 17 . . . 10_1101-2020_12_31_424971 215 1 To to TO 10_1101-2020_12_31_424971 215 2 complement complement VB 10_1101-2020_12_31_424971 215 3 the the DT 10_1101-2020_12_31_424971 215 4 directed direct VBN 10_1101-2020_12_31_424971 215 5 evolution evolution NN 10_1101-2020_12_31_424971 215 6 approach approach NN 10_1101-2020_12_31_424971 215 7 , , , 10_1101-2020_12_31_424971 215 8 we -PRON- PRP 10_1101-2020_12_31_424971 215 9 suggest suggest VBP 10_1101-2020_12_31_424971 215 10 that that IN 10_1101-2020_12_31_424971 215 11 our -PRON- PRP$ 10_1101-2020_12_31_424971 215 12 recently recently RB 10_1101-2020_12_31_424971 215 13 537 537 CD 10_1101-2020_12_31_424971 215 14 developed develop VBN 10_1101-2020_12_31_424971 215 15 time time NN 10_1101-2020_12_31_424971 215 16 - - HYPH 10_1101-2020_12_31_424971 215 17 resolved resolve VBN 10_1101-2020_12_31_424971 215 18 cryogenic cryogenic JJ 10_1101-2020_12_31_424971 215 19 electron electron NN 10_1101-2020_12_31_424971 215 20 microscopy microscopy NN 10_1101-2020_12_31_424971 215 21 ( ( -LRB- 10_1101-2020_12_31_424971 215 22 TR TR NNP 10_1101-2020_12_31_424971 215 23 cryo cryo JJ 10_1101-2020_12_31_424971 215 24 - - HYPH 10_1101-2020_12_31_424971 215 25 EM em NN 10_1101-2020_12_31_424971 215 26 ) ) -RRB- 10_1101-2020_12_31_424971 215 27 method56,57 method56,57 NN 10_1101-2020_12_31_424971 215 28 can can MD 10_1101-2020_12_31_424971 215 29 be be VB 10_1101-2020_12_31_424971 215 30 used use VBN 10_1101-2020_12_31_424971 215 31 538 538 CD 10_1101-2020_12_31_424971 215 32 to to TO 10_1101-2020_12_31_424971 215 33 obtain obtain VB 10_1101-2020_12_31_424971 215 34 structures structure NNS 10_1101-2020_12_31_424971 215 35 of of IN 10_1101-2020_12_31_424971 215 36 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 215 37 and and CC 10_1101-2020_12_31_424971 215 38 ProL ProL NNP 10_1101-2020_12_31_424971 215 39 in in IN 10_1101-2020_12_31_424971 215 40 EF EF NNP 10_1101-2020_12_31_424971 215 41 - - HYPH 10_1101-2020_12_31_424971 215 42 G G NNP 10_1101-2020_12_31_424971 215 43 - - HYPH 10_1101-2020_12_31_424971 215 44 bound bind VBN 10_1101-2020_12_31_424971 215 45 PRE PRE NNP 10_1101-2020_12_31_424971 215 46 complexes complex NNS 10_1101-2020_12_31_424971 215 47 captured capture VBN 10_1101-2020_12_31_424971 215 48 in in IN 10_1101-2020_12_31_424971 215 49 intermediate intermediate JJ 10_1101-2020_12_31_424971 215 50 539 539 CD 10_1101-2020_12_31_424971 215 51 states state NNS 10_1101-2020_12_31_424971 215 52 of of IN 10_1101-2020_12_31_424971 215 53 translocation translocation NN 10_1101-2020_12_31_424971 215 54 . . . 10_1101-2020_12_31_424971 216 1 Such such JJ 10_1101-2020_12_31_424971 216 2 cryo cryo JJ 10_1101-2020_12_31_424971 216 3 - - HYPH 10_1101-2020_12_31_424971 216 4 EM em NN 10_1101-2020_12_31_424971 216 5 structures structure NNS 10_1101-2020_12_31_424971 216 6 would would MD 10_1101-2020_12_31_424971 216 7 help help VB 10_1101-2020_12_31_424971 216 8 further far RBR 10_1101-2020_12_31_424971 216 9 define define VB 10_1101-2020_12_31_424971 216 10 how how WRB 10_1101-2020_12_31_424971 216 11 the the DT 10_1101-2020_12_31_424971 216 12 two two CD 10_1101-2020_12_31_424971 216 13 hinges hinge NNS 10_1101-2020_12_31_424971 216 14 540 540 CD 10_1101-2020_12_31_424971 216 15 that that DT 10_1101-2020_12_31_424971 216 16 control control NN 10_1101-2020_12_31_424971 216 17 head head NN 10_1101-2020_12_31_424971 216 18 swiveling swivel VBG 10_1101-2020_12_31_424971 216 19 are be VBP 10_1101-2020_12_31_424971 216 20 differentially differentially RB 10_1101-2020_12_31_424971 216 21 modulated modulate VBN 10_1101-2020_12_31_424971 216 22 during during IN 10_1101-2020_12_31_424971 216 23 translocation translocation NN 10_1101-2020_12_31_424971 216 24 of of IN 10_1101-2020_12_31_424971 216 25 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 216 26 vs. vs. IN 10_1101-2020_12_31_424971 216 27 ProL ProL NNP 10_1101-2020_12_31_424971 216 28 to to IN 10_1101-2020_12_31_424971 216 29 541 541 CD 10_1101-2020_12_31_424971 216 30 provide provide VB 10_1101-2020_12_31_424971 216 31 a a DT 10_1101-2020_12_31_424971 216 32 structure structure NN 10_1101-2020_12_31_424971 216 33 - - HYPH 10_1101-2020_12_31_424971 216 34 based base VBN 10_1101-2020_12_31_424971 216 35 roadmap roadmap NNP 10_1101-2020_12_31_424971 216 36 for for IN 10_1101-2020_12_31_424971 216 37 engineering engineer VBG 10_1101-2020_12_31_424971 216 38 them -PRON- PRP 10_1101-2020_12_31_424971 216 39 . . . 10_1101-2020_12_31_424971 217 1 In in IN 10_1101-2020_12_31_424971 217 2 addition addition NN 10_1101-2020_12_31_424971 217 3 , , , 10_1101-2020_12_31_424971 217 4 detailed detailed JJ 10_1101-2020_12_31_424971 217 5 comparison comparison NN 10_1101-2020_12_31_424971 217 6 of of IN 10_1101-2020_12_31_424971 217 7 such such JJ 10_1101-2020_12_31_424971 217 8 542 542 CD 10_1101-2020_12_31_424971 217 9 structures structure NNS 10_1101-2020_12_31_424971 217 10 would would MD 10_1101-2020_12_31_424971 217 11 offer offer VB 10_1101-2020_12_31_424971 217 12 the the DT 10_1101-2020_12_31_424971 217 13 opportunity opportunity NN 10_1101-2020_12_31_424971 217 14 to to TO 10_1101-2020_12_31_424971 217 15 identify identify VB 10_1101-2020_12_31_424971 217 16 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 217 17 structural structural JJ 10_1101-2020_12_31_424971 217 18 elements element NNS 10_1101-2020_12_31_424971 217 19 beyond beyond IN 10_1101-2020_12_31_424971 217 20 the the DT 10_1101-2020_12_31_424971 217 21 two two CD 10_1101-2020_12_31_424971 217 22 543 543 CD 10_1101-2020_12_31_424971 217 23 hinges hinge NNS 10_1101-2020_12_31_424971 217 24 that that WDT 10_1101-2020_12_31_424971 217 25 play play VBP 10_1101-2020_12_31_424971 217 26 a a DT 10_1101-2020_12_31_424971 217 27 role role NN 10_1101-2020_12_31_424971 217 28 in in IN 10_1101-2020_12_31_424971 217 29 +1 +1 NNS 10_1101-2020_12_31_424971 217 30 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 217 31 and and CC 10_1101-2020_12_31_424971 217 32 can can MD 10_1101-2020_12_31_424971 217 33 thus thus RB 10_1101-2020_12_31_424971 217 34 serve serve VB 10_1101-2020_12_31_424971 217 35 as as IN 10_1101-2020_12_31_424971 217 36 additional additional JJ 10_1101-2020_12_31_424971 217 37 targets target NNS 10_1101-2020_12_31_424971 217 38 for for IN 10_1101-2020_12_31_424971 217 39 engineering engineering NN 10_1101-2020_12_31_424971 217 40 . . . 10_1101-2020_12_31_424971 218 1 544 544 CD 10_1101-2020_12_31_424971 218 2 Furthermore furthermore RB 10_1101-2020_12_31_424971 218 3 , , , 10_1101-2020_12_31_424971 218 4 antibiotics antibiotic NNS 10_1101-2020_12_31_424971 218 5 that that WDT 10_1101-2020_12_31_424971 218 6 bind bind VBP 10_1101-2020_12_31_424971 218 7 to to IN 10_1101-2020_12_31_424971 218 8 the the DT 10_1101-2020_12_31_424971 218 9 30S 30s CD 10_1101-2020_12_31_424971 218 10 subunit subunit NN 10_1101-2020_12_31_424971 218 11 and and CC 10_1101-2020_12_31_424971 218 12 act act VB 10_1101-2020_12_31_424971 218 13 as as IN 10_1101-2020_12_31_424971 218 14 translocation translocation NN 10_1101-2020_12_31_424971 218 15 modulators modulator NNS 10_1101-2020_12_31_424971 218 16 can can MD 10_1101-2020_12_31_424971 218 17 be be VB 10_1101-2020_12_31_424971 218 18 545 545 CD 10_1101-2020_12_31_424971 218 19 exploited exploit VBN 10_1101-2020_12_31_424971 218 20 to to TO 10_1101-2020_12_31_424971 218 21 further further RB 10_1101-2020_12_31_424971 218 22 increase increase VB 10_1101-2020_12_31_424971 218 23 the the DT 10_1101-2020_12_31_424971 218 24 +1-frameshifting +1-frameshifting JJ 10_1101-2020_12_31_424971 218 25 efficiency efficiency NN 10_1101-2020_12_31_424971 218 26 at at IN 10_1101-2020_12_31_424971 218 27 a a DT 10_1101-2020_12_31_424971 218 28 quadruplet quadruplet NN 10_1101-2020_12_31_424971 218 29 codon codon NN 10_1101-2020_12_31_424971 218 30 with with IN 10_1101-2020_12_31_424971 218 31 either either DT 10_1101-2020_12_31_424971 218 32 546 546 CD 10_1101-2020_12_31_424971 218 33 wildtype wildtype NN 10_1101-2020_12_31_424971 218 34 or or CC 10_1101-2020_12_31_424971 218 35 highly highly RB 10_1101-2020_12_31_424971 218 36 efficient efficient JJ 10_1101-2020_12_31_424971 218 37 30S 30s CD 10_1101-2020_12_31_424971 218 38 subunit subunit NN 10_1101-2020_12_31_424971 218 39 variants variant NNS 10_1101-2020_12_31_424971 218 40 . . . 10_1101-2020_12_31_424971 219 1 Implemented implement VBN 10_1101-2020_12_31_424971 219 2 in in IN 10_1101-2020_12_31_424971 219 3 combination combination NN 10_1101-2020_12_31_424971 219 4 and and CC 10_1101-2020_12_31_424971 219 5 integrated integrate VBN 10_1101-2020_12_31_424971 219 6 into into IN 10_1101-2020_12_31_424971 219 7 547 547 CD 10_1101-2020_12_31_424971 219 8 a a DT 10_1101-2020_12_31_424971 219 9 recently recently RB 10_1101-2020_12_31_424971 219 10 described describe VBN 10_1101-2020_12_31_424971 219 11 in in IN 10_1101-2020_12_31_424971 219 12 vivo vivo NNP 10_1101-2020_12_31_424971 219 13 ‘ ' `` 10_1101-2020_12_31_424971 219 14 designer designer NN 10_1101-2020_12_31_424971 219 15 organelle organelle NNP 10_1101-2020_12_31_424971 219 16 ’ ' '' 10_1101-2020_12_31_424971 219 17 strategy58 strategy58 NNP 10_1101-2020_12_31_424971 219 18 , , , 10_1101-2020_12_31_424971 219 19 these these DT 10_1101-2020_12_31_424971 219 20 approaches approach NNS 10_1101-2020_12_31_424971 219 21 should should MD 10_1101-2020_12_31_424971 219 22 provide provide VB 10_1101-2020_12_31_424971 219 23 a a DT 10_1101-2020_12_31_424971 219 24 548 548 CD 10_1101-2020_12_31_424971 219 25 novel novel NN 10_1101-2020_12_31_424971 219 26 and and CC 10_1101-2020_12_31_424971 219 27 powerful powerful JJ 10_1101-2020_12_31_424971 219 28 platform platform NN 10_1101-2020_12_31_424971 219 29 for for IN 10_1101-2020_12_31_424971 219 30 increasing increase VBG 10_1101-2020_12_31_424971 219 31 the the DT 10_1101-2020_12_31_424971 219 32 efficiency efficiency NN 10_1101-2020_12_31_424971 219 33 of of IN 10_1101-2020_12_31_424971 219 34 genome genome JJ 10_1101-2020_12_31_424971 219 35 recoding recode VBG 10_1101-2020_12_31_424971 219 36 at at IN 10_1101-2020_12_31_424971 219 37 quadruplet quadruplet NN 10_1101-2020_12_31_424971 219 38 codons codon NNS 10_1101-2020_12_31_424971 219 39 549 549 CD 10_1101-2020_12_31_424971 219 40 with with IN 10_1101-2020_12_31_424971 219 41 minimal minimal JJ 10_1101-2020_12_31_424971 219 42 off off JJ 10_1101-2020_12_31_424971 219 43 - - HYPH 10_1101-2020_12_31_424971 219 44 target target NN 10_1101-2020_12_31_424971 219 45 effects effect NNS 10_1101-2020_12_31_424971 219 46 . . . 10_1101-2020_12_31_424971 220 1 550 550 CD 10_1101-2020_12_31_424971 220 2 .CC .CC : 10_1101-2020_12_31_424971 220 3 - - HYPH 10_1101-2020_12_31_424971 220 4 BY by IN 10_1101-2020_12_31_424971 220 5 - - HYPH 10_1101-2020_12_31_424971 220 6 NC NC NNP 10_1101-2020_12_31_424971 220 7 - - HYPH 10_1101-2020_12_31_424971 220 8 ND ND NNP 10_1101-2020_12_31_424971 220 9 4.0 4.0 CD 10_1101-2020_12_31_424971 220 10 International International NNP 10_1101-2020_12_31_424971 220 11 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 220 12 under under IN 10_1101-2020_12_31_424971 220 13 a a DT 10_1101-2020_12_31_424971 220 14 ( ( -LRB- 10_1101-2020_12_31_424971 220 15 which which WDT 10_1101-2020_12_31_424971 220 16 was be VBD 10_1101-2020_12_31_424971 220 17 not not RB 10_1101-2020_12_31_424971 220 18 certified certify VBN 10_1101-2020_12_31_424971 220 19 by by IN 10_1101-2020_12_31_424971 220 20 peer peer NN 10_1101-2020_12_31_424971 220 21 review review NN 10_1101-2020_12_31_424971 220 22 ) ) -RRB- 10_1101-2020_12_31_424971 220 23 is be VBZ 10_1101-2020_12_31_424971 220 24 the the DT 10_1101-2020_12_31_424971 220 25 author author NN 10_1101-2020_12_31_424971 220 26 / / SYM 10_1101-2020_12_31_424971 220 27 funder funder NN 10_1101-2020_12_31_424971 220 28 , , , 10_1101-2020_12_31_424971 220 29 who who WP 10_1101-2020_12_31_424971 220 30 has have VBZ 10_1101-2020_12_31_424971 220 31 granted grant VBN 10_1101-2020_12_31_424971 220 32 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 220 33 a a DT 10_1101-2020_12_31_424971 220 34 license license NN 10_1101-2020_12_31_424971 220 35 to to TO 10_1101-2020_12_31_424971 220 36 display display VB 10_1101-2020_12_31_424971 220 37 the the DT 10_1101-2020_12_31_424971 220 38 preprint preprint NN 10_1101-2020_12_31_424971 220 39 in in IN 10_1101-2020_12_31_424971 220 40 perpetuity perpetuity NN 10_1101-2020_12_31_424971 220 41 . . . 10_1101-2020_12_31_424971 221 1 It -PRON- PRP 10_1101-2020_12_31_424971 221 2 is be VBZ 10_1101-2020_12_31_424971 221 3 made make VBN 10_1101-2020_12_31_424971 221 4 The the DT 10_1101-2020_12_31_424971 221 5 copyright copyright NN 10_1101-2020_12_31_424971 221 6 holder holder NN 10_1101-2020_12_31_424971 221 7 for for IN 10_1101-2020_12_31_424971 221 8 this this DT 10_1101-2020_12_31_424971 221 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 221 10 version version NN 10_1101-2020_12_31_424971 221 11 posted post VBD 10_1101-2020_12_31_424971 221 12 January January NNP 10_1101-2020_12_31_424971 221 13 2 2 CD 10_1101-2020_12_31_424971 221 14 , , , 10_1101-2020_12_31_424971 221 15 2021 2021 CD 10_1101-2020_12_31_424971 221 16 . . . 10_1101-2020_12_31_424971 221 17 ; ; : 10_1101-2020_12_31_424971 221 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 221 19 : : : 10_1101-2020_12_31_424971 221 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 221 21 preprint preprint NN 10_1101-2020_12_31_424971 221 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 221 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 221 24 23 23 CD 10_1101-2020_12_31_424971 221 25 METHODS METHODS NNP 10_1101-2020_12_31_424971 221 26 551 551 CD 10_1101-2020_12_31_424971 221 27 Construction construction NN 10_1101-2020_12_31_424971 221 28 of of IN 10_1101-2020_12_31_424971 221 29 E. E. NNP 10_1101-2020_12_31_424971 221 30 coli coli NNS 10_1101-2020_12_31_424971 221 31 strains strain NNS 10_1101-2020_12_31_424971 221 32 . . . 10_1101-2020_12_31_424971 222 1 E. E. NNP 10_1101-2020_12_31_424971 222 2 coli coli NNS 10_1101-2020_12_31_424971 222 3 strains strain NNS 10_1101-2020_12_31_424971 222 4 that that WDT 10_1101-2020_12_31_424971 222 5 expressed express VBD 10_1101-2020_12_31_424971 222 6 a a DT 10_1101-2020_12_31_424971 222 7 plasmid plasmid NN 10_1101-2020_12_31_424971 222 8 - - HYPH 10_1101-2020_12_31_424971 222 9 borne bear VBN 10_1101-2020_12_31_424971 222 10 ProL ProL NNP 10_1101-2020_12_31_424971 222 11 or or CC 10_1101-2020_12_31_424971 222 12 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 222 13 for for IN 10_1101-2020_12_31_424971 222 14 552 552 CD 10_1101-2020_12_31_424971 222 15 isolation isolation NN 10_1101-2020_12_31_424971 222 16 of of IN 10_1101-2020_12_31_424971 222 17 native native JJ 10_1101-2020_12_31_424971 222 18 - - HYPH 10_1101-2020_12_31_424971 222 19 state state NN 10_1101-2020_12_31_424971 222 20 tRNAs trna NNS 10_1101-2020_12_31_424971 222 21 were be VBD 10_1101-2020_12_31_424971 222 22 made make VBN 10_1101-2020_12_31_424971 222 23 in in IN 10_1101-2020_12_31_424971 222 24 a a DT 10_1101-2020_12_31_424971 222 25 ProL ProL NNP 10_1101-2020_12_31_424971 222 26 - - HYPH 10_1101-2020_12_31_424971 222 27 KO KO NNP 10_1101-2020_12_31_424971 222 28 strain strain NN 10_1101-2020_12_31_424971 222 29 , , , 10_1101-2020_12_31_424971 222 30 which which WDT 10_1101-2020_12_31_424971 222 31 was be VBD 10_1101-2020_12_31_424971 222 32 constructed construct VBN 10_1101-2020_12_31_424971 222 33 by by IN 10_1101-2020_12_31_424971 222 34 inserting insert VBG 10_1101-2020_12_31_424971 222 35 553 553 CD 10_1101-2020_12_31_424971 222 36 the the DT 10_1101-2020_12_31_424971 222 37 Kan Kan NNP 10_1101-2020_12_31_424971 222 38 - - HYPH 10_1101-2020_12_31_424971 222 39 resistance resistance NNP 10_1101-2020_12_31_424971 222 40 ( ( -LRB- 10_1101-2020_12_31_424971 222 41 Kan Kan NNP 10_1101-2020_12_31_424971 222 42 - - HYPH 10_1101-2020_12_31_424971 222 43 R R NNP 10_1101-2020_12_31_424971 222 44 ) ) -RRB- 10_1101-2020_12_31_424971 222 45 gene gene NN 10_1101-2020_12_31_424971 222 46 , , , 10_1101-2020_12_31_424971 222 47 amplified amplify VBN 10_1101-2020_12_31_424971 222 48 by by IN 10_1101-2020_12_31_424971 222 49 PCR PCR NNP 10_1101-2020_12_31_424971 222 50 primers primer NNS 10_1101-2020_12_31_424971 222 51 from from IN 10_1101-2020_12_31_424971 222 52 pKD4 pKD4 NNS 10_1101-2020_12_31_424971 222 53 , , , 10_1101-2020_12_31_424971 222 54 into into IN 10_1101-2020_12_31_424971 222 55 the the DT 10_1101-2020_12_31_424971 222 56 ProL ProL NNP 10_1101-2020_12_31_424971 222 57 locus locus NN 10_1101-2020_12_31_424971 222 58 of of IN 10_1101-2020_12_31_424971 222 59 E. E. NNP 10_1101-2020_12_31_424971 222 60 554 554 CD 10_1101-2020_12_31_424971 222 61 coli coli NNS 10_1101-2020_12_31_424971 222 62 BL21(DE3 BL21(DE3 '' 10_1101-2020_12_31_424971 222 63 ) ) -RRB- 10_1101-2020_12_31_424971 222 64 using use VBG 10_1101-2020_12_31_424971 222 65 the the DT 10_1101-2020_12_31_424971 222 66 l l NNP 10_1101-2020_12_31_424971 222 67 - - HYPH 10_1101-2020_12_31_424971 222 68 Red Red NNP 10_1101-2020_12_31_424971 222 69 recombination recombination NN 10_1101-2020_12_31_424971 222 70 method59 method59 NN 10_1101-2020_12_31_424971 222 71 , , , 10_1101-2020_12_31_424971 222 72 followed follow VBN 10_1101-2020_12_31_424971 222 73 by by IN 10_1101-2020_12_31_424971 222 74 removal removal NN 10_1101-2020_12_31_424971 222 75 of of IN 10_1101-2020_12_31_424971 222 76 the the DT 10_1101-2020_12_31_424971 222 77 Kan Kan NNP 10_1101-2020_12_31_424971 222 78 - - HYPH 10_1101-2020_12_31_424971 222 79 R R NNP 10_1101-2020_12_31_424971 222 80 gene gene NN 10_1101-2020_12_31_424971 222 81 555 555 CD 10_1101-2020_12_31_424971 222 82 using use VBG 10_1101-2020_12_31_424971 222 83 FLP FLP NNP 10_1101-2020_12_31_424971 222 84 recombination30 recombination30 NNP 10_1101-2020_12_31_424971 222 85 . . . 10_1101-2020_12_31_424971 223 1 The the DT 10_1101-2020_12_31_424971 223 2 pKK223-SufB2 pKK223-SufB2 NNP 10_1101-2020_12_31_424971 223 3 plasmid plasmid NN 10_1101-2020_12_31_424971 223 4 was be VBD 10_1101-2020_12_31_424971 223 5 made make VBN 10_1101-2020_12_31_424971 223 6 by by IN 10_1101-2020_12_31_424971 223 7 site site NN 10_1101-2020_12_31_424971 223 8 - - HYPH 10_1101-2020_12_31_424971 223 9 directed direct VBN 10_1101-2020_12_31_424971 223 10 mutagenesis mutagenesis NN 10_1101-2020_12_31_424971 223 11 556 556 CD 10_1101-2020_12_31_424971 223 12 to to TO 10_1101-2020_12_31_424971 223 13 introduce introduce VB 10_1101-2020_12_31_424971 223 14 G37a g37a NN 10_1101-2020_12_31_424971 223 15 into into IN 10_1101-2020_12_31_424971 223 16 the the DT 10_1101-2020_12_31_424971 223 17 pKK223-ProL pKK223-ProL NNP 10_1101-2020_12_31_424971 223 18 plasmid29 plasmid29 NN 10_1101-2020_12_31_424971 223 19 . . . 10_1101-2020_12_31_424971 224 1 E. E. NNP 10_1101-2020_12_31_424971 224 2 coli coli NNS 10_1101-2020_12_31_424971 224 3 strains strain VBZ 10_1101-2020_12_31_424971 224 4 that that WDT 10_1101-2020_12_31_424971 224 5 expressed express VBD 10_1101-2020_12_31_424971 224 6 ProL ProL NNP 10_1101-2020_12_31_424971 224 7 or or CC 10_1101-2020_12_31_424971 224 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 224 9 557 557 CD 10_1101-2020_12_31_424971 224 10 from from IN 10_1101-2020_12_31_424971 224 11 the the DT 10_1101-2020_12_31_424971 224 12 chromosome chromosome NN 10_1101-2020_12_31_424971 224 13 as as IN 10_1101-2020_12_31_424971 224 14 an an DT 10_1101-2020_12_31_424971 224 15 isogenic isogenic JJ 10_1101-2020_12_31_424971 224 16 pair pair NN 10_1101-2020_12_31_424971 224 17 for for IN 10_1101-2020_12_31_424971 224 18 reporter reporter NN 10_1101-2020_12_31_424971 224 19 assays assay NNS 10_1101-2020_12_31_424971 224 20 were be VBD 10_1101-2020_12_31_424971 224 21 made make VBN 10_1101-2020_12_31_424971 224 22 using use VBG 10_1101-2020_12_31_424971 224 23 the the DT 10_1101-2020_12_31_424971 224 24 l l NNP 10_1101-2020_12_31_424971 224 25 - - HYPH 10_1101-2020_12_31_424971 224 26 Red Red NNP 10_1101-2020_12_31_424971 224 27 558 558 CD 10_1101-2020_12_31_424971 224 28 technique30 technique30 NNS 10_1101-2020_12_31_424971 224 29 . . . 10_1101-2020_12_31_424971 225 1 To to TO 10_1101-2020_12_31_424971 225 2 construct construct VB 10_1101-2020_12_31_424971 225 3 the the DT 10_1101-2020_12_31_424971 225 4 E. E. NNP 10_1101-2020_12_31_424971 225 5 coli coli VB 10_1101-2020_12_31_424971 225 6 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 225 7 strain strain NN 10_1101-2020_12_31_424971 225 8 , , , 10_1101-2020_12_31_424971 225 9 the the DT 10_1101-2020_12_31_424971 225 10 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 225 11 gene gene NN 10_1101-2020_12_31_424971 225 12 was be VBD 10_1101-2020_12_31_424971 225 13 PCR PCR NNP 10_1101-2020_12_31_424971 225 14 - - HYPH 10_1101-2020_12_31_424971 225 15 amplified amplify VBN 10_1101-2020_12_31_424971 225 16 from from IN 10_1101-2020_12_31_424971 225 17 559 559 CD 10_1101-2020_12_31_424971 225 18 pKK223-SufB2 pKK223-SufB2 NNS 10_1101-2020_12_31_424971 225 19 , , , 10_1101-2020_12_31_424971 225 20 and and CC 10_1101-2020_12_31_424971 225 21 the the DT 10_1101-2020_12_31_424971 225 22 5 5 CD 10_1101-2020_12_31_424971 225 23 ' ' '' 10_1101-2020_12_31_424971 225 24 end end NN 10_1101-2020_12_31_424971 225 25 of of IN 10_1101-2020_12_31_424971 225 26 the the DT 10_1101-2020_12_31_424971 225 27 amplified amplified JJ 10_1101-2020_12_31_424971 225 28 gene gene NN 10_1101-2020_12_31_424971 225 29 was be VBD 10_1101-2020_12_31_424971 225 30 joined join VBN 10_1101-2020_12_31_424971 225 31 with with IN 10_1101-2020_12_31_424971 225 32 Kan Kan NNP 10_1101-2020_12_31_424971 225 33 - - HYPH 10_1101-2020_12_31_424971 225 34 R R NNP 10_1101-2020_12_31_424971 225 35 ( ( -LRB- 10_1101-2020_12_31_424971 225 36 from from IN 10_1101-2020_12_31_424971 225 37 pKD4 pKD4 NNS 10_1101-2020_12_31_424971 225 38 ) ) -RRB- 10_1101-2020_12_31_424971 225 39 by by IN 10_1101-2020_12_31_424971 225 40 PCR PCR NNP 10_1101-2020_12_31_424971 225 41 560 560 CD 10_1101-2020_12_31_424971 225 42 using use VBG 10_1101-2020_12_31_424971 225 43 reverse-2 reverse-2 NNP 10_1101-2020_12_31_424971 225 44 primer primer NN 10_1101-2020_12_31_424971 225 45 , , , 10_1101-2020_12_31_424971 225 46 while while IN 10_1101-2020_12_31_424971 225 47 the the DT 10_1101-2020_12_31_424971 225 48 3 3 CD 10_1101-2020_12_31_424971 225 49 ' ' POS 10_1101-2020_12_31_424971 225 50 end end NN 10_1101-2020_12_31_424971 225 51 was be VBD 10_1101-2020_12_31_424971 225 52 homologous homologous JJ 10_1101-2020_12_31_424971 225 53 to to IN 10_1101-2020_12_31_424971 225 54 the the DT 10_1101-2020_12_31_424971 225 55 ProL prol NN 10_1101-2020_12_31_424971 225 56 3 3 CD 10_1101-2020_12_31_424971 225 57 ' ' POS 10_1101-2020_12_31_424971 225 58 flanking flanking NN 10_1101-2020_12_31_424971 225 59 region region NN 10_1101-2020_12_31_424971 225 60 . . . 10_1101-2020_12_31_424971 226 1 The the DT 10_1101-2020_12_31_424971 226 2 PCR-561 pcr-561 NN 10_1101-2020_12_31_424971 226 3 amplified amplify VBD 10_1101-2020_12_31_424971 226 4 SufB2-Kan sufb2-kan ADD 10_1101-2020_12_31_424971 226 5 product product NN 10_1101-2020_12_31_424971 226 6 was be VBD 10_1101-2020_12_31_424971 226 7 used use VBN 10_1101-2020_12_31_424971 226 8 to to TO 10_1101-2020_12_31_424971 226 9 replace replace VB 10_1101-2020_12_31_424971 226 10 ProL ProL NNP 10_1101-2020_12_31_424971 226 11 in in IN 10_1101-2020_12_31_424971 226 12 l l NNP 10_1101-2020_12_31_424971 226 13 - - HYPH 10_1101-2020_12_31_424971 226 14 Red red NN 10_1101-2020_12_31_424971 226 15 expressing express VBG 10_1101-2020_12_31_424971 226 16 cells cell NNS 10_1101-2020_12_31_424971 226 17 . . . 10_1101-2020_12_31_424971 227 1 An an DT 10_1101-2020_12_31_424971 227 2 isogenic isogenic JJ 10_1101-2020_12_31_424971 227 3 562 562 CD 10_1101-2020_12_31_424971 227 4 counterpart counterpart NN 10_1101-2020_12_31_424971 227 5 strain strain NN 10_1101-2020_12_31_424971 227 6 expressing express VBG 10_1101-2020_12_31_424971 227 7 ProL ProL NNP 10_1101-2020_12_31_424971 227 8 - - HYPH 10_1101-2020_12_31_424971 227 9 Kan Kan NNP 10_1101-2020_12_31_424971 227 10 was be VBD 10_1101-2020_12_31_424971 227 11 also also RB 10_1101-2020_12_31_424971 227 12 made make VBN 10_1101-2020_12_31_424971 227 13 . . . 10_1101-2020_12_31_424971 228 1 These these DT 10_1101-2020_12_31_424971 228 2 ProL ProL NNP 10_1101-2020_12_31_424971 228 3 - - HYPH 10_1101-2020_12_31_424971 228 4 Kan Kan NNP 10_1101-2020_12_31_424971 228 5 and and CC 10_1101-2020_12_31_424971 228 6 SufB2-Kan sufb2-kan ADD 10_1101-2020_12_31_424971 228 7 loci loci NN 10_1101-2020_12_31_424971 228 8 563 563 CD 10_1101-2020_12_31_424971 228 9 were be VBD 10_1101-2020_12_31_424971 228 10 independently independently RB 10_1101-2020_12_31_424971 228 11 transferred transfer VBN 10_1101-2020_12_31_424971 228 12 to to IN 10_1101-2020_12_31_424971 228 13 the the DT 10_1101-2020_12_31_424971 228 14 trmD trmd CD 10_1101-2020_12_31_424971 228 15 - - HYPH 10_1101-2020_12_31_424971 228 16 KO KO NNP 10_1101-2020_12_31_424971 228 17 strain29 strain29 NN 10_1101-2020_12_31_424971 228 18 by by IN 10_1101-2020_12_31_424971 228 19 P1 P1 NNP 10_1101-2020_12_31_424971 228 20 transduction transduction NN 10_1101-2020_12_31_424971 228 21 , , , 10_1101-2020_12_31_424971 228 22 followed follow VBN 10_1101-2020_12_31_424971 228 23 by by IN 10_1101-2020_12_31_424971 228 24 pCP20 pcp20 JJ 10_1101-2020_12_31_424971 228 25 - - : 10_1101-2020_12_31_424971 228 26 564 564 CD 10_1101-2020_12_31_424971 228 27 dependent dependent JJ 10_1101-2020_12_31_424971 228 28 FLP FLP NNP 10_1101-2020_12_31_424971 228 29 recombination recombination NN 10_1101-2020_12_31_424971 228 30 , , , 10_1101-2020_12_31_424971 228 31 generating generate VBG 10_1101-2020_12_31_424971 228 32 the the DT 10_1101-2020_12_31_424971 228 33 isogenic isogenic JJ 10_1101-2020_12_31_424971 228 34 pair pair NN 10_1101-2020_12_31_424971 228 35 of of IN 10_1101-2020_12_31_424971 228 36 ProL ProL NNP 10_1101-2020_12_31_424971 228 37 and and CC 10_1101-2020_12_31_424971 228 38 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 228 39 strains strain NNS 10_1101-2020_12_31_424971 228 40 in in IN 10_1101-2020_12_31_424971 228 41 the the DT 10_1101-2020_12_31_424971 228 42 trmD-565 trmD-565 NNP 10_1101-2020_12_31_424971 228 43 KO KO NNP 10_1101-2020_12_31_424971 228 44 background background NN 10_1101-2020_12_31_424971 228 45 . . . 10_1101-2020_12_31_424971 229 1 These these DT 10_1101-2020_12_31_424971 229 2 strains strain NNS 10_1101-2020_12_31_424971 229 3 were be VBD 10_1101-2020_12_31_424971 229 4 transformed transform VBN 10_1101-2020_12_31_424971 229 5 with with IN 10_1101-2020_12_31_424971 229 6 pKK223 pKK223 NNP 10_1101-2020_12_31_424971 229 7 - - HYPH 10_1101-2020_12_31_424971 229 8 3-lacZ 3-lacz CD 10_1101-2020_12_31_424971 229 9 reporter reporter NN 10_1101-2020_12_31_424971 229 10 plasmid plasmid NN 10_1101-2020_12_31_424971 229 11 that that WDT 10_1101-2020_12_31_424971 229 12 has have VBZ 10_1101-2020_12_31_424971 229 13 566 566 CD 10_1101-2020_12_31_424971 229 14 the the DT 10_1101-2020_12_31_424971 229 15 CCC CCC NNP 10_1101-2020_12_31_424971 229 16 - - HYPH 10_1101-2020_12_31_424971 229 17 C C NNP 10_1101-2020_12_31_424971 229 18 motif motif NN 10_1101-2020_12_31_424971 229 19 at at IN 10_1101-2020_12_31_424971 229 20 the the DT 10_1101-2020_12_31_424971 229 21 2nd 2nd JJ 10_1101-2020_12_31_424971 229 22 codon codon NNP 10_1101-2020_12_31_424971 229 23 position position NN 10_1101-2020_12_31_424971 229 24 of of IN 10_1101-2020_12_31_424971 229 25 the the DT 10_1101-2020_12_31_424971 229 26 lacZ lacz CD 10_1101-2020_12_31_424971 229 27 gene gene NN 10_1101-2020_12_31_424971 229 28 , , , 10_1101-2020_12_31_424971 229 29 and and CC 10_1101-2020_12_31_424971 229 30 the the DT 10_1101-2020_12_31_424971 229 31 b b NNP 10_1101-2020_12_31_424971 229 32 - - HYPH 10_1101-2020_12_31_424971 229 33 Gal Gal NNP 10_1101-2020_12_31_424971 229 34 activity activity NN 10_1101-2020_12_31_424971 229 35 was be VBD 10_1101-2020_12_31_424971 229 36 measured29 measured29 VBN 10_1101-2020_12_31_424971 229 37 . . . 10_1101-2020_12_31_424971 230 1 567 567 CD 10_1101-2020_12_31_424971 230 2 All all DT 10_1101-2020_12_31_424971 230 3 primer primer JJ 10_1101-2020_12_31_424971 230 4 sequences sequence NNS 10_1101-2020_12_31_424971 230 5 used use VBN 10_1101-2020_12_31_424971 230 6 in in IN 10_1101-2020_12_31_424971 230 7 this this DT 10_1101-2020_12_31_424971 230 8 work work NN 10_1101-2020_12_31_424971 230 9 are be VBP 10_1101-2020_12_31_424971 230 10 shown show VBN 10_1101-2020_12_31_424971 230 11 in in IN 10_1101-2020_12_31_424971 230 12 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 230 13 Table Table NNP 10_1101-2020_12_31_424971 230 14 1 1 CD 10_1101-2020_12_31_424971 230 15 . . . 10_1101-2020_12_31_424971 231 1 568 568 CD 10_1101-2020_12_31_424971 231 2 569 569 CD 10_1101-2020_12_31_424971 231 3 Preparation Preparation NNP 10_1101-2020_12_31_424971 231 4 of of IN 10_1101-2020_12_31_424971 231 5 translation translation NN 10_1101-2020_12_31_424971 231 6 components component NNS 10_1101-2020_12_31_424971 231 7 for for IN 10_1101-2020_12_31_424971 231 8 ensemble ensemble JJ 10_1101-2020_12_31_424971 231 9 biochemical biochemical JJ 10_1101-2020_12_31_424971 231 10 experiments experiment NNS 10_1101-2020_12_31_424971 231 11 . . . 10_1101-2020_12_31_424971 232 1 The the DT 10_1101-2020_12_31_424971 232 2 mRNA mrna CD 10_1101-2020_12_31_424971 232 3 570 570 CD 10_1101-2020_12_31_424971 232 4 used use VBN 10_1101-2020_12_31_424971 232 5 for for IN 10_1101-2020_12_31_424971 232 6 most most JJS 10_1101-2020_12_31_424971 232 7 in in FW 10_1101-2020_12_31_424971 232 8 vitro vitro FW 10_1101-2020_12_31_424971 232 9 translation translation NN 10_1101-2020_12_31_424971 232 10 reactions reaction NNS 10_1101-2020_12_31_424971 232 11 is be VBZ 10_1101-2020_12_31_424971 232 12 shown show VBN 10_1101-2020_12_31_424971 232 13 below below IN 10_1101-2020_12_31_424971 232 14 , , , 10_1101-2020_12_31_424971 232 15 including include VBG 10_1101-2020_12_31_424971 232 16 the the DT 10_1101-2020_12_31_424971 232 17 Shine Shine NNP 10_1101-2020_12_31_424971 232 18 - - HYPH 10_1101-2020_12_31_424971 232 19 Dalgarno Dalgarno NNP 10_1101-2020_12_31_424971 232 20 571 571 CD 10_1101-2020_12_31_424971 232 21 sequence sequence NN 10_1101-2020_12_31_424971 232 22 , , , 10_1101-2020_12_31_424971 232 23 the the DT 10_1101-2020_12_31_424971 232 24 AUG AUG NNP 10_1101-2020_12_31_424971 232 25 start start VBP 10_1101-2020_12_31_424971 232 26 codon codon NNP 10_1101-2020_12_31_424971 232 27 , , , 10_1101-2020_12_31_424971 232 28 and and CC 10_1101-2020_12_31_424971 232 29 the the DT 10_1101-2020_12_31_424971 232 30 CCC CCC NNP 10_1101-2020_12_31_424971 232 31 - - HYPH 10_1101-2020_12_31_424971 232 32 C C NNP 10_1101-2020_12_31_424971 232 33 motif motif NN 10_1101-2020_12_31_424971 232 34 : : : 10_1101-2020_12_31_424971 232 35 572 572 CD 10_1101-2020_12_31_424971 232 36 5'-GGGAAGGAGGUAAAAAUGCCCCGUUCUAAG(CAC)7 5'-GGGAAGGAGGUAAAAAUGCCCCGUUCUAAG(CAC)7 NNS 10_1101-2020_12_31_424971 232 37 . . . 10_1101-2020_12_31_424971 233 1 573 573 CD 10_1101-2020_12_31_424971 233 2 Variants variant NNS 10_1101-2020_12_31_424971 233 3 of of IN 10_1101-2020_12_31_424971 233 4 this this DT 10_1101-2020_12_31_424971 233 5 mRNA mrna NN 10_1101-2020_12_31_424971 233 6 had have VBD 10_1101-2020_12_31_424971 233 7 a a DT 10_1101-2020_12_31_424971 233 8 base base JJ 10_1101-2020_12_31_424971 233 9 substitution substitution NN 10_1101-2020_12_31_424971 233 10 in in IN 10_1101-2020_12_31_424971 233 11 the the DT 10_1101-2020_12_31_424971 233 12 CCC CCC NNP 10_1101-2020_12_31_424971 233 13 - - HYPH 10_1101-2020_12_31_424971 233 14 C C NNP 10_1101-2020_12_31_424971 233 15 motif motif NN 10_1101-2020_12_31_424971 233 16 . . . 10_1101-2020_12_31_424971 234 1 All all DT 10_1101-2020_12_31_424971 234 2 mRNAs mRNAs '' 10_1101-2020_12_31_424971 234 3 were be VBD 10_1101-2020_12_31_424971 234 4 transcribed transcribed NNP 10_1101-2020_12_31_424971 234 5 574 574 CD 10_1101-2020_12_31_424971 234 6 from from IN 10_1101-2020_12_31_424971 234 7 double double JJ 10_1101-2020_12_31_424971 234 8 - - HYPH 10_1101-2020_12_31_424971 234 9 stranded strand VBN 10_1101-2020_12_31_424971 234 10 DNA dna NN 10_1101-2020_12_31_424971 234 11 templates template NNS 10_1101-2020_12_31_424971 234 12 with with IN 10_1101-2020_12_31_424971 234 13 T7 T7 NNP 10_1101-2020_12_31_424971 234 14 RNA RNA NNP 10_1101-2020_12_31_424971 234 15 polymerase polymerase NN 10_1101-2020_12_31_424971 234 16 and and CC 10_1101-2020_12_31_424971 234 17 purified purify VBN 10_1101-2020_12_31_424971 234 18 by by IN 10_1101-2020_12_31_424971 234 19 gel gel NN 10_1101-2020_12_31_424971 234 20 575 575 CD 10_1101-2020_12_31_424971 234 21 .CC .CC : 10_1101-2020_12_31_424971 234 22 - - : 10_1101-2020_12_31_424971 234 23 BY by IN 10_1101-2020_12_31_424971 234 24 - - HYPH 10_1101-2020_12_31_424971 234 25 NC NC NNP 10_1101-2020_12_31_424971 234 26 - - HYPH 10_1101-2020_12_31_424971 234 27 ND ND NNP 10_1101-2020_12_31_424971 234 28 4.0 4.0 CD 10_1101-2020_12_31_424971 234 29 International International NNP 10_1101-2020_12_31_424971 234 30 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 234 31 under under IN 10_1101-2020_12_31_424971 234 32 a a DT 10_1101-2020_12_31_424971 234 33 ( ( -LRB- 10_1101-2020_12_31_424971 234 34 which which WDT 10_1101-2020_12_31_424971 234 35 was be VBD 10_1101-2020_12_31_424971 234 36 not not RB 10_1101-2020_12_31_424971 234 37 certified certify VBN 10_1101-2020_12_31_424971 234 38 by by IN 10_1101-2020_12_31_424971 234 39 peer peer NN 10_1101-2020_12_31_424971 234 40 review review NN 10_1101-2020_12_31_424971 234 41 ) ) -RRB- 10_1101-2020_12_31_424971 234 42 is be VBZ 10_1101-2020_12_31_424971 234 43 the the DT 10_1101-2020_12_31_424971 234 44 author author NN 10_1101-2020_12_31_424971 234 45 / / SYM 10_1101-2020_12_31_424971 234 46 funder funder NN 10_1101-2020_12_31_424971 234 47 , , , 10_1101-2020_12_31_424971 234 48 who who WP 10_1101-2020_12_31_424971 234 49 has have VBZ 10_1101-2020_12_31_424971 234 50 granted grant VBN 10_1101-2020_12_31_424971 234 51 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 234 52 a a DT 10_1101-2020_12_31_424971 234 53 license license NN 10_1101-2020_12_31_424971 234 54 to to TO 10_1101-2020_12_31_424971 234 55 display display VB 10_1101-2020_12_31_424971 234 56 the the DT 10_1101-2020_12_31_424971 234 57 preprint preprint NN 10_1101-2020_12_31_424971 234 58 in in IN 10_1101-2020_12_31_424971 234 59 perpetuity perpetuity NN 10_1101-2020_12_31_424971 234 60 . . . 10_1101-2020_12_31_424971 235 1 It -PRON- PRP 10_1101-2020_12_31_424971 235 2 is be VBZ 10_1101-2020_12_31_424971 235 3 made make VBN 10_1101-2020_12_31_424971 235 4 The the DT 10_1101-2020_12_31_424971 235 5 copyright copyright NN 10_1101-2020_12_31_424971 235 6 holder holder NN 10_1101-2020_12_31_424971 235 7 for for IN 10_1101-2020_12_31_424971 235 8 this this DT 10_1101-2020_12_31_424971 235 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 235 10 version version NN 10_1101-2020_12_31_424971 235 11 posted post VBD 10_1101-2020_12_31_424971 235 12 January January NNP 10_1101-2020_12_31_424971 235 13 2 2 CD 10_1101-2020_12_31_424971 235 14 , , , 10_1101-2020_12_31_424971 235 15 2021 2021 CD 10_1101-2020_12_31_424971 235 16 . . . 10_1101-2020_12_31_424971 235 17 ; ; : 10_1101-2020_12_31_424971 235 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 235 19 : : : 10_1101-2020_12_31_424971 235 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 235 21 preprint preprint NN 10_1101-2020_12_31_424971 235 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 235 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 235 24 24 24 CD 10_1101-2020_12_31_424971 235 25 electrophoresis electrophoresis NN 10_1101-2020_12_31_424971 235 26 . . . 10_1101-2020_12_31_424971 236 1 E. E. NNP 10_1101-2020_12_31_424971 236 2 coli coli NNS 10_1101-2020_12_31_424971 236 3 strains strain VBZ 10_1101-2020_12_31_424971 236 4 over over RB 10_1101-2020_12_31_424971 236 5 - - HYPH 10_1101-2020_12_31_424971 236 6 expressing express VBG 10_1101-2020_12_31_424971 236 7 native native JJ 10_1101-2020_12_31_424971 236 8 - - HYPH 10_1101-2020_12_31_424971 236 9 state state NN 10_1101-2020_12_31_424971 236 10 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 236 11 , , , 10_1101-2020_12_31_424971 236 12 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 236 13 ( ( -LRB- 10_1101-2020_12_31_424971 236 14 anticodon anticodon NNP 10_1101-2020_12_31_424971 236 15 ICG ICG NNP 10_1101-2020_12_31_424971 236 16 , , , 10_1101-2020_12_31_424971 236 17 576 576 CD 10_1101-2020_12_31_424971 236 18 where where WRB 10_1101-2020_12_31_424971 236 19 I -PRON- PRP 10_1101-2020_12_31_424971 236 20 = = SYM 10_1101-2020_12_31_424971 236 21 inosine inosine NNP 10_1101-2020_12_31_424971 236 22 ) ) -RRB- 10_1101-2020_12_31_424971 236 23 , , , 10_1101-2020_12_31_424971 236 24 and and CC 10_1101-2020_12_31_424971 236 25 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 236 26 ( ( -LRB- 10_1101-2020_12_31_424971 236 27 anticodon anticodon NNP 10_1101-2020_12_31_424971 236 28 U*AC U*AC NNP 10_1101-2020_12_31_424971 236 29 , , , 10_1101-2020_12_31_424971 236 30 where where WRB 10_1101-2020_12_31_424971 236 31 U u NN 10_1101-2020_12_31_424971 236 32 * * NFP 10_1101-2020_12_31_424971 236 33 = = SYM 10_1101-2020_12_31_424971 236 34 cmo5U cmo5u NN 10_1101-2020_12_31_424971 236 35 ) ) -RRB- 10_1101-2020_12_31_424971 236 36 were be VBD 10_1101-2020_12_31_424971 236 37 grown grow VBN 10_1101-2020_12_31_424971 236 38 to to IN 10_1101-2020_12_31_424971 236 39 saturation saturation NN 10_1101-2020_12_31_424971 236 40 577 577 CD 10_1101-2020_12_31_424971 236 41 and and CC 10_1101-2020_12_31_424971 236 42 were be VBD 10_1101-2020_12_31_424971 236 43 used use VBN 10_1101-2020_12_31_424971 236 44 to to TO 10_1101-2020_12_31_424971 236 45 isolate isolate VB 10_1101-2020_12_31_424971 236 46 total total JJ 10_1101-2020_12_31_424971 236 47 tRNA trna NN 10_1101-2020_12_31_424971 236 48 . . . 10_1101-2020_12_31_424971 237 1 The the DT 10_1101-2020_12_31_424971 237 2 over over RB 10_1101-2020_12_31_424971 237 3 - - HYPH 10_1101-2020_12_31_424971 237 4 expressed express VBN 10_1101-2020_12_31_424971 237 5 tRNA trna NN 10_1101-2020_12_31_424971 237 6 species specie NNS 10_1101-2020_12_31_424971 237 7 in in IN 10_1101-2020_12_31_424971 237 8 each each DT 10_1101-2020_12_31_424971 237 9 total total JJ 10_1101-2020_12_31_424971 237 10 tRNA trna NN 10_1101-2020_12_31_424971 237 11 sample sample NN 10_1101-2020_12_31_424971 237 12 578 578 CD 10_1101-2020_12_31_424971 237 13 was be VBD 10_1101-2020_12_31_424971 237 14 aminoacylated aminoacylate VBN 10_1101-2020_12_31_424971 237 15 by by IN 10_1101-2020_12_31_424971 237 16 the the DT 10_1101-2020_12_31_424971 237 17 cognate cognate NN 10_1101-2020_12_31_424971 237 18 aminoacyl aminoacyl NNP 10_1101-2020_12_31_424971 237 19 - - HYPH 10_1101-2020_12_31_424971 237 20 tRNA trna NN 10_1101-2020_12_31_424971 237 21 synthetase synthetase NN 10_1101-2020_12_31_424971 237 22 and and CC 10_1101-2020_12_31_424971 237 23 used use VBD 10_1101-2020_12_31_424971 237 24 directly directly RB 10_1101-2020_12_31_424971 237 25 in in IN 10_1101-2020_12_31_424971 237 26 the the DT 10_1101-2020_12_31_424971 237 27 TC TC NNP 10_1101-2020_12_31_424971 237 28 579 579 CD 10_1101-2020_12_31_424971 237 29 formation formation NN 10_1101-2020_12_31_424971 237 30 reaction reaction NN 10_1101-2020_12_31_424971 237 31 and and CC 10_1101-2020_12_31_424971 237 32 subsequent subsequent JJ 10_1101-2020_12_31_424971 237 33 TC TC NNP 10_1101-2020_12_31_424971 237 34 delivery delivery NN 10_1101-2020_12_31_424971 237 35 to to IN 10_1101-2020_12_31_424971 237 36 70S 70S NNP 10_1101-2020_12_31_424971 237 37 ICs ic NNS 10_1101-2020_12_31_424971 237 38 or or CC 10_1101-2020_12_31_424971 237 39 POST POST NNP 10_1101-2020_12_31_424971 237 40 complexes complex NNS 10_1101-2020_12_31_424971 237 41 . . . 10_1101-2020_12_31_424971 238 1 E. E. NNP 10_1101-2020_12_31_424971 238 2 coli coli VBZ 10_1101-2020_12_31_424971 238 3 tRNASer tRNASer NNP 10_1101-2020_12_31_424971 238 4 580 580 CD 10_1101-2020_12_31_424971 238 5 ( ( -LRB- 10_1101-2020_12_31_424971 238 6 anticodon anticodon NNP 10_1101-2020_12_31_424971 238 7 ACU ACU NNP 10_1101-2020_12_31_424971 238 8 ) ) -RRB- 10_1101-2020_12_31_424971 238 9 was be VBD 10_1101-2020_12_31_424971 238 10 prepared prepare VBN 10_1101-2020_12_31_424971 238 11 by by IN 10_1101-2020_12_31_424971 238 12 in in IN 10_1101-2020_12_31_424971 238 13 vitro vitro FW 10_1101-2020_12_31_424971 238 14 transcription transcription NN 10_1101-2020_12_31_424971 238 15 . . . 10_1101-2020_12_31_424971 239 1 Aminoacyl Aminoacyl NNP 10_1101-2020_12_31_424971 239 2 - - HYPH 10_1101-2020_12_31_424971 239 3 tRNAs trnas CD 10_1101-2020_12_31_424971 239 4 with with IN 10_1101-2020_12_31_424971 239 5 the the DT 10_1101-2020_12_31_424971 239 6 cognate cognate NN 10_1101-2020_12_31_424971 239 7 581 581 CD 10_1101-2020_12_31_424971 239 8 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 239 9 amino amino NN 10_1101-2020_12_31_424971 239 10 acid acid NN 10_1101-2020_12_31_424971 239 11 were be VBD 10_1101-2020_12_31_424971 239 12 prepared prepare VBN 10_1101-2020_12_31_424971 239 13 using use VBG 10_1101-2020_12_31_424971 239 14 the the DT 10_1101-2020_12_31_424971 239 15 respective respective JJ 10_1101-2020_12_31_424971 239 16 aminoacyl aminoacyl NN 10_1101-2020_12_31_424971 239 17 - - HYPH 10_1101-2020_12_31_424971 239 18 tRNA trna NN 10_1101-2020_12_31_424971 239 19 synthetase synthetase NN 10_1101-2020_12_31_424971 239 20 and and CC 10_1101-2020_12_31_424971 239 21 582 582 CD 10_1101-2020_12_31_424971 239 22 those those DT 10_1101-2020_12_31_424971 239 23 with with IN 10_1101-2020_12_31_424971 239 24 a a DT 10_1101-2020_12_31_424971 239 25 non non JJ 10_1101-2020_12_31_424971 239 26 - - JJ 10_1101-2020_12_31_424971 239 27 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 239 28 amino amino NN 10_1101-2020_12_31_424971 239 29 acid acid NN 10_1101-2020_12_31_424971 239 30 were be VBD 10_1101-2020_12_31_424971 239 31 prepared prepare VBN 10_1101-2020_12_31_424971 239 32 using use VBG 10_1101-2020_12_31_424971 239 33 the the DT 10_1101-2020_12_31_424971 239 34 dFx dFx `` 10_1101-2020_12_31_424971 239 35 Flexizyme Flexizyme NNPS 10_1101-2020_12_31_424971 239 36 and and CC 10_1101-2020_12_31_424971 239 37 the the DT 10_1101-2020_12_31_424971 239 38 3,5 3,5 CD 10_1101-2020_12_31_424971 239 39 - - SYM 10_1101-2020_12_31_424971 239 40 583 583 CD 10_1101-2020_12_31_424971 239 41 dinitobenzyl dinitobenzyl NN 10_1101-2020_12_31_424971 239 42 ester ester NN 10_1101-2020_12_31_424971 239 43 ( ( -LRB- 10_1101-2020_12_31_424971 239 44 DBE DBE NNP 10_1101-2020_12_31_424971 239 45 ) ) -RRB- 10_1101-2020_12_31_424971 239 46 of of IN 10_1101-2020_12_31_424971 239 47 the the DT 10_1101-2020_12_31_424971 239 48 respective respective JJ 10_1101-2020_12_31_424971 239 49 amino amino NN 10_1101-2020_12_31_424971 239 50 acid acid NN 10_1101-2020_12_31_424971 239 51 ( ( -LRB- 10_1101-2020_12_31_424971 239 52 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 239 53 Figure Figure NNP 10_1101-2020_12_31_424971 239 54 1 1 CD 10_1101-2020_12_31_424971 239 55 ) ) -RRB- 10_1101-2020_12_31_424971 239 56 . . . 10_1101-2020_12_31_424971 240 1 Aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 240 2 584 584 CD 10_1101-2020_12_31_424971 240 3 and and CC 10_1101-2020_12_31_424971 240 4 formylation formylation NN 10_1101-2020_12_31_424971 240 5 of of IN 10_1101-2020_12_31_424971 240 6 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 240 7 were be VBD 10_1101-2020_12_31_424971 240 8 performed perform VBN 10_1101-2020_12_31_424971 240 9 in in IN 10_1101-2020_12_31_424971 240 10 a a DT 10_1101-2020_12_31_424971 240 11 one one CD 10_1101-2020_12_31_424971 240 12 - - HYPH 10_1101-2020_12_31_424971 240 13 step step NN 10_1101-2020_12_31_424971 240 14 reaction reaction NN 10_1101-2020_12_31_424971 240 15 in in IN 10_1101-2020_12_31_424971 240 16 which which WDT 10_1101-2020_12_31_424971 240 17 formyl formyl NN 10_1101-2020_12_31_424971 240 18 transferase transferase NN 10_1101-2020_12_31_424971 240 19 and and CC 10_1101-2020_12_31_424971 240 20 585 585 CD 10_1101-2020_12_31_424971 240 21 the the DT 10_1101-2020_12_31_424971 240 22 methyl methyl NN 10_1101-2020_12_31_424971 240 23 donor donor NN 10_1101-2020_12_31_424971 240 24 10-formyltetrahydrofolate 10-formyltetrahydrofolate CD 10_1101-2020_12_31_424971 240 25 were be VBD 10_1101-2020_12_31_424971 240 26 added add VBN 10_1101-2020_12_31_424971 240 27 to to IN 10_1101-2020_12_31_424971 240 28 the the DT 10_1101-2020_12_31_424971 240 29 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 240 30 reaction29 reaction29 JJ 10_1101-2020_12_31_424971 240 31 . . . 10_1101-2020_12_31_424971 241 1 586 586 CD 10_1101-2020_12_31_424971 241 2 Aminoacyl Aminoacyl NNP 10_1101-2020_12_31_424971 241 3 - - HYPH 10_1101-2020_12_31_424971 241 4 tRNAs tRNAs NNP 10_1101-2020_12_31_424971 241 5 were be VBD 10_1101-2020_12_31_424971 241 6 stored store VBN 10_1101-2020_12_31_424971 241 7 in in IN 10_1101-2020_12_31_424971 241 8 25 25 CD 10_1101-2020_12_31_424971 241 9 mM mm NN 10_1101-2020_12_31_424971 241 10 sodium sodium NN 10_1101-2020_12_31_424971 241 11 acetate acetate NN 10_1101-2020_12_31_424971 241 12 ( ( -LRB- 10_1101-2020_12_31_424971 241 13 NaOAc NaOAc NNPS 10_1101-2020_12_31_424971 241 14 ) ) -RRB- 10_1101-2020_12_31_424971 241 15 ( ( -LRB- 10_1101-2020_12_31_424971 241 16 pH pH NNP 10_1101-2020_12_31_424971 241 17 5 5 CD 10_1101-2020_12_31_424971 241 18 ) ) -RRB- 10_1101-2020_12_31_424971 241 19 at at IN 10_1101-2020_12_31_424971 241 20 – – : 10_1101-2020_12_31_424971 241 21 70 70 CD 10_1101-2020_12_31_424971 241 22 ° ° , 10_1101-2020_12_31_424971 241 23 C C NNP 10_1101-2020_12_31_424971 241 24 , , , 10_1101-2020_12_31_424971 241 25 as as IN 10_1101-2020_12_31_424971 241 26 were be VBD 10_1101-2020_12_31_424971 241 27 six-587 six-587 NNP 10_1101-2020_12_31_424971 241 28 His -PRON- PRP$ 10_1101-2020_12_31_424971 241 29 - - HYPH 10_1101-2020_12_31_424971 241 30 tagged tag VBN 10_1101-2020_12_31_424971 241 31 E. E. NNP 10_1101-2020_12_31_424971 241 32 coli coli NNS 10_1101-2020_12_31_424971 241 33 initiation initiation NN 10_1101-2020_12_31_424971 241 34 and and CC 10_1101-2020_12_31_424971 241 35 elongation elongation NN 10_1101-2020_12_31_424971 241 36 factors factor NNS 10_1101-2020_12_31_424971 241 37 and and CC 10_1101-2020_12_31_424971 241 38 tight tight RB 10_1101-2020_12_31_424971 241 39 - - HYPH 10_1101-2020_12_31_424971 241 40 coupled couple VBN 10_1101-2020_12_31_424971 241 41 70S 70S NNP 10_1101-2020_12_31_424971 241 42 ribosomes ribosome NNS 10_1101-2020_12_31_424971 241 43 isolated isolate VBN 10_1101-2020_12_31_424971 241 44 from from IN 10_1101-2020_12_31_424971 241 45 588 588 CD 10_1101-2020_12_31_424971 241 46 E. E. NNP 10_1101-2020_12_31_424971 241 47 coli coli NNS 10_1101-2020_12_31_424971 241 48 MRE600 MRE600 NNP 10_1101-2020_12_31_424971 241 49 cells cell NNS 10_1101-2020_12_31_424971 241 50 . . . 10_1101-2020_12_31_424971 242 1 Recombinant recombinant VB 10_1101-2020_12_31_424971 242 2 His -PRON- PRP$ 10_1101-2020_12_31_424971 242 3 - - HYPH 10_1101-2020_12_31_424971 242 4 tagged tag VBN 10_1101-2020_12_31_424971 242 5 E. E. NNP 10_1101-2020_12_31_424971 242 6 coli coli NNS 10_1101-2020_12_31_424971 242 7 EF EF NNP 10_1101-2020_12_31_424971 242 8 - - HYPH 10_1101-2020_12_31_424971 242 9 P p NN 10_1101-2020_12_31_424971 242 10 bearing bear VBG 10_1101-2020_12_31_424971 242 11 a a DT 10_1101-2020_12_31_424971 242 12 b b NN 10_1101-2020_12_31_424971 242 13 - - HYPH 10_1101-2020_12_31_424971 242 14 lysyl lysyl JJ 10_1101-2020_12_31_424971 242 15 - - HYPH 10_1101-2020_12_31_424971 242 16 K34 K34 NNP 10_1101-2020_12_31_424971 242 17 was be VBD 10_1101-2020_12_31_424971 242 18 expressed express VBN 10_1101-2020_12_31_424971 242 19 589 589 CD 10_1101-2020_12_31_424971 242 20 and and CC 10_1101-2020_12_31_424971 242 21 purified purify VBN 10_1101-2020_12_31_424971 242 22 from from IN 10_1101-2020_12_31_424971 242 23 cells cell NNS 10_1101-2020_12_31_424971 242 24 co co JJ 10_1101-2020_12_31_424971 242 25 - - JJ 10_1101-2020_12_31_424971 242 26 expressing expressing JJ 10_1101-2020_12_31_424971 242 27 efp efp NN 10_1101-2020_12_31_424971 242 28 , , , 10_1101-2020_12_31_424971 242 29 yjeA yjea CD 10_1101-2020_12_31_424971 242 30 , , , 10_1101-2020_12_31_424971 242 31 and and CC 10_1101-2020_12_31_424971 242 32 yjeK yjeK , 10_1101-2020_12_31_424971 242 33 and and CC 10_1101-2020_12_31_424971 242 34 stored store VBD 10_1101-2020_12_31_424971 242 35 at at IN 10_1101-2020_12_31_424971 242 36 – – : 10_1101-2020_12_31_424971 242 37 20 20 CD 10_1101-2020_12_31_424971 242 38 ° ° NNS 10_1101-2020_12_31_424971 242 39 C29 C29 NNP 10_1101-2020_12_31_424971 242 40 . . . 10_1101-2020_12_31_424971 243 1 590 590 CD 10_1101-2020_12_31_424971 243 2 591 591 CD 10_1101-2020_12_31_424971 243 3 Preparation preparation NN 10_1101-2020_12_31_424971 243 4 of of IN 10_1101-2020_12_31_424971 243 5 translation translation NN 10_1101-2020_12_31_424971 243 6 components component NNS 10_1101-2020_12_31_424971 243 7 for for IN 10_1101-2020_12_31_424971 243 8 smFRET smfret NN 10_1101-2020_12_31_424971 243 9 experiments experiment NNS 10_1101-2020_12_31_424971 243 10 . . . 10_1101-2020_12_31_424971 244 1 30S 30s CD 10_1101-2020_12_31_424971 244 2 subunits subunit NNS 10_1101-2020_12_31_424971 244 3 and and CC 10_1101-2020_12_31_424971 244 4 50S 50s CD 10_1101-2020_12_31_424971 244 5 592 592 CD 10_1101-2020_12_31_424971 244 6 subunits subunit NNS 10_1101-2020_12_31_424971 244 7 lacking lack VBG 10_1101-2020_12_31_424971 244 8 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 244 9 proteins protein NNS 10_1101-2020_12_31_424971 244 10 bL9 bL9 NNP 10_1101-2020_12_31_424971 244 11 and and CC 10_1101-2020_12_31_424971 244 12 uL1 uL1 NNP 10_1101-2020_12_31_424971 244 13 were be VBD 10_1101-2020_12_31_424971 244 14 purified purify VBN 10_1101-2020_12_31_424971 244 15 from from IN 10_1101-2020_12_31_424971 244 16 a a DT 10_1101-2020_12_31_424971 244 17 previously previously RB 10_1101-2020_12_31_424971 244 18 described describe VBN 10_1101-2020_12_31_424971 244 19 bL9 bL9 NNP 10_1101-2020_12_31_424971 244 20 - - : 10_1101-2020_12_31_424971 244 21 593 593 NNP 10_1101-2020_12_31_424971 244 22 uL1 ul1 RB 10_1101-2020_12_31_424971 244 23 double double JJ 10_1101-2020_12_31_424971 244 24 deletion deletion NN 10_1101-2020_12_31_424971 244 25 E. E. NNP 10_1101-2020_12_31_424971 244 26 coli coli NNS 10_1101-2020_12_31_424971 244 27 strain35,60 strain35,60 NNP 10_1101-2020_12_31_424971 244 28 using use VBG 10_1101-2020_12_31_424971 244 29 previously previously RB 10_1101-2020_12_31_424971 244 30 described describe VBN 10_1101-2020_12_31_424971 244 31 protocols35,37,60 protocols35,37,60 NNP 10_1101-2020_12_31_424971 244 32 . . . 10_1101-2020_12_31_424971 245 1 A a DT 10_1101-2020_12_31_424971 245 2 previously previously RB 10_1101-2020_12_31_424971 245 3 594 594 CD 10_1101-2020_12_31_424971 245 4 described describe VBN 10_1101-2020_12_31_424971 245 5 single single JJ 10_1101-2020_12_31_424971 245 6 - - HYPH 10_1101-2020_12_31_424971 245 7 cysteine cysteine NN 10_1101-2020_12_31_424971 245 8 variant variant NN 10_1101-2020_12_31_424971 245 9 of of IN 10_1101-2020_12_31_424971 245 10 bL9 bl9 NN 10_1101-2020_12_31_424971 245 11 carrying carry VBG 10_1101-2020_12_31_424971 245 12 a a DT 10_1101-2020_12_31_424971 245 13 Gln Gln NNP 10_1101-2020_12_31_424971 245 14 - - HYPH 10_1101-2020_12_31_424971 245 15 to to IN 10_1101-2020_12_31_424971 245 16 - - HYPH 10_1101-2020_12_31_424971 245 17 Cys cys NN 10_1101-2020_12_31_424971 245 18 substitution substitution NN 10_1101-2020_12_31_424971 245 19 mutation mutation NN 10_1101-2020_12_31_424971 245 20 at at IN 10_1101-2020_12_31_424971 245 21 residue residue NN 10_1101-2020_12_31_424971 245 22 595 595 CD 10_1101-2020_12_31_424971 245 23 position position NN 10_1101-2020_12_31_424971 245 24 18 18 CD 10_1101-2020_12_31_424971 245 25 ( ( -LRB- 10_1101-2020_12_31_424971 245 26 bL9(Q18C))35 bL9(Q18C))35 NNP 10_1101-2020_12_31_424971 245 27 and and CC 10_1101-2020_12_31_424971 245 28 a a DT 10_1101-2020_12_31_424971 245 29 previously previously RB 10_1101-2020_12_31_424971 245 30 described describe VBN 10_1101-2020_12_31_424971 245 31 single single JJ 10_1101-2020_12_31_424971 245 32 - - HYPH 10_1101-2020_12_31_424971 245 33 cysteine cysteine NN 10_1101-2020_12_31_424971 245 34 variant variant NN 10_1101-2020_12_31_424971 245 35 of of IN 10_1101-2020_12_31_424971 245 36 uL1 uL1 NNS 10_1101-2020_12_31_424971 245 37 carrying carry VBG 10_1101-2020_12_31_424971 245 38 a a DT 10_1101-2020_12_31_424971 245 39 596 596 CD 10_1101-2020_12_31_424971 245 40 Thr Thr NNP 10_1101-2020_12_31_424971 245 41 - - HYPH 10_1101-2020_12_31_424971 245 42 to to IN 10_1101-2020_12_31_424971 245 43 - - HYPH 10_1101-2020_12_31_424971 245 44 Cys cys NN 10_1101-2020_12_31_424971 245 45 substitution substitution NN 10_1101-2020_12_31_424971 245 46 mutation mutation NN 10_1101-2020_12_31_424971 245 47 at at IN 10_1101-2020_12_31_424971 245 48 residue residue JJ 10_1101-2020_12_31_424971 245 49 position position NN 10_1101-2020_12_31_424971 245 50 202 202 CD 10_1101-2020_12_31_424971 245 51 ( ( -LRB- 10_1101-2020_12_31_424971 245 52 uL1(T202C))35,37 uL1(T202C))35,37 NNP 10_1101-2020_12_31_424971 245 53 were be VBD 10_1101-2020_12_31_424971 245 54 purified purify VBN 10_1101-2020_12_31_424971 245 55 , , , 10_1101-2020_12_31_424971 245 56 labeled label VBD 10_1101-2020_12_31_424971 245 57 597 597 CD 10_1101-2020_12_31_424971 245 58 with with IN 10_1101-2020_12_31_424971 245 59 Cy3- cy3- CD 10_1101-2020_12_31_424971 245 60 and and CC 10_1101-2020_12_31_424971 245 61 Cy5-maleimide Cy5-maleimide NNP 10_1101-2020_12_31_424971 245 62 , , , 10_1101-2020_12_31_424971 245 63 respectively respectively RB 10_1101-2020_12_31_424971 245 64 , , , 10_1101-2020_12_31_424971 245 65 to to TO 10_1101-2020_12_31_424971 245 66 generate generate VB 10_1101-2020_12_31_424971 245 67 bL9(Cy3 bl9(cy3 NN 10_1101-2020_12_31_424971 245 68 ) ) -RRB- 10_1101-2020_12_31_424971 245 69 and and CC 10_1101-2020_12_31_424971 245 70 uL1(Cy5 ul1(cy5 NN 10_1101-2020_12_31_424971 245 71 ) ) -RRB- 10_1101-2020_12_31_424971 245 72 , , , 10_1101-2020_12_31_424971 245 73 and and CC 10_1101-2020_12_31_424971 245 74 598 598 CD 10_1101-2020_12_31_424971 245 75 reconstituted reconstitute VBN 10_1101-2020_12_31_424971 245 76 into into IN 10_1101-2020_12_31_424971 245 77 the the DT 10_1101-2020_12_31_424971 245 78 50S 50S NNP 10_1101-2020_12_31_424971 245 79 subunits subunit NNS 10_1101-2020_12_31_424971 245 80 lacking lack VBG 10_1101-2020_12_31_424971 245 81 bL9 bl9 NN 10_1101-2020_12_31_424971 245 82 and and CC 10_1101-2020_12_31_424971 245 83 uL1 ul1 RB 10_1101-2020_12_31_424971 245 84 following follow VBG 10_1101-2020_12_31_424971 245 85 previously previously RB 10_1101-2020_12_31_424971 245 86 described describe VBN 10_1101-2020_12_31_424971 245 87 599 599 CD 10_1101-2020_12_31_424971 245 88 protocols35 protocols35 NNS 10_1101-2020_12_31_424971 245 89 . . . 10_1101-2020_12_31_424971 246 1 The the DT 10_1101-2020_12_31_424971 246 2 reconstituted reconstituted JJ 10_1101-2020_12_31_424971 246 3 bL9(Cy3)- bl9(cy3)- NN 10_1101-2020_12_31_424971 246 4 and and CC 10_1101-2020_12_31_424971 246 5 uL1(Cy5)-labeled ul1(cy5)-labeled CD 10_1101-2020_12_31_424971 246 6 50S 50s CD 10_1101-2020_12_31_424971 246 7 subunits subunit NNS 10_1101-2020_12_31_424971 246 8 were be VBD 10_1101-2020_12_31_424971 246 9 then then RB 10_1101-2020_12_31_424971 246 10 re re NN 10_1101-2020_12_31_424971 246 11 - - VBN 10_1101-2020_12_31_424971 246 12 purified purify VBD 10_1101-2020_12_31_424971 246 13 600 600 CD 10_1101-2020_12_31_424971 246 14 .CC .CC , 10_1101-2020_12_31_424971 246 15 - - HYPH 10_1101-2020_12_31_424971 246 16 BY by IN 10_1101-2020_12_31_424971 246 17 - - HYPH 10_1101-2020_12_31_424971 246 18 NC NC NNP 10_1101-2020_12_31_424971 246 19 - - HYPH 10_1101-2020_12_31_424971 246 20 ND ND NNP 10_1101-2020_12_31_424971 246 21 4.0 4.0 CD 10_1101-2020_12_31_424971 246 22 International International NNP 10_1101-2020_12_31_424971 246 23 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 246 24 under under IN 10_1101-2020_12_31_424971 246 25 a a DT 10_1101-2020_12_31_424971 246 26 ( ( -LRB- 10_1101-2020_12_31_424971 246 27 which which WDT 10_1101-2020_12_31_424971 246 28 was be VBD 10_1101-2020_12_31_424971 246 29 not not RB 10_1101-2020_12_31_424971 246 30 certified certify VBN 10_1101-2020_12_31_424971 246 31 by by IN 10_1101-2020_12_31_424971 246 32 peer peer NN 10_1101-2020_12_31_424971 246 33 review review NN 10_1101-2020_12_31_424971 246 34 ) ) -RRB- 10_1101-2020_12_31_424971 246 35 is be VBZ 10_1101-2020_12_31_424971 246 36 the the DT 10_1101-2020_12_31_424971 246 37 author author NN 10_1101-2020_12_31_424971 246 38 / / SYM 10_1101-2020_12_31_424971 246 39 funder funder NN 10_1101-2020_12_31_424971 246 40 , , , 10_1101-2020_12_31_424971 246 41 who who WP 10_1101-2020_12_31_424971 246 42 has have VBZ 10_1101-2020_12_31_424971 246 43 granted grant VBN 10_1101-2020_12_31_424971 246 44 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 246 45 a a DT 10_1101-2020_12_31_424971 246 46 license license NN 10_1101-2020_12_31_424971 246 47 to to TO 10_1101-2020_12_31_424971 246 48 display display VB 10_1101-2020_12_31_424971 246 49 the the DT 10_1101-2020_12_31_424971 246 50 preprint preprint NN 10_1101-2020_12_31_424971 246 51 in in IN 10_1101-2020_12_31_424971 246 52 perpetuity perpetuity NN 10_1101-2020_12_31_424971 246 53 . . . 10_1101-2020_12_31_424971 247 1 It -PRON- PRP 10_1101-2020_12_31_424971 247 2 is be VBZ 10_1101-2020_12_31_424971 247 3 made make VBN 10_1101-2020_12_31_424971 247 4 The the DT 10_1101-2020_12_31_424971 247 5 copyright copyright NN 10_1101-2020_12_31_424971 247 6 holder holder NN 10_1101-2020_12_31_424971 247 7 for for IN 10_1101-2020_12_31_424971 247 8 this this DT 10_1101-2020_12_31_424971 247 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 247 10 version version NN 10_1101-2020_12_31_424971 247 11 posted post VBD 10_1101-2020_12_31_424971 247 12 January January NNP 10_1101-2020_12_31_424971 247 13 2 2 CD 10_1101-2020_12_31_424971 247 14 , , , 10_1101-2020_12_31_424971 247 15 2021 2021 CD 10_1101-2020_12_31_424971 247 16 . . . 10_1101-2020_12_31_424971 247 17 ; ; : 10_1101-2020_12_31_424971 247 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 247 19 : : : 10_1101-2020_12_31_424971 247 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 247 21 preprint preprint NN 10_1101-2020_12_31_424971 247 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 247 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 247 24 25 25 CD 10_1101-2020_12_31_424971 247 25 using use VBG 10_1101-2020_12_31_424971 247 26 sucrose sucrose JJ 10_1101-2020_12_31_424971 247 27 density density NN 10_1101-2020_12_31_424971 247 28 gradient gradient NN 10_1101-2020_12_31_424971 247 29 ultracentrifugation35,43 ultracentrifugation35,43 NNP 10_1101-2020_12_31_424971 247 30 . . . 10_1101-2020_12_31_424971 248 1 50S 50s CD 10_1101-2020_12_31_424971 248 2 subunits subunit NNS 10_1101-2020_12_31_424971 248 3 lacking lack VBG 10_1101-2020_12_31_424971 248 4 bL9(Cy3 bl9(cy3 NN 10_1101-2020_12_31_424971 248 5 ) ) -RRB- 10_1101-2020_12_31_424971 248 6 and/or and/or CC 10_1101-2020_12_31_424971 248 7 601 601 CD 10_1101-2020_12_31_424971 248 8 uL1(Cy5 ul1(cy5 CD 10_1101-2020_12_31_424971 248 9 ) ) -RRB- 10_1101-2020_12_31_424971 248 10 or or CC 10_1101-2020_12_31_424971 248 11 harboring harbor VBG 10_1101-2020_12_31_424971 248 12 unlabeled unlabeled JJ 10_1101-2020_12_31_424971 248 13 bL9 bl9 NN 10_1101-2020_12_31_424971 248 14 and/or and/or CC 10_1101-2020_12_31_424971 248 15 uL1 ul1 NN 10_1101-2020_12_31_424971 248 16 do do VBP 10_1101-2020_12_31_424971 248 17 not not RB 10_1101-2020_12_31_424971 248 18 generate generate VB 10_1101-2020_12_31_424971 248 19 bL9(Cy3)-uL1(Cy5 bl9(cy3)-ul1(cy5 NN 10_1101-2020_12_31_424971 248 20 ) ) -RRB- 10_1101-2020_12_31_424971 248 21 smFRET smfret NN 10_1101-2020_12_31_424971 248 22 602 602 CD 10_1101-2020_12_31_424971 248 23 signals signal NNS 10_1101-2020_12_31_424971 248 24 and and CC 10_1101-2020_12_31_424971 248 25 therefore therefore RB 10_1101-2020_12_31_424971 248 26 do do VBP 10_1101-2020_12_31_424971 248 27 not not RB 10_1101-2020_12_31_424971 248 28 affect affect VB 10_1101-2020_12_31_424971 248 29 data datum NNS 10_1101-2020_12_31_424971 248 30 collection collection NN 10_1101-2020_12_31_424971 248 31 or or CC 10_1101-2020_12_31_424971 248 32 analysis analysis NN 10_1101-2020_12_31_424971 248 33 . . . 10_1101-2020_12_31_424971 249 1 Previously previously RB 10_1101-2020_12_31_424971 249 2 , , , 10_1101-2020_12_31_424971 249 3 we -PRON- PRP 10_1101-2020_12_31_424971 249 4 have have VBP 10_1101-2020_12_31_424971 249 5 shown show VBN 10_1101-2020_12_31_424971 249 6 that that IN 10_1101-2020_12_31_424971 249 7 603 603 CD 10_1101-2020_12_31_424971 249 8 70S 70S NNP 10_1101-2020_12_31_424971 249 9 ICs ic NNS 10_1101-2020_12_31_424971 249 10 formed form VBD 10_1101-2020_12_31_424971 249 11 with with IN 10_1101-2020_12_31_424971 249 12 these these DT 10_1101-2020_12_31_424971 249 13 bL9(Cy3)- bL9(Cy3)- NNS 10_1101-2020_12_31_424971 249 14 and and CC 10_1101-2020_12_31_424971 249 15 uL1(Cy5)-containing ul1(cy5)-containing CD 10_1101-2020_12_31_424971 249 16 50S 50s CD 10_1101-2020_12_31_424971 249 17 subunits subunit NNS 10_1101-2020_12_31_424971 249 18 can can MD 10_1101-2020_12_31_424971 249 19 undergo undergo VB 10_1101-2020_12_31_424971 249 20 604 604 CD 10_1101-2020_12_31_424971 249 21 peptide peptide NN 10_1101-2020_12_31_424971 249 22 - - HYPH 10_1101-2020_12_31_424971 249 23 bond bond NN 10_1101-2020_12_31_424971 249 24 formation formation NN 10_1101-2020_12_31_424971 249 25 and and CC 10_1101-2020_12_31_424971 249 26 two two CD 10_1101-2020_12_31_424971 249 27 rounds round NNS 10_1101-2020_12_31_424971 249 28 of of IN 10_1101-2020_12_31_424971 249 29 translocation translocation NN 10_1101-2020_12_31_424971 249 30 elongation elongation NN 10_1101-2020_12_31_424971 249 31 with with IN 10_1101-2020_12_31_424971 249 32 similar similar JJ 10_1101-2020_12_31_424971 249 33 efficiency efficiency NN 10_1101-2020_12_31_424971 249 34 as as IN 10_1101-2020_12_31_424971 249 35 70S 70S NNP 10_1101-2020_12_31_424971 249 36 605 605 CD 10_1101-2020_12_31_424971 249 37 ICs ic NNS 10_1101-2020_12_31_424971 249 38 formed form VBN 10_1101-2020_12_31_424971 249 39 with with IN 10_1101-2020_12_31_424971 249 40 wild wild JJ 10_1101-2020_12_31_424971 249 41 - - HYPH 10_1101-2020_12_31_424971 249 42 type type NN 10_1101-2020_12_31_424971 249 43 50S 50S NNP 10_1101-2020_12_31_424971 249 44 subunits35 subunits35 NNP 10_1101-2020_12_31_424971 249 45 . . . 10_1101-2020_12_31_424971 250 1 606 606 CD 10_1101-2020_12_31_424971 250 2 The the DT 10_1101-2020_12_31_424971 250 3 sequence sequence NN 10_1101-2020_12_31_424971 250 4 of of IN 10_1101-2020_12_31_424971 250 5 the the DT 10_1101-2020_12_31_424971 250 6 mRNA mrna NN 10_1101-2020_12_31_424971 250 7 used use VBN 10_1101-2020_12_31_424971 250 8 for for IN 10_1101-2020_12_31_424971 250 9 assembling assemble VBG 10_1101-2020_12_31_424971 250 10 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 250 11 complexes complex NNS 10_1101-2020_12_31_424971 250 12 for for IN 10_1101-2020_12_31_424971 250 13 smFRET smFRET NNP 10_1101-2020_12_31_424971 250 14 studies study NNS 10_1101-2020_12_31_424971 250 15 607 607 CD 10_1101-2020_12_31_424971 250 16 is be VBZ 10_1101-2020_12_31_424971 250 17 shown show VBN 10_1101-2020_12_31_424971 250 18 below below IN 10_1101-2020_12_31_424971 250 19 , , , 10_1101-2020_12_31_424971 250 20 including include VBG 10_1101-2020_12_31_424971 250 21 the the DT 10_1101-2020_12_31_424971 250 22 Shine Shine NNP 10_1101-2020_12_31_424971 250 23 - - HYPH 10_1101-2020_12_31_424971 250 24 Dalgarno Dalgarno NNP 10_1101-2020_12_31_424971 250 25 sequence sequence NN 10_1101-2020_12_31_424971 250 26 , , , 10_1101-2020_12_31_424971 250 27 the the DT 10_1101-2020_12_31_424971 250 28 AUG AUG NNP 10_1101-2020_12_31_424971 250 29 start start VBP 10_1101-2020_12_31_424971 250 30 codon codon NNP 10_1101-2020_12_31_424971 250 31 , , , 10_1101-2020_12_31_424971 250 32 and and CC 10_1101-2020_12_31_424971 250 33 the the DT 10_1101-2020_12_31_424971 250 34 CCC CCC NNP 10_1101-2020_12_31_424971 250 35 - - HYPH 10_1101-2020_12_31_424971 250 36 C C NNP 10_1101-2020_12_31_424971 250 37 608 608 CD 10_1101-2020_12_31_424971 250 38 motif motif NN 10_1101-2020_12_31_424971 250 39 : : : 10_1101-2020_12_31_424971 250 40 609 609 CD 10_1101-2020_12_31_424971 250 41 5'-GCAACCUAAAACUCACACAGGGCCCUAAGGACAUAAAAAUGCCCCGUU 5'-gcaaccuaaaacucacacagggcccuaaggacauaaaaaugccccguu CD 10_1101-2020_12_31_424971 250 42 610 610 CD 10_1101-2020_12_31_424971 250 43 AUCCUCCUGCUGCACUCGCUGCACAAAUCGCUCAACGGCAAUUAAGGA AUCCUCCUGCUGCACUCGCUGCACAAAUCGCUCAACGGCAAUUAAGGA NNP 10_1101-2020_12_31_424971 250 44 . . . 10_1101-2020_12_31_424971 251 1 611 611 CD 10_1101-2020_12_31_424971 251 2 The the DT 10_1101-2020_12_31_424971 251 3 mRNA mrna NN 10_1101-2020_12_31_424971 251 4 was be VBD 10_1101-2020_12_31_424971 251 5 synthesized synthesize VBN 10_1101-2020_12_31_424971 251 6 by by IN 10_1101-2020_12_31_424971 251 7 in in IN 10_1101-2020_12_31_424971 251 8 vitro vitro FW 10_1101-2020_12_31_424971 251 9 transcription transcription NN 10_1101-2020_12_31_424971 251 10 using use VBG 10_1101-2020_12_31_424971 251 11 T7 T7 NNP 10_1101-2020_12_31_424971 251 12 RNA RNA NNP 10_1101-2020_12_31_424971 251 13 polymerase polymerase NN 10_1101-2020_12_31_424971 251 14 , , , 10_1101-2020_12_31_424971 251 15 and and CC 10_1101-2020_12_31_424971 251 16 then then RB 10_1101-2020_12_31_424971 251 17 612 612 CD 10_1101-2020_12_31_424971 251 18 hybridized hybridize VBD 10_1101-2020_12_31_424971 251 19 to to IN 10_1101-2020_12_31_424971 251 20 a a DT 10_1101-2020_12_31_424971 251 21 previously previously RB 10_1101-2020_12_31_424971 251 22 described describe VBN 10_1101-2020_12_31_424971 251 23 3’-biotinylated 3’-biotinylated CD 10_1101-2020_12_31_424971 251 24 DNA dna NN 10_1101-2020_12_31_424971 251 25 oligonucleotide oligonucleotide RB 10_1101-2020_12_31_424971 251 26 ( ( -LRB- 10_1101-2020_12_31_424971 251 27 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 251 28 Table Table NNP 10_1101-2020_12_31_424971 251 29 613 613 CD 10_1101-2020_12_31_424971 251 30 1 1 CD 10_1101-2020_12_31_424971 251 31 ) ) -RRB- 10_1101-2020_12_31_424971 251 32 that that WDT 10_1101-2020_12_31_424971 251 33 was be VBD 10_1101-2020_12_31_424971 251 34 complementary complementary JJ 10_1101-2020_12_31_424971 251 35 to to IN 10_1101-2020_12_31_424971 251 36 the the DT 10_1101-2020_12_31_424971 251 37 5 5 CD 10_1101-2020_12_31_424971 251 38 ' ' '' 10_1101-2020_12_31_424971 251 39 end end NN 10_1101-2020_12_31_424971 251 40 of of IN 10_1101-2020_12_31_424971 251 41 the the DT 10_1101-2020_12_31_424971 251 42 mRNA mrna NN 10_1101-2020_12_31_424971 251 43 and and CC 10_1101-2020_12_31_424971 251 44 was be VBD 10_1101-2020_12_31_424971 251 45 chemically chemically RB 10_1101-2020_12_31_424971 251 46 synthesized synthesize VBN 10_1101-2020_12_31_424971 251 47 by by IN 10_1101-2020_12_31_424971 251 48 614 614 CD 10_1101-2020_12_31_424971 251 49 Integrated Integrated NNP 10_1101-2020_12_31_424971 251 50 DNA DNA NNP 10_1101-2020_12_31_424971 251 51 Technologies60 Technologies60 NNP 10_1101-2020_12_31_424971 251 52 . . . 10_1101-2020_12_31_424971 252 1 Hybridized hybridize VBN 10_1101-2020_12_31_424971 252 2 mRNA mrna RB 10_1101-2020_12_31_424971 252 3 : : : 10_1101-2020_12_31_424971 252 4 DNA dna NN 10_1101-2020_12_31_424971 252 5 - - HYPH 10_1101-2020_12_31_424971 252 6 biotin biotin JJ 10_1101-2020_12_31_424971 252 7 complexes complex NNS 10_1101-2020_12_31_424971 252 8 were be VBD 10_1101-2020_12_31_424971 252 9 stored store VBN 10_1101-2020_12_31_424971 252 10 in in IN 10_1101-2020_12_31_424971 252 11 10 10 CD 10_1101-2020_12_31_424971 252 12 mM mM NNP 10_1101-2020_12_31_424971 252 13 615 615 CD 10_1101-2020_12_31_424971 252 14 Tris Tris NNP 10_1101-2020_12_31_424971 252 15 - - HYPH 10_1101-2020_12_31_424971 252 16 OAc OAc NNS 10_1101-2020_12_31_424971 252 17 ( ( -LRB- 10_1101-2020_12_31_424971 252 18 pH pH NNP 10_1101-2020_12_31_424971 252 19 = = SYM 10_1101-2020_12_31_424971 252 20 7.5 7.5 CD 10_1101-2020_12_31_424971 252 21 at at IN 10_1101-2020_12_31_424971 252 22 37 37 CD 10_1101-2020_12_31_424971 252 23 ºC ºC NNP 10_1101-2020_12_31_424971 252 24 ) ) -RRB- 10_1101-2020_12_31_424971 252 25 , , , 10_1101-2020_12_31_424971 252 26 1 1 CD 10_1101-2020_12_31_424971 252 27 mM mm NN 10_1101-2020_12_31_424971 252 28 EDTA edta NN 10_1101-2020_12_31_424971 252 29 , , , 10_1101-2020_12_31_424971 252 30 and and CC 10_1101-2020_12_31_424971 252 31 10 10 CD 10_1101-2020_12_31_424971 252 32 mM mm NN 10_1101-2020_12_31_424971 252 33 KCl KCl NNS 10_1101-2020_12_31_424971 252 34 at at IN 10_1101-2020_12_31_424971 252 35 – – : 10_1101-2020_12_31_424971 252 36 80 80 CD 10_1101-2020_12_31_424971 252 37 ºC ºC NNP 10_1101-2020_12_31_424971 252 38 until until IN 10_1101-2020_12_31_424971 252 39 they -PRON- PRP 10_1101-2020_12_31_424971 252 40 were be VBD 10_1101-2020_12_31_424971 252 41 used use VBN 10_1101-2020_12_31_424971 252 42 in in IN 10_1101-2020_12_31_424971 252 43 616 616 CD 10_1101-2020_12_31_424971 252 44 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 252 45 complex complex JJ 10_1101-2020_12_31_424971 252 46 assembly assembly NN 10_1101-2020_12_31_424971 252 47 . . . 10_1101-2020_12_31_424971 253 1 Aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 253 2 and and CC 10_1101-2020_12_31_424971 253 3 formylation formylation NN 10_1101-2020_12_31_424971 253 4 of of IN 10_1101-2020_12_31_424971 253 5 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 253 6 ( ( -LRB- 10_1101-2020_12_31_424971 253 7 purchased purchase VBN 10_1101-2020_12_31_424971 253 8 from from IN 10_1101-2020_12_31_424971 253 9 MP MP NNP 10_1101-2020_12_31_424971 253 10 617 617 CD 10_1101-2020_12_31_424971 253 11 Biomedicals Biomedicals NNPS 10_1101-2020_12_31_424971 253 12 ) ) -RRB- 10_1101-2020_12_31_424971 253 13 was be VBD 10_1101-2020_12_31_424971 253 14 achieved achieve VBN 10_1101-2020_12_31_424971 253 15 simultaneously simultaneously RB 10_1101-2020_12_31_424971 253 16 using use VBG 10_1101-2020_12_31_424971 253 17 E. E. NNP 10_1101-2020_12_31_424971 253 18 coli coli NNS 10_1101-2020_12_31_424971 253 19 methionyl methionyl NN 10_1101-2020_12_31_424971 253 20 - - HYPH 10_1101-2020_12_31_424971 253 21 tRNA trna JJ 10_1101-2020_12_31_424971 253 22 synthetase synthetase NN 10_1101-2020_12_31_424971 253 23 and and CC 10_1101-2020_12_31_424971 253 24 E. E. NNP 10_1101-2020_12_31_424971 253 25 coli coli VBZ 10_1101-2020_12_31_424971 253 26 618 618 CD 10_1101-2020_12_31_424971 253 27 formylmethionyl formylmethionyl NN 10_1101-2020_12_31_424971 253 28 - - HYPH 10_1101-2020_12_31_424971 253 29 tRNA trna JJ 10_1101-2020_12_31_424971 253 30 formyltransferase60 formyltransferase60 NN 10_1101-2020_12_31_424971 253 31 . . . 10_1101-2020_12_31_424971 254 1 Expression expression NN 10_1101-2020_12_31_424971 254 2 and and CC 10_1101-2020_12_31_424971 254 3 purification purification NN 10_1101-2020_12_31_424971 254 4 of of IN 10_1101-2020_12_31_424971 254 5 IF1 IF1 NNP 10_1101-2020_12_31_424971 254 6 , , , 10_1101-2020_12_31_424971 254 7 IF2 IF2 NNP 10_1101-2020_12_31_424971 254 8 , , , 10_1101-2020_12_31_424971 254 9 IF3 IF3 NNP 10_1101-2020_12_31_424971 254 10 , , , 10_1101-2020_12_31_424971 254 11 EF EF NNP 10_1101-2020_12_31_424971 254 12 - - HYPH 10_1101-2020_12_31_424971 254 13 Tu Tu NNP 10_1101-2020_12_31_424971 254 14 , , , 10_1101-2020_12_31_424971 254 15 619 619 CD 10_1101-2020_12_31_424971 254 16 EF EF NNP 10_1101-2020_12_31_424971 254 17 - - HYPH 10_1101-2020_12_31_424971 254 18 Ts Ts NNP 10_1101-2020_12_31_424971 254 19 , , , 10_1101-2020_12_31_424971 254 20 and and CC 10_1101-2020_12_31_424971 254 21 EF EF NNP 10_1101-2020_12_31_424971 254 22 - - HYPH 10_1101-2020_12_31_424971 254 23 G G NNP 10_1101-2020_12_31_424971 254 24 were be VBD 10_1101-2020_12_31_424971 254 25 following follow VBG 10_1101-2020_12_31_424971 254 26 previously previously RB 10_1101-2020_12_31_424971 254 27 published publish VBN 10_1101-2020_12_31_424971 254 28 procedures60 procedures60 NNS 10_1101-2020_12_31_424971 254 29 . . . 10_1101-2020_12_31_424971 255 1 620 620 CD 10_1101-2020_12_31_424971 255 2 621 621 CD 10_1101-2020_12_31_424971 255 3 Preparation preparation NN 10_1101-2020_12_31_424971 255 4 and and CC 10_1101-2020_12_31_424971 255 5 purification purification NN 10_1101-2020_12_31_424971 255 6 of of IN 10_1101-2020_12_31_424971 255 7 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 255 8 and and CC 10_1101-2020_12_31_424971 255 9 ProL. ProL. . 10_1101-2020_12_31_424971 256 1 Native native JJ 10_1101-2020_12_31_424971 256 2 - - HYPH 10_1101-2020_12_31_424971 256 3 state state NN 10_1101-2020_12_31_424971 256 4 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 256 5 was be VBD 10_1101-2020_12_31_424971 256 6 isolated isolate VBN 10_1101-2020_12_31_424971 256 7 from from IN 10_1101-2020_12_31_424971 256 8 a a DT 10_1101-2020_12_31_424971 256 9 622 622 CD 10_1101-2020_12_31_424971 256 10 derivative derivative NN 10_1101-2020_12_31_424971 256 11 of of IN 10_1101-2020_12_31_424971 256 12 E. E. NNP 10_1101-2020_12_31_424971 256 13 coli coli NNS 10_1101-2020_12_31_424971 256 14 JM109 JM109 NNP 10_1101-2020_12_31_424971 256 15 lacking lack VBG 10_1101-2020_12_31_424971 256 16 the the DT 10_1101-2020_12_31_424971 256 17 endogenous endogenous JJ 10_1101-2020_12_31_424971 256 18 ProL prol NN 10_1101-2020_12_31_424971 256 19 , , , 10_1101-2020_12_31_424971 256 20 but but CC 10_1101-2020_12_31_424971 256 21 expressing express VBG 10_1101-2020_12_31_424971 256 22 SufB2 sufb2 NN 10_1101-2020_12_31_424971 256 23 from from IN 10_1101-2020_12_31_424971 256 24 the the DT 10_1101-2020_12_31_424971 256 25 pKK223 pKK223 NNP 10_1101-2020_12_31_424971 256 26 - - HYPH 10_1101-2020_12_31_424971 256 27 623 623 CD 10_1101-2020_12_31_424971 256 28 3 3 CD 10_1101-2020_12_31_424971 256 29 plasmid plasmid NN 10_1101-2020_12_31_424971 256 30 ( ( -LRB- 10_1101-2020_12_31_424971 256 31 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 256 32 Table Table NNP 10_1101-2020_12_31_424971 256 33 1 1 CD 10_1101-2020_12_31_424971 256 34 ) ) -RRB- 10_1101-2020_12_31_424971 256 35 , , , 10_1101-2020_12_31_424971 256 36 while while IN 10_1101-2020_12_31_424971 256 37 native native JJ 10_1101-2020_12_31_424971 256 38 - - HYPH 10_1101-2020_12_31_424971 256 39 state state NN 10_1101-2020_12_31_424971 256 40 ProL ProL NNP 10_1101-2020_12_31_424971 256 41 was be VBD 10_1101-2020_12_31_424971 256 42 purified purify VBN 10_1101-2020_12_31_424971 256 43 from from IN 10_1101-2020_12_31_424971 256 44 total total JJ 10_1101-2020_12_31_424971 256 45 tRNA trna NN 10_1101-2020_12_31_424971 256 46 isolated isolate VBD 10_1101-2020_12_31_424971 256 47 624 624 CD 10_1101-2020_12_31_424971 256 48 from from IN 10_1101-2020_12_31_424971 256 49 E. E. NNP 10_1101-2020_12_31_424971 256 50 coli coli NNS 10_1101-2020_12_31_424971 256 51 JM109 JM109 NNP 10_1101-2020_12_31_424971 256 52 cells cell NNS 10_1101-2020_12_31_424971 256 53 over over IN 10_1101-2020_12_31_424971 256 54 - - HYPH 10_1101-2020_12_31_424971 256 55 expressing express VBG 10_1101-2020_12_31_424971 256 56 ProL prol NN 10_1101-2020_12_31_424971 256 57 from from IN 10_1101-2020_12_31_424971 256 58 the the DT 10_1101-2020_12_31_424971 256 59 pKK223 pKK223 NNP 10_1101-2020_12_31_424971 256 60 - - HYPH 10_1101-2020_12_31_424971 256 61 3 3 CD 10_1101-2020_12_31_424971 256 62 plasmid plasmid NN 10_1101-2020_12_31_424971 256 63 . . . 10_1101-2020_12_31_424971 257 1 The the DT 10_1101-2020_12_31_424971 257 2 ProL ProL NNP 10_1101-2020_12_31_424971 257 3 - - HYPH 10_1101-2020_12_31_424971 257 4 KO KO NNP 10_1101-2020_12_31_424971 257 5 strain strain VBP 10_1101-2020_12_31_424971 257 6 625 625 CD 10_1101-2020_12_31_424971 257 7 lacking lack VBG 10_1101-2020_12_31_424971 257 8 the the DT 10_1101-2020_12_31_424971 257 9 endogenous endogenous JJ 10_1101-2020_12_31_424971 257 10 ProL ProL NNP 10_1101-2020_12_31_424971 257 11 was be VBD 10_1101-2020_12_31_424971 257 12 described describe VBN 10_1101-2020_12_31_424971 257 13 previously30 previously30 NNP 10_1101-2020_12_31_424971 257 14 . . . 10_1101-2020_12_31_424971 258 1 Each each DT 10_1101-2020_12_31_424971 258 2 native native JJ 10_1101-2020_12_31_424971 258 3 - - HYPH 10_1101-2020_12_31_424971 258 4 state state NN 10_1101-2020_12_31_424971 258 5 tRNA trna NN 10_1101-2020_12_31_424971 258 6 was be VBD 10_1101-2020_12_31_424971 258 7 isolated isolate VBN 10_1101-2020_12_31_424971 258 8 626 626 CD 10_1101-2020_12_31_424971 258 9 .CC .CC , 10_1101-2020_12_31_424971 258 10 - - HYPH 10_1101-2020_12_31_424971 258 11 BY by IN 10_1101-2020_12_31_424971 258 12 - - HYPH 10_1101-2020_12_31_424971 258 13 NC NC NNP 10_1101-2020_12_31_424971 258 14 - - HYPH 10_1101-2020_12_31_424971 258 15 ND ND NNP 10_1101-2020_12_31_424971 258 16 4.0 4.0 CD 10_1101-2020_12_31_424971 258 17 International International NNP 10_1101-2020_12_31_424971 258 18 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 258 19 under under IN 10_1101-2020_12_31_424971 258 20 a a DT 10_1101-2020_12_31_424971 258 21 ( ( -LRB- 10_1101-2020_12_31_424971 258 22 which which WDT 10_1101-2020_12_31_424971 258 23 was be VBD 10_1101-2020_12_31_424971 258 24 not not RB 10_1101-2020_12_31_424971 258 25 certified certify VBN 10_1101-2020_12_31_424971 258 26 by by IN 10_1101-2020_12_31_424971 258 27 peer peer NN 10_1101-2020_12_31_424971 258 28 review review NN 10_1101-2020_12_31_424971 258 29 ) ) -RRB- 10_1101-2020_12_31_424971 258 30 is be VBZ 10_1101-2020_12_31_424971 258 31 the the DT 10_1101-2020_12_31_424971 258 32 author author NN 10_1101-2020_12_31_424971 258 33 / / SYM 10_1101-2020_12_31_424971 258 34 funder funder NN 10_1101-2020_12_31_424971 258 35 , , , 10_1101-2020_12_31_424971 258 36 who who WP 10_1101-2020_12_31_424971 258 37 has have VBZ 10_1101-2020_12_31_424971 258 38 granted grant VBN 10_1101-2020_12_31_424971 258 39 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 258 40 a a DT 10_1101-2020_12_31_424971 258 41 license license NN 10_1101-2020_12_31_424971 258 42 to to TO 10_1101-2020_12_31_424971 258 43 display display VB 10_1101-2020_12_31_424971 258 44 the the DT 10_1101-2020_12_31_424971 258 45 preprint preprint NN 10_1101-2020_12_31_424971 258 46 in in IN 10_1101-2020_12_31_424971 258 47 perpetuity perpetuity NN 10_1101-2020_12_31_424971 258 48 . . . 10_1101-2020_12_31_424971 259 1 It -PRON- PRP 10_1101-2020_12_31_424971 259 2 is be VBZ 10_1101-2020_12_31_424971 259 3 made make VBN 10_1101-2020_12_31_424971 259 4 The the DT 10_1101-2020_12_31_424971 259 5 copyright copyright NN 10_1101-2020_12_31_424971 259 6 holder holder NN 10_1101-2020_12_31_424971 259 7 for for IN 10_1101-2020_12_31_424971 259 8 this this DT 10_1101-2020_12_31_424971 259 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 259 10 version version NN 10_1101-2020_12_31_424971 259 11 posted post VBD 10_1101-2020_12_31_424971 259 12 January January NNP 10_1101-2020_12_31_424971 259 13 2 2 CD 10_1101-2020_12_31_424971 259 14 , , , 10_1101-2020_12_31_424971 259 15 2021 2021 CD 10_1101-2020_12_31_424971 259 16 . . . 10_1101-2020_12_31_424971 259 17 ; ; : 10_1101-2020_12_31_424971 259 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 259 19 : : : 10_1101-2020_12_31_424971 259 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 259 21 preprint preprint NN 10_1101-2020_12_31_424971 259 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 259 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 259 24 26 26 CD 10_1101-2020_12_31_424971 259 25 by by IN 10_1101-2020_12_31_424971 259 26 a a DT 10_1101-2020_12_31_424971 259 27 biotinylated biotinylated JJ 10_1101-2020_12_31_424971 259 28 capture capture NN 10_1101-2020_12_31_424971 259 29 probe probe NN 10_1101-2020_12_31_424971 259 30 attached attach VBN 10_1101-2020_12_31_424971 259 31 to to IN 10_1101-2020_12_31_424971 259 32 streptavidin streptavidin NNP 10_1101-2020_12_31_424971 259 33 - - HYPH 10_1101-2020_12_31_424971 259 34 derivatized derivatize VBN 10_1101-2020_12_31_424971 259 35 Sepharose Sepharose NNP 10_1101-2020_12_31_424971 259 36 beads29 beads29 NNP 10_1101-2020_12_31_424971 259 37 . . . 10_1101-2020_12_31_424971 260 1 G37-state G37-state NNP 10_1101-2020_12_31_424971 260 2 627 627 CD 10_1101-2020_12_31_424971 260 3 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 260 4 and and CC 10_1101-2020_12_31_424971 260 5 ProL ProL NNP 10_1101-2020_12_31_424971 260 6 were be VBD 10_1101-2020_12_31_424971 260 7 also also RB 10_1101-2020_12_31_424971 260 8 prepared prepare VBN 10_1101-2020_12_31_424971 260 9 by by IN 10_1101-2020_12_31_424971 260 10 in in IN 10_1101-2020_12_31_424971 260 11 vitro vitro FW 10_1101-2020_12_31_424971 260 12 transcription transcription NN 10_1101-2020_12_31_424971 260 13 . . . 10_1101-2020_12_31_424971 261 1 Each each DT 10_1101-2020_12_31_424971 261 2 primary primary JJ 10_1101-2020_12_31_424971 261 3 transcript transcript NN 10_1101-2020_12_31_424971 261 4 contained contain VBD 10_1101-2020_12_31_424971 261 5 a a DT 10_1101-2020_12_31_424971 261 6 628 628 CD 10_1101-2020_12_31_424971 261 7 ribozyme ribozyme NNS 10_1101-2020_12_31_424971 261 8 domain domain NN 10_1101-2020_12_31_424971 261 9 on on IN 10_1101-2020_12_31_424971 261 10 the the DT 10_1101-2020_12_31_424971 261 11 5'-side 5'-side NN 10_1101-2020_12_31_424971 261 12 of of IN 10_1101-2020_12_31_424971 261 13 the the DT 10_1101-2020_12_31_424971 261 14 tRNA trna NN 10_1101-2020_12_31_424971 261 15 sequence sequence NN 10_1101-2020_12_31_424971 261 16 , , , 10_1101-2020_12_31_424971 261 17 which which WDT 10_1101-2020_12_31_424971 261 18 self self NN 10_1101-2020_12_31_424971 261 19 - - HYPH 10_1101-2020_12_31_424971 261 20 cleaved cleave VBN 10_1101-2020_12_31_424971 261 21 to to TO 10_1101-2020_12_31_424971 261 22 release release VB 10_1101-2020_12_31_424971 261 23 the the DT 10_1101-2020_12_31_424971 261 24 tRNA trna NN 10_1101-2020_12_31_424971 261 25 . . . 10_1101-2020_12_31_424971 262 1 629 629 CD 10_1101-2020_12_31_424971 262 2 m1G37-state m1G37-state NNP 10_1101-2020_12_31_424971 262 3 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 262 4 and and CC 10_1101-2020_12_31_424971 262 5 ProL ProL NNP 10_1101-2020_12_31_424971 262 6 were be VBD 10_1101-2020_12_31_424971 262 7 prepared prepare VBN 10_1101-2020_12_31_424971 262 8 by by IN 10_1101-2020_12_31_424971 262 9 TrmD TrmD NNP 10_1101-2020_12_31_424971 262 10 - - HYPH 10_1101-2020_12_31_424971 262 11 catalyzed catalyze VBN 10_1101-2020_12_31_424971 262 12 and and CC 10_1101-2020_12_31_424971 262 13 S S NNP 10_1101-2020_12_31_424971 262 14 - - HYPH 10_1101-2020_12_31_424971 262 15 adenosyl adenosyl NNP 10_1101-2020_12_31_424971 262 16 methionine methionine NNP 10_1101-2020_12_31_424971 262 17 630 630 CD 10_1101-2020_12_31_424971 262 18 ( ( -LRB- 10_1101-2020_12_31_424971 262 19 AdoMet)-dependent adomet)-dependent IN 10_1101-2020_12_31_424971 262 20 methylation methylation NN 10_1101-2020_12_31_424971 262 21 of of IN 10_1101-2020_12_31_424971 262 22 each each DT 10_1101-2020_12_31_424971 262 23 G37-state G37-state NNP 10_1101-2020_12_31_424971 262 24 tRNA trna NN 10_1101-2020_12_31_424971 262 25 . . . 10_1101-2020_12_31_424971 263 1 Due due IN 10_1101-2020_12_31_424971 263 2 to to IN 10_1101-2020_12_31_424971 263 3 the the DT 10_1101-2020_12_31_424971 263 4 lability lability NN 10_1101-2020_12_31_424971 263 5 of of IN 10_1101-2020_12_31_424971 263 6 the the DT 10_1101-2020_12_31_424971 263 7 aminoacyl aminoacyl JJ 10_1101-2020_12_31_424971 263 8 631 631 CD 10_1101-2020_12_31_424971 263 9 linkage linkage NN 10_1101-2020_12_31_424971 263 10 to to IN 10_1101-2020_12_31_424971 263 11 Pro Pro NNP 10_1101-2020_12_31_424971 263 12 , , , 10_1101-2020_12_31_424971 263 13 stocks stock NNS 10_1101-2020_12_31_424971 263 14 of of IN 10_1101-2020_12_31_424971 263 15 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 263 16 and and CC 10_1101-2020_12_31_424971 263 17 ProL ProL NNP 10_1101-2020_12_31_424971 263 18 aminoacylated aminoacylate VBD 10_1101-2020_12_31_424971 263 19 with with IN 10_1101-2020_12_31_424971 263 20 Pro Pro NNP 10_1101-2020_12_31_424971 263 21 were be VBD 10_1101-2020_12_31_424971 263 22 either either CC 10_1101-2020_12_31_424971 263 23 used use VBN 10_1101-2020_12_31_424971 263 24 immediately immediately RB 10_1101-2020_12_31_424971 263 25 632 632 CD 10_1101-2020_12_31_424971 263 26 or or CC 10_1101-2020_12_31_424971 263 27 stored store VBN 10_1101-2020_12_31_424971 263 28 no no RB 10_1101-2020_12_31_424971 263 29 longer long RBR 10_1101-2020_12_31_424971 263 30 than than IN 10_1101-2020_12_31_424971 263 31 2 2 CD 10_1101-2020_12_31_424971 263 32 - - SYM 10_1101-2020_12_31_424971 263 33 3 3 CD 10_1101-2020_12_31_424971 263 34 weeks week NNS 10_1101-2020_12_31_424971 263 35 at at IN 10_1101-2020_12_31_424971 263 36 – – : 10_1101-2020_12_31_424971 263 37 70 70 CD 10_1101-2020_12_31_424971 263 38 ° ° , 10_1101-2020_12_31_424971 263 39 C C NNP 10_1101-2020_12_31_424971 263 40 in in IN 10_1101-2020_12_31_424971 263 41 25 25 CD 10_1101-2020_12_31_424971 263 42 mM mM NNP 10_1101-2020_12_31_424971 263 43 NaOAc NaOAc NNS 10_1101-2020_12_31_424971 263 44 ( ( -LRB- 10_1101-2020_12_31_424971 263 45 pH pH NNP 10_1101-2020_12_31_424971 263 46 5.0 5.0 CD 10_1101-2020_12_31_424971 263 47 ) ) -RRB- 10_1101-2020_12_31_424971 263 48 . . . 10_1101-2020_12_31_424971 264 1 633 633 CD 10_1101-2020_12_31_424971 264 2 634 634 CD 10_1101-2020_12_31_424971 264 3 Primer Primer NNP 10_1101-2020_12_31_424971 264 4 extension extension NN 10_1101-2020_12_31_424971 264 5 inhibition inhibition NN 10_1101-2020_12_31_424971 264 6 assays assays RB 10_1101-2020_12_31_424971 264 7 . . . 10_1101-2020_12_31_424971 265 1 Primer primer JJ 10_1101-2020_12_31_424971 265 2 extension extension NN 10_1101-2020_12_31_424971 265 3 inhibition inhibition NN 10_1101-2020_12_31_424971 265 4 analyses analysis NNS 10_1101-2020_12_31_424971 265 5 of of IN 10_1101-2020_12_31_424971 265 6 native- native- NN 10_1101-2020_12_31_424971 265 7 , , , 10_1101-2020_12_31_424971 265 8 G37- G37- NNP 10_1101-2020_12_31_424971 265 9 , , , 10_1101-2020_12_31_424971 265 10 and and CC 10_1101-2020_12_31_424971 265 11 635 635 CD 10_1101-2020_12_31_424971 265 12 m1G37-state m1G37-state NNP 10_1101-2020_12_31_424971 265 13 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 265 14 and and CC 10_1101-2020_12_31_424971 265 15 ProL ProL NNP 10_1101-2020_12_31_424971 265 16 were be VBD 10_1101-2020_12_31_424971 265 17 performed perform VBN 10_1101-2020_12_31_424971 265 18 as as IN 10_1101-2020_12_31_424971 265 19 described30 described30 JJ 10_1101-2020_12_31_424971 265 20 . . . 10_1101-2020_12_31_424971 266 1 A a DT 10_1101-2020_12_31_424971 266 2 DNA dna NN 10_1101-2020_12_31_424971 266 3 primer primer NN 10_1101-2020_12_31_424971 266 4 complementary complementary JJ 10_1101-2020_12_31_424971 266 5 to to IN 10_1101-2020_12_31_424971 266 6 636 636 CD 10_1101-2020_12_31_424971 266 7 the the DT 10_1101-2020_12_31_424971 266 8 sequence sequence NN 10_1101-2020_12_31_424971 266 9 of of IN 10_1101-2020_12_31_424971 266 10 C41 C41 NNP 10_1101-2020_12_31_424971 266 11 to to IN 10_1101-2020_12_31_424971 266 12 A57 A57 NNP 10_1101-2020_12_31_424971 266 13 of of IN 10_1101-2020_12_31_424971 266 14 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 266 15 and and CC 10_1101-2020_12_31_424971 266 16 ProL ProL NNP 10_1101-2020_12_31_424971 266 17 was be VBD 10_1101-2020_12_31_424971 266 18 chemically chemically RB 10_1101-2020_12_31_424971 266 19 synthesized synthesize VBN 10_1101-2020_12_31_424971 266 20 , , , 10_1101-2020_12_31_424971 266 21 32P 32P NNP 10_1101-2020_12_31_424971 266 22 - - HYPH 10_1101-2020_12_31_424971 266 23 labeled label VBN 10_1101-2020_12_31_424971 266 24 at at IN 10_1101-2020_12_31_424971 266 25 the the DT 10_1101-2020_12_31_424971 266 26 637 637 CD 10_1101-2020_12_31_424971 266 27 5'-end 5'-end CD 10_1101-2020_12_31_424971 266 28 by by IN 10_1101-2020_12_31_424971 266 29 T4 T4 NNP 10_1101-2020_12_31_424971 266 30 polynucleotide polynucleotide NN 10_1101-2020_12_31_424971 266 31 kinase kinase NNP 10_1101-2020_12_31_424971 266 32 , , , 10_1101-2020_12_31_424971 266 33 annealed anneal VBD 10_1101-2020_12_31_424971 266 34 to to IN 10_1101-2020_12_31_424971 266 35 each each DT 10_1101-2020_12_31_424971 266 36 tRNA trna NN 10_1101-2020_12_31_424971 266 37 , , , 10_1101-2020_12_31_424971 266 38 and and CC 10_1101-2020_12_31_424971 266 39 was be VBD 10_1101-2020_12_31_424971 266 40 extended extend VBN 10_1101-2020_12_31_424971 266 41 by by IN 10_1101-2020_12_31_424971 266 42 Superscript Superscript NNP 10_1101-2020_12_31_424971 266 43 III III NNP 10_1101-2020_12_31_424971 266 44 638 638 CD 10_1101-2020_12_31_424971 266 45 reverse reverse JJ 10_1101-2020_12_31_424971 266 46 transcriptase transcriptase NN 10_1101-2020_12_31_424971 266 47 ( ( -LRB- 10_1101-2020_12_31_424971 266 48 Invitrogen Invitrogen NNP 10_1101-2020_12_31_424971 266 49 ) ) -RRB- 10_1101-2020_12_31_424971 266 50 at at IN 10_1101-2020_12_31_424971 266 51 200 200 CD 10_1101-2020_12_31_424971 266 52 units/µL units/µL NNS 10_1101-2020_12_31_424971 266 53 with with IN 10_1101-2020_12_31_424971 266 54 6 6 CD 10_1101-2020_12_31_424971 266 55 µM µM NNP 10_1101-2020_12_31_424971 266 56 each each DT 10_1101-2020_12_31_424971 266 57 dNTP dntp NN 10_1101-2020_12_31_424971 266 58 in in IN 10_1101-2020_12_31_424971 266 59 50 50 CD 10_1101-2020_12_31_424971 266 60 mM mM NNP 10_1101-2020_12_31_424971 266 61 Tris Tris NNP 10_1101-2020_12_31_424971 266 62 - - HYPH 10_1101-2020_12_31_424971 266 63 HCl HCl NNP 10_1101-2020_12_31_424971 266 64 ( ( -LRB- 10_1101-2020_12_31_424971 266 65 pH pH NNP 10_1101-2020_12_31_424971 266 66 639 639 CD 10_1101-2020_12_31_424971 266 67 8.3 8.3 CD 10_1101-2020_12_31_424971 266 68 ) ) -RRB- 10_1101-2020_12_31_424971 266 69 , , , 10_1101-2020_12_31_424971 266 70 3 3 CD 10_1101-2020_12_31_424971 266 71 mM mM NNP 10_1101-2020_12_31_424971 266 72 MgCl2 mgcl2 NN 10_1101-2020_12_31_424971 266 73 , , , 10_1101-2020_12_31_424971 266 74 75 75 CD 10_1101-2020_12_31_424971 266 75 mM mM NNP 10_1101-2020_12_31_424971 266 76 KCl KCl NNS 10_1101-2020_12_31_424971 266 77 , , , 10_1101-2020_12_31_424971 266 78 and and CC 10_1101-2020_12_31_424971 266 79 1 1 CD 10_1101-2020_12_31_424971 266 80 mM mM NNP 10_1101-2020_12_31_424971 266 81 DTT DTT NNP 10_1101-2020_12_31_424971 266 82 at at IN 10_1101-2020_12_31_424971 266 83 55 55 CD 10_1101-2020_12_31_424971 266 84 ° ° NNS 10_1101-2020_12_31_424971 266 85 C C NNP 10_1101-2020_12_31_424971 266 86 for for IN 10_1101-2020_12_31_424971 266 87 30 30 CD 10_1101-2020_12_31_424971 266 88 min min NN 10_1101-2020_12_31_424971 266 89 , , , 10_1101-2020_12_31_424971 266 90 and and CC 10_1101-2020_12_31_424971 266 91 terminated terminate VBN 10_1101-2020_12_31_424971 266 92 by by IN 10_1101-2020_12_31_424971 266 93 heating heat VBG 10_1101-2020_12_31_424971 266 94 at at IN 10_1101-2020_12_31_424971 266 95 640 640 CD 10_1101-2020_12_31_424971 266 96 70 70 CD 10_1101-2020_12_31_424971 266 97 ° ° , 10_1101-2020_12_31_424971 266 98 C C NNP 10_1101-2020_12_31_424971 266 99 for for IN 10_1101-2020_12_31_424971 266 100 15 15 CD 10_1101-2020_12_31_424971 266 101 min min NN 10_1101-2020_12_31_424971 266 102 . . . 10_1101-2020_12_31_424971 267 1 Extension extension NN 10_1101-2020_12_31_424971 267 2 was be VBD 10_1101-2020_12_31_424971 267 3 quenched quench VBN 10_1101-2020_12_31_424971 267 4 with with IN 10_1101-2020_12_31_424971 267 5 10 10 CD 10_1101-2020_12_31_424971 267 6 mM mm NN 10_1101-2020_12_31_424971 267 7 EDTA EDTA NNS 10_1101-2020_12_31_424971 267 8 and and CC 10_1101-2020_12_31_424971 267 9 products product NNS 10_1101-2020_12_31_424971 267 10 of of IN 10_1101-2020_12_31_424971 267 11 extension extension NN 10_1101-2020_12_31_424971 267 12 were be VBD 10_1101-2020_12_31_424971 267 13 641 641 CD 10_1101-2020_12_31_424971 267 14 separated separate VBN 10_1101-2020_12_31_424971 267 15 by by IN 10_1101-2020_12_31_424971 267 16 12 12 CD 10_1101-2020_12_31_424971 267 17 % % NN 10_1101-2020_12_31_424971 267 18 denaturing denature VBG 10_1101-2020_12_31_424971 267 19 polyacrylamide polyacrylamide NN 10_1101-2020_12_31_424971 267 20 gel gel NN 10_1101-2020_12_31_424971 267 21 electrophoresis electrophoresis NN 10_1101-2020_12_31_424971 267 22 ( ( -LRB- 10_1101-2020_12_31_424971 267 23 PAGE/7 PAGE/7 NNP 10_1101-2020_12_31_424971 267 24 M M NNP 10_1101-2020_12_31_424971 267 25 urea urea NN 10_1101-2020_12_31_424971 267 26 ) ) -RRB- 10_1101-2020_12_31_424971 267 27 and and CC 10_1101-2020_12_31_424971 267 28 analyzed analyze VBD 10_1101-2020_12_31_424971 267 29 642 642 CD 10_1101-2020_12_31_424971 267 30 by by IN 10_1101-2020_12_31_424971 267 31 phosphorimaging phosphorimage VBG 10_1101-2020_12_31_424971 267 32 . . . 10_1101-2020_12_31_424971 268 1 In in IN 10_1101-2020_12_31_424971 268 2 these these DT 10_1101-2020_12_31_424971 268 3 assays assay NNS 10_1101-2020_12_31_424971 268 4 , , , 10_1101-2020_12_31_424971 268 5 the the DT 10_1101-2020_12_31_424971 268 6 length length NN 10_1101-2020_12_31_424971 268 7 of of IN 10_1101-2020_12_31_424971 268 8 the the DT 10_1101-2020_12_31_424971 268 9 read read VB 10_1101-2020_12_31_424971 268 10 - - HYPH 10_1101-2020_12_31_424971 268 11 through through RP 10_1101-2020_12_31_424971 268 12 cDNA cdna NN 10_1101-2020_12_31_424971 268 13 is be VBZ 10_1101-2020_12_31_424971 268 14 54 54 CD 10_1101-2020_12_31_424971 268 15 - - SYM 10_1101-2020_12_31_424971 268 16 55 55 CD 10_1101-2020_12_31_424971 268 17 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 268 18 , , , 10_1101-2020_12_31_424971 268 19 643 643 CD 10_1101-2020_12_31_424971 268 20 as as IN 10_1101-2020_12_31_424971 268 21 in in IN 10_1101-2020_12_31_424971 268 22 the the DT 10_1101-2020_12_31_424971 268 23 case case NN 10_1101-2020_12_31_424971 268 24 of of IN 10_1101-2020_12_31_424971 268 25 the the DT 10_1101-2020_12_31_424971 268 26 G37-state G37-state NNP 10_1101-2020_12_31_424971 268 27 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 268 28 and and CC 10_1101-2020_12_31_424971 268 29 ProL ProL NNP 10_1101-2020_12_31_424971 268 30 , , , 10_1101-2020_12_31_424971 268 31 whereas whereas IN 10_1101-2020_12_31_424971 268 32 the the DT 10_1101-2020_12_31_424971 268 33 length length NN 10_1101-2020_12_31_424971 268 34 of of IN 10_1101-2020_12_31_424971 268 35 the the DT 10_1101-2020_12_31_424971 268 36 primer primer NN 10_1101-2020_12_31_424971 268 37 - - HYPH 10_1101-2020_12_31_424971 268 38 extension extension NN 10_1101-2020_12_31_424971 268 39 644 644 CD 10_1101-2020_12_31_424971 268 40 inhibited inhibit VBN 10_1101-2020_12_31_424971 268 41 cDNA cdna NN 10_1101-2020_12_31_424971 268 42 products product NNS 10_1101-2020_12_31_424971 268 43 is be VBZ 10_1101-2020_12_31_424971 268 44 21 21 CD 10_1101-2020_12_31_424971 268 45 - - SYM 10_1101-2020_12_31_424971 268 46 22 22 CD 10_1101-2020_12_31_424971 268 47 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 268 48 , , , 10_1101-2020_12_31_424971 268 49 as as IN 10_1101-2020_12_31_424971 268 50 in in IN 10_1101-2020_12_31_424971 268 51 the the DT 10_1101-2020_12_31_424971 268 52 case case NN 10_1101-2020_12_31_424971 268 53 of of IN 10_1101-2020_12_31_424971 268 54 the the DT 10_1101-2020_12_31_424971 268 55 m1G37-state m1G37-state NNP 10_1101-2020_12_31_424971 268 56 and and CC 10_1101-2020_12_31_424971 268 57 native native JJ 10_1101-2020_12_31_424971 268 58 - - HYPH 10_1101-2020_12_31_424971 268 59 state state NN 10_1101-2020_12_31_424971 268 60 . . . 10_1101-2020_12_31_424971 269 1 645 645 CD 10_1101-2020_12_31_424971 269 2 646 646 CD 10_1101-2020_12_31_424971 269 3 RNase RNase NNP 10_1101-2020_12_31_424971 269 4 T1 t1 NN 10_1101-2020_12_31_424971 269 5 cleavage cleavage NN 10_1101-2020_12_31_424971 269 6 inhibition inhibition NN 10_1101-2020_12_31_424971 269 7 assays assays RB 10_1101-2020_12_31_424971 269 8 . . . 10_1101-2020_12_31_424971 270 1 RNase rnase UH 10_1101-2020_12_31_424971 270 2 T1 T1 NNP 10_1101-2020_12_31_424971 270 3 cleaves cleave NNS 10_1101-2020_12_31_424971 270 4 on on IN 10_1101-2020_12_31_424971 270 5 the the DT 10_1101-2020_12_31_424971 270 6 3'-side 3'-side NNP 10_1101-2020_12_31_424971 270 7 of of IN 10_1101-2020_12_31_424971 270 8 G G NNP 10_1101-2020_12_31_424971 270 9 , , , 10_1101-2020_12_31_424971 270 10 but but CC 10_1101-2020_12_31_424971 270 11 not not RB 10_1101-2020_12_31_424971 270 12 m1G. m1g. VB 10_1101-2020_12_31_424971 271 1 647 647 CD 10_1101-2020_12_31_424971 271 2 Cleavage cleavage NN 10_1101-2020_12_31_424971 271 3 of of IN 10_1101-2020_12_31_424971 271 4 tRNAs trnas CD 10_1101-2020_12_31_424971 271 5 was be VBD 10_1101-2020_12_31_424971 271 6 performed perform VBN 10_1101-2020_12_31_424971 271 7 as as IN 10_1101-2020_12_31_424971 271 8 previously previously RB 10_1101-2020_12_31_424971 271 9 described29 described29 NNP 10_1101-2020_12_31_424971 271 10 . . . 10_1101-2020_12_31_424971 272 1 Each each DT 10_1101-2020_12_31_424971 272 2 tRNA trna NN 10_1101-2020_12_31_424971 272 3 ( ( -LRB- 10_1101-2020_12_31_424971 272 4 1 1 CD 10_1101-2020_12_31_424971 272 5 µg µg IN 10_1101-2020_12_31_424971 272 6 ) ) -RRB- 10_1101-2020_12_31_424971 272 7 was be VBD 10_1101-2020_12_31_424971 272 8 3'-end 3'-end CD 10_1101-2020_12_31_424971 272 9 648 648 CD 10_1101-2020_12_31_424971 272 10 labeled label VBN 10_1101-2020_12_31_424971 272 11 using use VBG 10_1101-2020_12_31_424971 272 12 Bacillus Bacillus NNP 10_1101-2020_12_31_424971 272 13 stearothermophilus stearothermophilus NN 10_1101-2020_12_31_424971 272 14 CCA CCA NNP 10_1101-2020_12_31_424971 272 15 - - HYPH 10_1101-2020_12_31_424971 272 16 adding add VBG 10_1101-2020_12_31_424971 272 17 enzyme enzyme NNS 10_1101-2020_12_31_424971 272 18 ( ( -LRB- 10_1101-2020_12_31_424971 272 19 10 10 CD 10_1101-2020_12_31_424971 272 20 nM nM NNP 10_1101-2020_12_31_424971 272 21 ) ) -RRB- 10_1101-2020_12_31_424971 272 22 with with IN 10_1101-2020_12_31_424971 272 23 [ [ -LRB- 10_1101-2020_12_31_424971 272 24 α-32P]ATP α-32p]atp NN 10_1101-2020_12_31_424971 272 25 at at IN 10_1101-2020_12_31_424971 272 26 60 60 CD 10_1101-2020_12_31_424971 272 27 ° ° NNS 10_1101-2020_12_31_424971 272 28 C C NNP 10_1101-2020_12_31_424971 272 29 649 649 CD 10_1101-2020_12_31_424971 272 30 in in IN 10_1101-2020_12_31_424971 272 31 100 100 CD 10_1101-2020_12_31_424971 272 32 mM mm NN 10_1101-2020_12_31_424971 272 33 glycine glycine NN 10_1101-2020_12_31_424971 272 34 ( ( -LRB- 10_1101-2020_12_31_424971 272 35 pH pH NNP 10_1101-2020_12_31_424971 272 36 9.0 9.0 CD 10_1101-2020_12_31_424971 272 37 ) ) -RRB- 10_1101-2020_12_31_424971 272 38 and and CC 10_1101-2020_12_31_424971 272 39 10 10 CD 10_1101-2020_12_31_424971 272 40 mM mM NNP 10_1101-2020_12_31_424971 272 41 MgCl2 mgcl2 NN 10_1101-2020_12_31_424971 272 42 . . . 10_1101-2020_12_31_424971 273 1 The the DT 10_1101-2020_12_31_424971 273 2 labeled label VBN 10_1101-2020_12_31_424971 273 3 tRNA trna NN 10_1101-2020_12_31_424971 273 4 was be VBD 10_1101-2020_12_31_424971 273 5 digested digest VBN 10_1101-2020_12_31_424971 273 6 by by IN 10_1101-2020_12_31_424971 273 7 RNase RNase NNP 10_1101-2020_12_31_424971 273 8 T1 T1 NNP 10_1101-2020_12_31_424971 273 9 650 650 CD 10_1101-2020_12_31_424971 273 10 ( ( -LRB- 10_1101-2020_12_31_424971 273 11 Roche Roche NNP 10_1101-2020_12_31_424971 273 12 , , , 10_1101-2020_12_31_424971 273 13 cat cat NN 10_1101-2020_12_31_424971 273 14 # # $ 10_1101-2020_12_31_424971 273 15 109193 109193 CD 10_1101-2020_12_31_424971 273 16 ) ) -RRB- 10_1101-2020_12_31_424971 273 17 at at IN 10_1101-2020_12_31_424971 273 18 a a DT 10_1101-2020_12_31_424971 273 19 final final JJ 10_1101-2020_12_31_424971 273 20 concentration concentration NN 10_1101-2020_12_31_424971 273 21 of of IN 10_1101-2020_12_31_424971 273 22 0.02 0.02 CD 10_1101-2020_12_31_424971 273 23 units/µL units/µL NNS 10_1101-2020_12_31_424971 273 24 for for IN 10_1101-2020_12_31_424971 273 25 20 20 CD 10_1101-2020_12_31_424971 273 26 min min NN 10_1101-2020_12_31_424971 273 27 at at IN 10_1101-2020_12_31_424971 273 28 50 50 CD 10_1101-2020_12_31_424971 273 29 ° ° NNS 10_1101-2020_12_31_424971 273 30 C C NNP 10_1101-2020_12_31_424971 273 31 in in IN 10_1101-2020_12_31_424971 273 32 20 20 CD 10_1101-2020_12_31_424971 273 33 mM mm NN 10_1101-2020_12_31_424971 273 34 651 651 CD 10_1101-2020_12_31_424971 273 35 .CC .CC , 10_1101-2020_12_31_424971 273 36 - - HYPH 10_1101-2020_12_31_424971 273 37 BY by IN 10_1101-2020_12_31_424971 273 38 - - HYPH 10_1101-2020_12_31_424971 273 39 NC NC NNP 10_1101-2020_12_31_424971 273 40 - - HYPH 10_1101-2020_12_31_424971 273 41 ND ND NNP 10_1101-2020_12_31_424971 273 42 4.0 4.0 CD 10_1101-2020_12_31_424971 273 43 International International NNP 10_1101-2020_12_31_424971 273 44 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 273 45 under under IN 10_1101-2020_12_31_424971 273 46 a a DT 10_1101-2020_12_31_424971 273 47 ( ( -LRB- 10_1101-2020_12_31_424971 273 48 which which WDT 10_1101-2020_12_31_424971 273 49 was be VBD 10_1101-2020_12_31_424971 273 50 not not RB 10_1101-2020_12_31_424971 273 51 certified certify VBN 10_1101-2020_12_31_424971 273 52 by by IN 10_1101-2020_12_31_424971 273 53 peer peer NN 10_1101-2020_12_31_424971 273 54 review review NN 10_1101-2020_12_31_424971 273 55 ) ) -RRB- 10_1101-2020_12_31_424971 273 56 is be VBZ 10_1101-2020_12_31_424971 273 57 the the DT 10_1101-2020_12_31_424971 273 58 author author NN 10_1101-2020_12_31_424971 273 59 / / SYM 10_1101-2020_12_31_424971 273 60 funder funder NN 10_1101-2020_12_31_424971 273 61 , , , 10_1101-2020_12_31_424971 273 62 who who WP 10_1101-2020_12_31_424971 273 63 has have VBZ 10_1101-2020_12_31_424971 273 64 granted grant VBN 10_1101-2020_12_31_424971 273 65 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 273 66 a a DT 10_1101-2020_12_31_424971 273 67 license license NN 10_1101-2020_12_31_424971 273 68 to to TO 10_1101-2020_12_31_424971 273 69 display display VB 10_1101-2020_12_31_424971 273 70 the the DT 10_1101-2020_12_31_424971 273 71 preprint preprint NN 10_1101-2020_12_31_424971 273 72 in in IN 10_1101-2020_12_31_424971 273 73 perpetuity perpetuity NN 10_1101-2020_12_31_424971 273 74 . . . 10_1101-2020_12_31_424971 274 1 It -PRON- PRP 10_1101-2020_12_31_424971 274 2 is be VBZ 10_1101-2020_12_31_424971 274 3 made make VBN 10_1101-2020_12_31_424971 274 4 The the DT 10_1101-2020_12_31_424971 274 5 copyright copyright NN 10_1101-2020_12_31_424971 274 6 holder holder NN 10_1101-2020_12_31_424971 274 7 for for IN 10_1101-2020_12_31_424971 274 8 this this DT 10_1101-2020_12_31_424971 274 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 274 10 version version NN 10_1101-2020_12_31_424971 274 11 posted post VBD 10_1101-2020_12_31_424971 274 12 January January NNP 10_1101-2020_12_31_424971 274 13 2 2 CD 10_1101-2020_12_31_424971 274 14 , , , 10_1101-2020_12_31_424971 274 15 2021 2021 CD 10_1101-2020_12_31_424971 274 16 . . . 10_1101-2020_12_31_424971 274 17 ; ; : 10_1101-2020_12_31_424971 274 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 274 19 : : : 10_1101-2020_12_31_424971 274 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 274 21 preprint preprint NN 10_1101-2020_12_31_424971 274 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 274 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 274 24 27 27 CD 10_1101-2020_12_31_424971 274 25 sodium sodium NN 10_1101-2020_12_31_424971 274 26 citrate citrate NN 10_1101-2020_12_31_424971 274 27 ( ( -LRB- 10_1101-2020_12_31_424971 274 28 pH pH NNP 10_1101-2020_12_31_424971 274 29 5.5 5.5 CD 10_1101-2020_12_31_424971 274 30 ) ) -RRB- 10_1101-2020_12_31_424971 274 31 and and CC 10_1101-2020_12_31_424971 274 32 1 1 CD 10_1101-2020_12_31_424971 274 33 mM mM NNP 10_1101-2020_12_31_424971 274 34 ethylene ethylene NN 10_1101-2020_12_31_424971 274 35 diamine diamine NN 10_1101-2020_12_31_424971 274 36 tetraacetic tetraacetic JJ 10_1101-2020_12_31_424971 274 37 acid acid NN 10_1101-2020_12_31_424971 274 38 ( ( -LRB- 10_1101-2020_12_31_424971 274 39 EDTA EDTA NNP 10_1101-2020_12_31_424971 274 40 ) ) -RRB- 10_1101-2020_12_31_424971 274 41 . . . 10_1101-2020_12_31_424971 275 1 The the DT 10_1101-2020_12_31_424971 275 2 RNA RNA NNP 10_1101-2020_12_31_424971 275 3 fragments fragment VBZ 10_1101-2020_12_31_424971 275 4 652 652 CD 10_1101-2020_12_31_424971 275 5 generated generate VBN 10_1101-2020_12_31_424971 275 6 from from IN 10_1101-2020_12_31_424971 275 7 cleavage cleavage NN 10_1101-2020_12_31_424971 275 8 were be VBD 10_1101-2020_12_31_424971 275 9 separated separate VBN 10_1101-2020_12_31_424971 275 10 by by IN 10_1101-2020_12_31_424971 275 11 12 12 CD 10_1101-2020_12_31_424971 275 12 % % NN 10_1101-2020_12_31_424971 275 13 PAGE/7 PAGE/7 NNP 10_1101-2020_12_31_424971 275 14 M M NNP 10_1101-2020_12_31_424971 275 15 urea urea NN 10_1101-2020_12_31_424971 275 16 along along IN 10_1101-2020_12_31_424971 275 17 with with IN 10_1101-2020_12_31_424971 275 18 an an DT 10_1101-2020_12_31_424971 275 19 RNA RNA NNP 10_1101-2020_12_31_424971 275 20 ladder ladder NN 10_1101-2020_12_31_424971 275 21 653 653 CD 10_1101-2020_12_31_424971 275 22 generated generate VBN 10_1101-2020_12_31_424971 275 23 by by IN 10_1101-2020_12_31_424971 275 24 alkali alkali JJ 10_1101-2020_12_31_424971 275 25 hydrolysis hydrolysis NN 10_1101-2020_12_31_424971 275 26 of of IN 10_1101-2020_12_31_424971 275 27 the the DT 10_1101-2020_12_31_424971 275 28 tRNA trna NN 10_1101-2020_12_31_424971 275 29 of of IN 10_1101-2020_12_31_424971 275 30 interest interest NN 10_1101-2020_12_31_424971 275 31 . . . 10_1101-2020_12_31_424971 276 1 Cleavage cleavage NN 10_1101-2020_12_31_424971 276 2 was be VBD 10_1101-2020_12_31_424971 276 3 analyzed analyze VBN 10_1101-2020_12_31_424971 276 4 by by IN 10_1101-2020_12_31_424971 276 5 654 654 CD 10_1101-2020_12_31_424971 276 6 phosphorimaging phosphorimaging NN 10_1101-2020_12_31_424971 276 7 . . . 10_1101-2020_12_31_424971 277 1 655 655 CD 10_1101-2020_12_31_424971 277 2 656 656 CD 10_1101-2020_12_31_424971 277 3 Methylation methylation NN 10_1101-2020_12_31_424971 277 4 assays assay NNS 10_1101-2020_12_31_424971 277 5 . . . 10_1101-2020_12_31_424971 278 1 Pre pre JJ 10_1101-2020_12_31_424971 278 2 - - JJ 10_1101-2020_12_31_424971 278 3 steady steady JJ 10_1101-2020_12_31_424971 278 4 - - HYPH 10_1101-2020_12_31_424971 278 5 state state NN 10_1101-2020_12_31_424971 278 6 assays assay NNS 10_1101-2020_12_31_424971 278 7 under under IN 10_1101-2020_12_31_424971 278 8 single single JJ 10_1101-2020_12_31_424971 278 9 - - HYPH 10_1101-2020_12_31_424971 278 10 turnover turnover NN 10_1101-2020_12_31_424971 278 11 conditions61 conditions61 NN 10_1101-2020_12_31_424971 278 12 were be VBD 10_1101-2020_12_31_424971 278 13 performed perform VBN 10_1101-2020_12_31_424971 278 14 657 657 CD 10_1101-2020_12_31_424971 278 15 on on IN 10_1101-2020_12_31_424971 278 16 a a DT 10_1101-2020_12_31_424971 278 17 rapid rapid JJ 10_1101-2020_12_31_424971 278 18 quench quench NN 10_1101-2020_12_31_424971 278 19 - - HYPH 10_1101-2020_12_31_424971 278 20 flow flow NN 10_1101-2020_12_31_424971 278 21 apparatus apparatus NN 10_1101-2020_12_31_424971 278 22 ( ( -LRB- 10_1101-2020_12_31_424971 278 23 Kintek Kintek NNP 10_1101-2020_12_31_424971 278 24 RQF-3 RQF-3 NNP 10_1101-2020_12_31_424971 278 25 ) ) -RRB- 10_1101-2020_12_31_424971 278 26 . . . 10_1101-2020_12_31_424971 279 1 The the DT 10_1101-2020_12_31_424971 279 2 tRNA trna NN 10_1101-2020_12_31_424971 279 3 substrate substrate NN 10_1101-2020_12_31_424971 279 4 was be VBD 10_1101-2020_12_31_424971 279 5 heated heat VBN 10_1101-2020_12_31_424971 279 6 to to IN 10_1101-2020_12_31_424971 279 7 85 85 CD 10_1101-2020_12_31_424971 279 8 ° ° , 10_1101-2020_12_31_424971 279 9 C C NNP 10_1101-2020_12_31_424971 279 10 for for IN 10_1101-2020_12_31_424971 279 11 658 658 CD 10_1101-2020_12_31_424971 279 12 2.5 2.5 CD 10_1101-2020_12_31_424971 279 13 min min NN 10_1101-2020_12_31_424971 279 14 followed follow VBN 10_1101-2020_12_31_424971 279 15 by by IN 10_1101-2020_12_31_424971 279 16 addition addition NN 10_1101-2020_12_31_424971 279 17 of of IN 10_1101-2020_12_31_424971 279 18 10 10 CD 10_1101-2020_12_31_424971 279 19 mM mM NNP 10_1101-2020_12_31_424971 279 20 MgCl2 mgcl2 NN 10_1101-2020_12_31_424971 279 21 , , , 10_1101-2020_12_31_424971 279 22 and and CC 10_1101-2020_12_31_424971 279 23 slowly slowly RB 10_1101-2020_12_31_424971 279 24 cooled cool VBD 10_1101-2020_12_31_424971 279 25 to to IN 10_1101-2020_12_31_424971 279 26 37 37 CD 10_1101-2020_12_31_424971 279 27 ° ° , 10_1101-2020_12_31_424971 279 28 C C NNP 10_1101-2020_12_31_424971 279 29 in in IN 10_1101-2020_12_31_424971 279 30 15 15 CD 10_1101-2020_12_31_424971 279 31 min min NN 10_1101-2020_12_31_424971 279 32 . . . 10_1101-2020_12_31_424971 280 1 N1-methylation n1-methylation DT 10_1101-2020_12_31_424971 280 2 659 659 CD 10_1101-2020_12_31_424971 280 3 of of IN 10_1101-2020_12_31_424971 280 4 G37 G37 NNP 10_1101-2020_12_31_424971 280 5 in in IN 10_1101-2020_12_31_424971 280 6 the the DT 10_1101-2020_12_31_424971 280 7 pre pre NN 10_1101-2020_12_31_424971 280 8 - - JJ 10_1101-2020_12_31_424971 280 9 annealed anneal VBN 10_1101-2020_12_31_424971 280 10 tRNA tRNA NNP 10_1101-2020_12_31_424971 280 11 ( ( -LRB- 10_1101-2020_12_31_424971 280 12 final final JJ 10_1101-2020_12_31_424971 280 13 concentration concentration NNP 10_1101-2020_12_31_424971 280 14 1 1 CD 10_1101-2020_12_31_424971 280 15 µM µM NNP 10_1101-2020_12_31_424971 280 16 ) ) -RRB- 10_1101-2020_12_31_424971 280 17 was be VBD 10_1101-2020_12_31_424971 280 18 initiated initiate VBN 10_1101-2020_12_31_424971 280 19 with with IN 10_1101-2020_12_31_424971 280 20 the the DT 10_1101-2020_12_31_424971 280 21 addition addition NN 10_1101-2020_12_31_424971 280 22 of of IN 10_1101-2020_12_31_424971 280 23 E. E. NNP 10_1101-2020_12_31_424971 280 24 660 660 CD 10_1101-2020_12_31_424971 280 25 coli coli NNS 10_1101-2020_12_31_424971 280 26 TrmD TrmD NNP 10_1101-2020_12_31_424971 280 27 ( ( -LRB- 10_1101-2020_12_31_424971 280 28 10 10 CD 10_1101-2020_12_31_424971 280 29 µM µM NNP 10_1101-2020_12_31_424971 280 30 ) ) -RRB- 10_1101-2020_12_31_424971 280 31 and and CC 10_1101-2020_12_31_424971 280 32 [ [ -LRB- 10_1101-2020_12_31_424971 280 33 3H]-AdoMet 3h]-adomet CD 10_1101-2020_12_31_424971 280 34 ( ( -LRB- 10_1101-2020_12_31_424971 280 35 Perkin Perkin NNP 10_1101-2020_12_31_424971 280 36 Elmer Elmer NNP 10_1101-2020_12_31_424971 280 37 , , , 10_1101-2020_12_31_424971 280 38 4200 4200 CD 10_1101-2020_12_31_424971 280 39 DPM DPM NNP 10_1101-2020_12_31_424971 280 40 / / SYM 10_1101-2020_12_31_424971 280 41 pmol pmol NN 10_1101-2020_12_31_424971 280 42 ) ) -RRB- 10_1101-2020_12_31_424971 280 43 at at IN 10_1101-2020_12_31_424971 280 44 a a DT 10_1101-2020_12_31_424971 280 45 final final JJ 10_1101-2020_12_31_424971 280 46 concentration concentration NN 10_1101-2020_12_31_424971 280 47 of of IN 10_1101-2020_12_31_424971 280 48 15 15 CD 10_1101-2020_12_31_424971 280 49 661 661 CD 10_1101-2020_12_31_424971 280 50 µM µM NNP 10_1101-2020_12_31_424971 280 51 in in IN 10_1101-2020_12_31_424971 280 52 a a DT 10_1101-2020_12_31_424971 280 53 buffer buffer NN 10_1101-2020_12_31_424971 280 54 containing contain VBG 10_1101-2020_12_31_424971 280 55 100 100 CD 10_1101-2020_12_31_424971 280 56 mM mM NNP 10_1101-2020_12_31_424971 280 57 Tris Tris NNP 10_1101-2020_12_31_424971 280 58 - - HYPH 10_1101-2020_12_31_424971 280 59 HCl HCl NNP 10_1101-2020_12_31_424971 280 60 ( ( -LRB- 10_1101-2020_12_31_424971 280 61 pH pH NNP 10_1101-2020_12_31_424971 280 62 8.0 8.0 CD 10_1101-2020_12_31_424971 280 63 ) ) -RRB- 10_1101-2020_12_31_424971 280 64 , , , 10_1101-2020_12_31_424971 280 65 24 24 CD 10_1101-2020_12_31_424971 280 66 mM mm NN 10_1101-2020_12_31_424971 280 67 NH4Cl nh4cl CD 10_1101-2020_12_31_424971 280 68 , , , 10_1101-2020_12_31_424971 280 69 6 6 CD 10_1101-2020_12_31_424971 280 70 mM mM NNP 10_1101-2020_12_31_424971 280 71 MgCl2 mgcl2 NN 10_1101-2020_12_31_424971 280 72 , , , 10_1101-2020_12_31_424971 280 73 4 4 CD 10_1101-2020_12_31_424971 280 74 mM mM NNP 10_1101-2020_12_31_424971 280 75 DTT DTT NNP 10_1101-2020_12_31_424971 280 76 , , , 10_1101-2020_12_31_424971 280 77 0.1 0.1 CD 10_1101-2020_12_31_424971 280 78 662 662 CD 10_1101-2020_12_31_424971 280 79 mM mm NN 10_1101-2020_12_31_424971 280 80 EDTA edta NN 10_1101-2020_12_31_424971 280 81 , , , 10_1101-2020_12_31_424971 280 82 and and CC 10_1101-2020_12_31_424971 280 83 0.024 0.024 CD 10_1101-2020_12_31_424971 280 84 mg mg NNP 10_1101-2020_12_31_424971 280 85 / / SYM 10_1101-2020_12_31_424971 280 86 mL mL NNP 10_1101-2020_12_31_424971 280 87 BSA BSA NNP 10_1101-2020_12_31_424971 280 88 in in IN 10_1101-2020_12_31_424971 280 89 a a DT 10_1101-2020_12_31_424971 280 90 reaction reaction NN 10_1101-2020_12_31_424971 280 91 of of IN 10_1101-2020_12_31_424971 280 92 30 30 CD 10_1101-2020_12_31_424971 280 93 µL. µl. NN 10_1101-2020_12_31_424971 281 1 The the DT 10_1101-2020_12_31_424971 281 2 buffer buffer NN 10_1101-2020_12_31_424971 281 3 used use VBN 10_1101-2020_12_31_424971 281 4 was be VBD 10_1101-2020_12_31_424971 281 5 optimized optimize VBN 10_1101-2020_12_31_424971 281 6 for for IN 10_1101-2020_12_31_424971 281 7 TrmD TrmD NNP 10_1101-2020_12_31_424971 281 8 663 663 CD 10_1101-2020_12_31_424971 281 9 in in IN 10_1101-2020_12_31_424971 281 10 order order NN 10_1101-2020_12_31_424971 281 11 to to TO 10_1101-2020_12_31_424971 281 12 evaluate evaluate VB 10_1101-2020_12_31_424971 281 13 its -PRON- PRP$ 10_1101-2020_12_31_424971 281 14 in in FW 10_1101-2020_12_31_424971 281 15 vitro vitro FW 10_1101-2020_12_31_424971 281 16 activity61 activity61 NNP 10_1101-2020_12_31_424971 281 17 . . . 10_1101-2020_12_31_424971 282 1 Reaction reaction NN 10_1101-2020_12_31_424971 282 2 aliquots aliquot NNS 10_1101-2020_12_31_424971 282 3 of of IN 10_1101-2020_12_31_424971 282 4 5 5 CD 10_1101-2020_12_31_424971 282 5 µL µl CD 10_1101-2020_12_31_424971 282 6 were be VBD 10_1101-2020_12_31_424971 282 7 removed remove VBN 10_1101-2020_12_31_424971 282 8 at at IN 10_1101-2020_12_31_424971 282 9 various various JJ 10_1101-2020_12_31_424971 282 10 time time NN 10_1101-2020_12_31_424971 282 11 664 664 CD 10_1101-2020_12_31_424971 282 12 points point NNS 10_1101-2020_12_31_424971 282 13 and and CC 10_1101-2020_12_31_424971 282 14 precipitated precipitate VBD 10_1101-2020_12_31_424971 282 15 in in IN 10_1101-2020_12_31_424971 282 16 5 5 CD 10_1101-2020_12_31_424971 282 17 % % NN 10_1101-2020_12_31_424971 282 18 ( ( -LRB- 10_1101-2020_12_31_424971 282 19 w w NNP 10_1101-2020_12_31_424971 282 20 / / SYM 10_1101-2020_12_31_424971 282 21 v v NNP 10_1101-2020_12_31_424971 282 22 ) ) -RRB- 10_1101-2020_12_31_424971 282 23 trichloroacetic trichloroacetic JJ 10_1101-2020_12_31_424971 282 24 acid acid NN 10_1101-2020_12_31_424971 282 25 ( ( -LRB- 10_1101-2020_12_31_424971 282 26 TCA TCA NNP 10_1101-2020_12_31_424971 282 27 ) ) -RRB- 10_1101-2020_12_31_424971 282 28 on on IN 10_1101-2020_12_31_424971 282 29 filter filter NN 10_1101-2020_12_31_424971 282 30 pads pad NNS 10_1101-2020_12_31_424971 282 31 for for IN 10_1101-2020_12_31_424971 282 32 10 10 CD 10_1101-2020_12_31_424971 282 33 min min NN 10_1101-2020_12_31_424971 282 34 twice twice RB 10_1101-2020_12_31_424971 282 35 . . . 10_1101-2020_12_31_424971 283 1 Filter Filter NNP 10_1101-2020_12_31_424971 283 2 665 665 CD 10_1101-2020_12_31_424971 283 3 pads pad NNS 10_1101-2020_12_31_424971 283 4 were be VBD 10_1101-2020_12_31_424971 283 5 washed wash VBN 10_1101-2020_12_31_424971 283 6 with with IN 10_1101-2020_12_31_424971 283 7 95 95 CD 10_1101-2020_12_31_424971 283 8 % % NN 10_1101-2020_12_31_424971 283 9 ethanol ethanol NN 10_1101-2020_12_31_424971 283 10 twice twice RB 10_1101-2020_12_31_424971 283 11 , , , 10_1101-2020_12_31_424971 283 12 with with IN 10_1101-2020_12_31_424971 283 13 ether ether NN 10_1101-2020_12_31_424971 283 14 once once RB 10_1101-2020_12_31_424971 283 15 , , , 10_1101-2020_12_31_424971 283 16 air air NN 10_1101-2020_12_31_424971 283 17 dried dry VBN 10_1101-2020_12_31_424971 283 18 , , , 10_1101-2020_12_31_424971 283 19 and and CC 10_1101-2020_12_31_424971 283 20 measured measure VBN 10_1101-2020_12_31_424971 283 21 for for IN 10_1101-2020_12_31_424971 283 22 666 666 CD 10_1101-2020_12_31_424971 283 23 radioactivity radioactivity NN 10_1101-2020_12_31_424971 283 24 in in IN 10_1101-2020_12_31_424971 283 25 an an DT 10_1101-2020_12_31_424971 283 26 LS6000 LS6000 NNP 10_1101-2020_12_31_424971 283 27 scintillation scintillation NN 10_1101-2020_12_31_424971 283 28 counter counter NN 10_1101-2020_12_31_424971 283 29 ( ( -LRB- 10_1101-2020_12_31_424971 283 30 Beckman Beckman NNP 10_1101-2020_12_31_424971 283 31 ) ) -RRB- 10_1101-2020_12_31_424971 283 32 . . . 10_1101-2020_12_31_424971 284 1 Counts count NNS 10_1101-2020_12_31_424971 284 2 were be VBD 10_1101-2020_12_31_424971 284 3 converted convert VBN 10_1101-2020_12_31_424971 284 4 to to IN 10_1101-2020_12_31_424971 284 5 pmoles pmole NNS 10_1101-2020_12_31_424971 284 6 667 667 CD 10_1101-2020_12_31_424971 284 7 using use VBG 10_1101-2020_12_31_424971 284 8 the the DT 10_1101-2020_12_31_424971 284 9 specific specific JJ 10_1101-2020_12_31_424971 284 10 activity activity NN 10_1101-2020_12_31_424971 284 11 of of IN 10_1101-2020_12_31_424971 284 12 the the DT 10_1101-2020_12_31_424971 284 13 [ [ -LRB- 10_1101-2020_12_31_424971 284 14 3H]-AdoMet 3h]-adomet CD 10_1101-2020_12_31_424971 284 15 after after IN 10_1101-2020_12_31_424971 284 16 correcting correct VBG 10_1101-2020_12_31_424971 284 17 for for IN 10_1101-2020_12_31_424971 284 18 the the DT 10_1101-2020_12_31_424971 284 19 signal signal NN 10_1101-2020_12_31_424971 284 20 quenching quenching NN 10_1101-2020_12_31_424971 284 21 by by IN 10_1101-2020_12_31_424971 284 22 filter filter NN 10_1101-2020_12_31_424971 284 23 668 668 CD 10_1101-2020_12_31_424971 284 24 pads pad NNS 10_1101-2020_12_31_424971 284 25 . . . 10_1101-2020_12_31_424971 285 1 In in IN 10_1101-2020_12_31_424971 285 2 these these DT 10_1101-2020_12_31_424971 285 3 assays assay NNS 10_1101-2020_12_31_424971 285 4 , , , 10_1101-2020_12_31_424971 285 5 a a DT 10_1101-2020_12_31_424971 285 6 negative negative JJ 10_1101-2020_12_31_424971 285 7 control control NN 10_1101-2020_12_31_424971 285 8 was be VBD 10_1101-2020_12_31_424971 285 9 always always RB 10_1101-2020_12_31_424971 285 10 included include VBN 10_1101-2020_12_31_424971 285 11 , , , 10_1101-2020_12_31_424971 285 12 in in IN 10_1101-2020_12_31_424971 285 13 which which WDT 10_1101-2020_12_31_424971 285 14 no no DT 10_1101-2020_12_31_424971 285 15 enzyme enzyme NNS 10_1101-2020_12_31_424971 285 16 was be VBD 10_1101-2020_12_31_424971 285 17 added add VBN 10_1101-2020_12_31_424971 285 18 669 669 CD 10_1101-2020_12_31_424971 285 19 to to IN 10_1101-2020_12_31_424971 285 20 the the DT 10_1101-2020_12_31_424971 285 21 reaction61 reaction61 NN 10_1101-2020_12_31_424971 285 22 , , , 10_1101-2020_12_31_424971 285 23 and and CC 10_1101-2020_12_31_424971 285 24 signal signal NN 10_1101-2020_12_31_424971 285 25 from from IN 10_1101-2020_12_31_424971 285 26 the the DT 10_1101-2020_12_31_424971 285 27 negative negative JJ 10_1101-2020_12_31_424971 285 28 control control NN 10_1101-2020_12_31_424971 285 29 was be VBD 10_1101-2020_12_31_424971 285 30 subtracted subtract VBN 10_1101-2020_12_31_424971 285 31 from from IN 10_1101-2020_12_31_424971 285 32 signal signal NN 10_1101-2020_12_31_424971 285 33 of of IN 10_1101-2020_12_31_424971 285 34 each each DT 10_1101-2020_12_31_424971 285 35 sample sample NN 10_1101-2020_12_31_424971 285 36 670 670 CD 10_1101-2020_12_31_424971 285 37 for for IN 10_1101-2020_12_31_424971 285 38 determining determine VBG 10_1101-2020_12_31_424971 285 39 the the DT 10_1101-2020_12_31_424971 285 40 level level NN 10_1101-2020_12_31_424971 285 41 of of IN 10_1101-2020_12_31_424971 285 42 methylation methylation NN 10_1101-2020_12_31_424971 285 43 . . . 10_1101-2020_12_31_424971 286 1 671 671 CD 10_1101-2020_12_31_424971 286 2 672 672 CD 10_1101-2020_12_31_424971 286 3 Aminoacylation Aminoacylation NNP 10_1101-2020_12_31_424971 286 4 assays assay NNS 10_1101-2020_12_31_424971 286 5 . . . 10_1101-2020_12_31_424971 287 1 Each each DT 10_1101-2020_12_31_424971 287 2 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 287 3 or or CC 10_1101-2020_12_31_424971 287 4 ProL ProL NNP 10_1101-2020_12_31_424971 287 5 tRNA trna NN 10_1101-2020_12_31_424971 287 6 was be VBD 10_1101-2020_12_31_424971 287 7 aminoacylated aminoacylate VBN 10_1101-2020_12_31_424971 287 8 with with IN 10_1101-2020_12_31_424971 287 9 Pro Pro NNP 10_1101-2020_12_31_424971 287 10 by by IN 10_1101-2020_12_31_424971 287 11 a a DT 10_1101-2020_12_31_424971 287 12 673 673 CD 10_1101-2020_12_31_424971 287 13 recombinant recombinant JJ 10_1101-2020_12_31_424971 287 14 E. E. NNP 10_1101-2020_12_31_424971 287 15 coli coli NNS 10_1101-2020_12_31_424971 287 16 ProRS ProRS NNP 10_1101-2020_12_31_424971 287 17 expressed express VBD 10_1101-2020_12_31_424971 287 18 from from IN 10_1101-2020_12_31_424971 287 19 the the DT 10_1101-2020_12_31_424971 287 20 plasmid plasmid NN 10_1101-2020_12_31_424971 287 21 pET22 pET22 NNS 10_1101-2020_12_31_424971 287 22 and and CC 10_1101-2020_12_31_424971 287 23 purified purify VBN 10_1101-2020_12_31_424971 287 24 from from IN 10_1101-2020_12_31_424971 287 25 E. E. NNP 10_1101-2020_12_31_424971 287 26 coli coli NNS 10_1101-2020_12_31_424971 287 27 BL21 BL21 NNP 10_1101-2020_12_31_424971 287 28 674 674 CD 10_1101-2020_12_31_424971 287 29 ( ( -LRB- 10_1101-2020_12_31_424971 287 30 DE3)62 DE3)62 NNP 10_1101-2020_12_31_424971 287 31 . . . 10_1101-2020_12_31_424971 288 1 Each each DT 10_1101-2020_12_31_424971 288 2 tRNA trna NN 10_1101-2020_12_31_424971 288 3 was be VBD 10_1101-2020_12_31_424971 288 4 heat heat NN 10_1101-2020_12_31_424971 288 5 - - HYPH 10_1101-2020_12_31_424971 288 6 denatured denature VBN 10_1101-2020_12_31_424971 288 7 at at IN 10_1101-2020_12_31_424971 288 8 80 80 CD 10_1101-2020_12_31_424971 288 9 ºC ºC NNP 10_1101-2020_12_31_424971 288 10 for for IN 10_1101-2020_12_31_424971 288 11 3 3 CD 10_1101-2020_12_31_424971 288 12 min min NN 10_1101-2020_12_31_424971 288 13 , , , 10_1101-2020_12_31_424971 288 14 and and CC 10_1101-2020_12_31_424971 288 15 re re NNS 10_1101-2020_12_31_424971 288 16 - - VBN 10_1101-2020_12_31_424971 288 17 annealed anneal VBD 10_1101-2020_12_31_424971 288 18 at at IN 10_1101-2020_12_31_424971 288 19 37 37 CD 10_1101-2020_12_31_424971 288 20 ºC ºC NNP 10_1101-2020_12_31_424971 288 21 for for IN 10_1101-2020_12_31_424971 288 22 15 15 CD 10_1101-2020_12_31_424971 288 23 min min NN 10_1101-2020_12_31_424971 288 24 . . . 10_1101-2020_12_31_424971 289 1 675 675 CD 10_1101-2020_12_31_424971 289 2 Aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 289 3 under under IN 10_1101-2020_12_31_424971 289 4 pre pre JJ 10_1101-2020_12_31_424971 289 5 - - JJ 10_1101-2020_12_31_424971 289 6 steady steady JJ 10_1101-2020_12_31_424971 289 7 state state NN 10_1101-2020_12_31_424971 289 8 conditions condition NNS 10_1101-2020_12_31_424971 289 9 was be VBD 10_1101-2020_12_31_424971 289 10 performed perform VBN 10_1101-2020_12_31_424971 289 11 at at IN 10_1101-2020_12_31_424971 289 12 37 37 CD 10_1101-2020_12_31_424971 289 13 ºC ºC NNP 10_1101-2020_12_31_424971 289 14 with with IN 10_1101-2020_12_31_424971 289 15 10 10 CD 10_1101-2020_12_31_424971 289 16 µM µM NNP 10_1101-2020_12_31_424971 289 17 tRNA trna NN 10_1101-2020_12_31_424971 289 18 , , , 10_1101-2020_12_31_424971 289 19 1 1 CD 10_1101-2020_12_31_424971 289 20 676 676 CD 10_1101-2020_12_31_424971 289 21 .CC .CC : 10_1101-2020_12_31_424971 289 22 - - HYPH 10_1101-2020_12_31_424971 289 23 BY by IN 10_1101-2020_12_31_424971 289 24 - - HYPH 10_1101-2020_12_31_424971 289 25 NC NC NNP 10_1101-2020_12_31_424971 289 26 - - HYPH 10_1101-2020_12_31_424971 289 27 ND ND NNP 10_1101-2020_12_31_424971 289 28 4.0 4.0 CD 10_1101-2020_12_31_424971 289 29 International International NNP 10_1101-2020_12_31_424971 289 30 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 289 31 under under IN 10_1101-2020_12_31_424971 289 32 a a DT 10_1101-2020_12_31_424971 289 33 ( ( -LRB- 10_1101-2020_12_31_424971 289 34 which which WDT 10_1101-2020_12_31_424971 289 35 was be VBD 10_1101-2020_12_31_424971 289 36 not not RB 10_1101-2020_12_31_424971 289 37 certified certify VBN 10_1101-2020_12_31_424971 289 38 by by IN 10_1101-2020_12_31_424971 289 39 peer peer NN 10_1101-2020_12_31_424971 289 40 review review NN 10_1101-2020_12_31_424971 289 41 ) ) -RRB- 10_1101-2020_12_31_424971 289 42 is be VBZ 10_1101-2020_12_31_424971 289 43 the the DT 10_1101-2020_12_31_424971 289 44 author author NN 10_1101-2020_12_31_424971 289 45 / / SYM 10_1101-2020_12_31_424971 289 46 funder funder NN 10_1101-2020_12_31_424971 289 47 , , , 10_1101-2020_12_31_424971 289 48 who who WP 10_1101-2020_12_31_424971 289 49 has have VBZ 10_1101-2020_12_31_424971 289 50 granted grant VBN 10_1101-2020_12_31_424971 289 51 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 289 52 a a DT 10_1101-2020_12_31_424971 289 53 license license NN 10_1101-2020_12_31_424971 289 54 to to TO 10_1101-2020_12_31_424971 289 55 display display VB 10_1101-2020_12_31_424971 289 56 the the DT 10_1101-2020_12_31_424971 289 57 preprint preprint NN 10_1101-2020_12_31_424971 289 58 in in IN 10_1101-2020_12_31_424971 289 59 perpetuity perpetuity NN 10_1101-2020_12_31_424971 289 60 . . . 10_1101-2020_12_31_424971 290 1 It -PRON- PRP 10_1101-2020_12_31_424971 290 2 is be VBZ 10_1101-2020_12_31_424971 290 3 made make VBN 10_1101-2020_12_31_424971 290 4 The the DT 10_1101-2020_12_31_424971 290 5 copyright copyright NN 10_1101-2020_12_31_424971 290 6 holder holder NN 10_1101-2020_12_31_424971 290 7 for for IN 10_1101-2020_12_31_424971 290 8 this this DT 10_1101-2020_12_31_424971 290 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 290 10 version version NN 10_1101-2020_12_31_424971 290 11 posted post VBD 10_1101-2020_12_31_424971 290 12 January January NNP 10_1101-2020_12_31_424971 290 13 2 2 CD 10_1101-2020_12_31_424971 290 14 , , , 10_1101-2020_12_31_424971 290 15 2021 2021 CD 10_1101-2020_12_31_424971 290 16 . . . 10_1101-2020_12_31_424971 290 17 ; ; : 10_1101-2020_12_31_424971 290 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 290 19 : : : 10_1101-2020_12_31_424971 290 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 290 21 preprint preprint NN 10_1101-2020_12_31_424971 290 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 290 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 290 24 28 28 CD 10_1101-2020_12_31_424971 290 25 µM µM NNP 10_1101-2020_12_31_424971 290 26 ProRS ProRS NNP 10_1101-2020_12_31_424971 290 27 , , , 10_1101-2020_12_31_424971 290 28 and and CC 10_1101-2020_12_31_424971 290 29 15 15 CD 10_1101-2020_12_31_424971 290 30 µM µM NNP 10_1101-2020_12_31_424971 290 31 [ [ -LRB- 10_1101-2020_12_31_424971 290 32 3H]-Pro 3H]-Pro NNP 10_1101-2020_12_31_424971 290 33 ( ( -LRB- 10_1101-2020_12_31_424971 290 34 Perkin Perkin NNP 10_1101-2020_12_31_424971 290 35 Elmer Elmer NNP 10_1101-2020_12_31_424971 290 36 , , , 10_1101-2020_12_31_424971 290 37 7.5 7.5 CD 10_1101-2020_12_31_424971 290 38 Ci Ci NNP 10_1101-2020_12_31_424971 290 39 / / SYM 10_1101-2020_12_31_424971 290 40 mmol mmol NN 10_1101-2020_12_31_424971 290 41 ) ) -RRB- 10_1101-2020_12_31_424971 290 42 in in IN 10_1101-2020_12_31_424971 290 43 a a DT 10_1101-2020_12_31_424971 290 44 buffer buffer NN 10_1101-2020_12_31_424971 290 45 containing contain VBG 10_1101-2020_12_31_424971 290 46 20 20 CD 10_1101-2020_12_31_424971 290 47 mM mm NN 10_1101-2020_12_31_424971 290 48 KCl KCl NNS 10_1101-2020_12_31_424971 290 49 , , , 10_1101-2020_12_31_424971 290 50 10 10 CD 10_1101-2020_12_31_424971 290 51 677 677 CD 10_1101-2020_12_31_424971 290 52 mM mM NNP 10_1101-2020_12_31_424971 290 53 MgCl2 mgcl2 NN 10_1101-2020_12_31_424971 290 54 , , , 10_1101-2020_12_31_424971 290 55 4 4 CD 10_1101-2020_12_31_424971 290 56 mM mM NNP 10_1101-2020_12_31_424971 290 57 dithiothreitol dithiothreitol NN 10_1101-2020_12_31_424971 290 58 ( ( -LRB- 10_1101-2020_12_31_424971 290 59 DTT DTT NNP 10_1101-2020_12_31_424971 290 60 ) ) -RRB- 10_1101-2020_12_31_424971 290 61 , , , 10_1101-2020_12_31_424971 290 62 0.2 0.2 CD 10_1101-2020_12_31_424971 290 63 mg mg NNP 10_1101-2020_12_31_424971 290 64 / / SYM 10_1101-2020_12_31_424971 290 65 mL mL NNP 10_1101-2020_12_31_424971 290 66 bovine bovine NN 10_1101-2020_12_31_424971 290 67 serum serum NN 10_1101-2020_12_31_424971 290 68 albumin albumin NN 10_1101-2020_12_31_424971 290 69 ( ( -LRB- 10_1101-2020_12_31_424971 290 70 BSA BSA NNP 10_1101-2020_12_31_424971 290 71 ) ) -RRB- 10_1101-2020_12_31_424971 290 72 , , , 10_1101-2020_12_31_424971 290 73 2 2 CD 10_1101-2020_12_31_424971 290 74 mM mm NN 10_1101-2020_12_31_424971 290 75 ATP atp NN 10_1101-2020_12_31_424971 290 76 ( ( -LRB- 10_1101-2020_12_31_424971 290 77 pH pH NNP 10_1101-2020_12_31_424971 290 78 678 678 CD 10_1101-2020_12_31_424971 290 79 8.0 8.0 CD 10_1101-2020_12_31_424971 290 80 ) ) -RRB- 10_1101-2020_12_31_424971 290 81 , , , 10_1101-2020_12_31_424971 290 82 and and CC 10_1101-2020_12_31_424971 290 83 50 50 CD 10_1101-2020_12_31_424971 290 84 mM mM NNP 10_1101-2020_12_31_424971 290 85 Tris Tris NNP 10_1101-2020_12_31_424971 290 86 - - HYPH 10_1101-2020_12_31_424971 290 87 HCl HCl NNP 10_1101-2020_12_31_424971 290 88 ( ( -LRB- 10_1101-2020_12_31_424971 290 89 pH pH NNP 10_1101-2020_12_31_424971 290 90 7.5 7.5 CD 10_1101-2020_12_31_424971 290 91 ) ) -RRB- 10_1101-2020_12_31_424971 290 92 in in IN 10_1101-2020_12_31_424971 290 93 a a DT 10_1101-2020_12_31_424971 290 94 reaction reaction NN 10_1101-2020_12_31_424971 290 95 of of IN 10_1101-2020_12_31_424971 290 96 30 30 CD 10_1101-2020_12_31_424971 290 97 µL. µl. NN 10_1101-2020_12_31_424971 291 1 Reaction reaction NN 10_1101-2020_12_31_424971 291 2 aliquots aliquot NNS 10_1101-2020_12_31_424971 291 3 of of IN 10_1101-2020_12_31_424971 291 4 5 5 CD 10_1101-2020_12_31_424971 291 5 µL µl CD 10_1101-2020_12_31_424971 291 6 were be VBD 10_1101-2020_12_31_424971 291 7 removed remove VBN 10_1101-2020_12_31_424971 291 8 679 679 CD 10_1101-2020_12_31_424971 291 9 at at IN 10_1101-2020_12_31_424971 291 10 different different JJ 10_1101-2020_12_31_424971 291 11 time time NN 10_1101-2020_12_31_424971 291 12 intervals interval NNS 10_1101-2020_12_31_424971 291 13 and and CC 10_1101-2020_12_31_424971 291 14 precipitated precipitate VBD 10_1101-2020_12_31_424971 291 15 with with IN 10_1101-2020_12_31_424971 291 16 5 5 CD 10_1101-2020_12_31_424971 291 17 % % NN 10_1101-2020_12_31_424971 291 18 ( ( -LRB- 10_1101-2020_12_31_424971 291 19 w w NNP 10_1101-2020_12_31_424971 291 20 / / SYM 10_1101-2020_12_31_424971 291 21 v v LS 10_1101-2020_12_31_424971 291 22 ) ) -RRB- 10_1101-2020_12_31_424971 291 23 TCA TCA NNP 10_1101-2020_12_31_424971 291 24 on on IN 10_1101-2020_12_31_424971 291 25 filter filter NN 10_1101-2020_12_31_424971 291 26 pads pad NNS 10_1101-2020_12_31_424971 291 27 for for IN 10_1101-2020_12_31_424971 291 28 10 10 CD 10_1101-2020_12_31_424971 291 29 min min NN 10_1101-2020_12_31_424971 291 30 twice twice RB 10_1101-2020_12_31_424971 291 31 . . . 10_1101-2020_12_31_424971 292 1 Filter Filter NNP 10_1101-2020_12_31_424971 292 2 680 680 CD 10_1101-2020_12_31_424971 292 3 pads pad NNS 10_1101-2020_12_31_424971 292 4 were be VBD 10_1101-2020_12_31_424971 292 5 washed wash VBN 10_1101-2020_12_31_424971 292 6 with with IN 10_1101-2020_12_31_424971 292 7 95 95 CD 10_1101-2020_12_31_424971 292 8 % % NN 10_1101-2020_12_31_424971 292 9 ethanol ethanol NN 10_1101-2020_12_31_424971 292 10 twice twice RB 10_1101-2020_12_31_424971 292 11 , , , 10_1101-2020_12_31_424971 292 12 with with IN 10_1101-2020_12_31_424971 292 13 ether ether NN 10_1101-2020_12_31_424971 292 14 once once RB 10_1101-2020_12_31_424971 292 15 , , , 10_1101-2020_12_31_424971 292 16 air air NN 10_1101-2020_12_31_424971 292 17 dried dry VBN 10_1101-2020_12_31_424971 292 18 , , , 10_1101-2020_12_31_424971 292 19 and and CC 10_1101-2020_12_31_424971 292 20 measured measure VBN 10_1101-2020_12_31_424971 292 21 for for IN 10_1101-2020_12_31_424971 292 22 681 681 CD 10_1101-2020_12_31_424971 292 23 radioactivity radioactivity NN 10_1101-2020_12_31_424971 292 24 in in IN 10_1101-2020_12_31_424971 292 25 an an DT 10_1101-2020_12_31_424971 292 26 LS6000 LS6000 NNP 10_1101-2020_12_31_424971 292 27 scintillation scintillation NN 10_1101-2020_12_31_424971 292 28 counter counter NN 10_1101-2020_12_31_424971 292 29 ( ( -LRB- 10_1101-2020_12_31_424971 292 30 Beckman Beckman NNP 10_1101-2020_12_31_424971 292 31 ) ) -RRB- 10_1101-2020_12_31_424971 292 32 . . . 10_1101-2020_12_31_424971 293 1 Counts count NNS 10_1101-2020_12_31_424971 293 2 were be VBD 10_1101-2020_12_31_424971 293 3 converted convert VBN 10_1101-2020_12_31_424971 293 4 to to IN 10_1101-2020_12_31_424971 293 5 pmoles pmole NNS 10_1101-2020_12_31_424971 293 6 682 682 CD 10_1101-2020_12_31_424971 293 7 using use VBG 10_1101-2020_12_31_424971 293 8 the the DT 10_1101-2020_12_31_424971 293 9 specific specific JJ 10_1101-2020_12_31_424971 293 10 activity activity NN 10_1101-2020_12_31_424971 293 11 of of IN 10_1101-2020_12_31_424971 293 12 the the DT 10_1101-2020_12_31_424971 293 13 [ [ -LRB- 10_1101-2020_12_31_424971 293 14 3H]-Pro 3h]-pro NN 10_1101-2020_12_31_424971 293 15 after after IN 10_1101-2020_12_31_424971 293 16 correcting correct VBG 10_1101-2020_12_31_424971 293 17 for for IN 10_1101-2020_12_31_424971 293 18 signal signal NN 10_1101-2020_12_31_424971 293 19 quenching quenching NN 10_1101-2020_12_31_424971 293 20 by by IN 10_1101-2020_12_31_424971 293 21 filter filter NN 10_1101-2020_12_31_424971 293 22 pads pad NNS 10_1101-2020_12_31_424971 293 23 . . . 10_1101-2020_12_31_424971 294 1 683 683 CD 10_1101-2020_12_31_424971 294 2 684 684 CD 10_1101-2020_12_31_424971 294 3 Cell Cell NNP 10_1101-2020_12_31_424971 294 4 - - HYPH 10_1101-2020_12_31_424971 294 5 based base VBN 10_1101-2020_12_31_424971 294 6 +1-frameshifting +1-frameshifting NNP 10_1101-2020_12_31_424971 294 7 reporter reporter NN 10_1101-2020_12_31_424971 294 8 assays assays RB 10_1101-2020_12_31_424971 294 9 . . . 10_1101-2020_12_31_424971 295 1 Isogenic Isogenic NNP 10_1101-2020_12_31_424971 295 2 E. E. NNP 10_1101-2020_12_31_424971 295 3 coli coli VBZ 10_1101-2020_12_31_424971 295 4 strains strain NNS 10_1101-2020_12_31_424971 295 5 expressing express VBG 10_1101-2020_12_31_424971 295 6 chromosomal chromosomal JJ 10_1101-2020_12_31_424971 295 7 685 685 CD 10_1101-2020_12_31_424971 295 8 copies copy NNS 10_1101-2020_12_31_424971 295 9 of of IN 10_1101-2020_12_31_424971 295 10 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 295 11 or or CC 10_1101-2020_12_31_424971 295 12 ProL ProL NNP 10_1101-2020_12_31_424971 295 13 were be VBD 10_1101-2020_12_31_424971 295 14 created create VBN 10_1101-2020_12_31_424971 295 15 in in IN 10_1101-2020_12_31_424971 295 16 a a DT 10_1101-2020_12_31_424971 295 17 previously previously RB 10_1101-2020_12_31_424971 295 18 developed develop VBN 10_1101-2020_12_31_424971 295 19 trmD trmd NN 10_1101-2020_12_31_424971 295 20 - - HYPH 10_1101-2020_12_31_424971 295 21 knockdown knockdown VBN 10_1101-2020_12_31_424971 295 22 ( ( -LRB- 10_1101-2020_12_31_424971 295 23 trmD trmD NNP 10_1101-2020_12_31_424971 295 24 - - HYPH 10_1101-2020_12_31_424971 295 25 KD KD NNP 10_1101-2020_12_31_424971 295 26 ) ) -RRB- 10_1101-2020_12_31_424971 295 27 686 686 CD 10_1101-2020_12_31_424971 295 28 background background NN 10_1101-2020_12_31_424971 295 29 , , , 10_1101-2020_12_31_424971 295 30 in in IN 10_1101-2020_12_31_424971 295 31 which which WDT 10_1101-2020_12_31_424971 295 32 the the DT 10_1101-2020_12_31_424971 295 33 chromosomal chromosomal NN 10_1101-2020_12_31_424971 295 34 trmD trmd CD 10_1101-2020_12_31_424971 295 35 is be VBZ 10_1101-2020_12_31_424971 295 36 deleted delete VBN 10_1101-2020_12_31_424971 295 37 but but CC 10_1101-2020_12_31_424971 295 38 cell cell NN 10_1101-2020_12_31_424971 295 39 viability viability NN 10_1101-2020_12_31_424971 295 40 is be VBZ 10_1101-2020_12_31_424971 295 41 maintained maintain VBN 10_1101-2020_12_31_424971 295 42 through through IN 10_1101-2020_12_31_424971 295 43 the the DT 10_1101-2020_12_31_424971 295 44 687 687 CD 10_1101-2020_12_31_424971 295 45 arabinose arabinose RB 10_1101-2020_12_31_424971 295 46 - - HYPH 10_1101-2020_12_31_424971 295 47 induced induce VBN 10_1101-2020_12_31_424971 295 48 expression expression NN 10_1101-2020_12_31_424971 295 49 of of IN 10_1101-2020_12_31_424971 295 50 a a DT 10_1101-2020_12_31_424971 295 51 plasmid plasmid NN 10_1101-2020_12_31_424971 295 52 - - HYPH 10_1101-2020_12_31_424971 295 53 borne bear VBN 10_1101-2020_12_31_424971 295 54 trm5 trm5 NN 10_1101-2020_12_31_424971 295 55 , , , 10_1101-2020_12_31_424971 295 56 the the DT 10_1101-2020_12_31_424971 295 57 human human JJ 10_1101-2020_12_31_424971 295 58 counterpart counterpart NN 10_1101-2020_12_31_424971 295 59 of of IN 10_1101-2020_12_31_424971 295 60 trmD29,30 trmD29,30 NNP 10_1101-2020_12_31_424971 295 61 that that DT 10_1101-2020_12_31_424971 295 62 is be VBZ 10_1101-2020_12_31_424971 295 63 688 688 CD 10_1101-2020_12_31_424971 295 64 competent competent JJ 10_1101-2020_12_31_424971 295 65 for for IN 10_1101-2020_12_31_424971 295 66 m1G37 m1g37 JJ 10_1101-2020_12_31_424971 295 67 synthesis synthesis NN 10_1101-2020_12_31_424971 295 68 to to TO 10_1101-2020_12_31_424971 295 69 support support VB 10_1101-2020_12_31_424971 295 70 bacterial bacterial JJ 10_1101-2020_12_31_424971 295 71 growth growth NN 10_1101-2020_12_31_424971 295 72 ( ( -LRB- 10_1101-2020_12_31_424971 295 73 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 295 74 Table Table NNP 10_1101-2020_12_31_424971 295 75 1 1 CD 10_1101-2020_12_31_424971 295 76 ) ) -RRB- 10_1101-2020_12_31_424971 295 77 . . . 10_1101-2020_12_31_424971 296 1 Due due IN 10_1101-2020_12_31_424971 296 2 to to IN 10_1101-2020_12_31_424971 296 3 the the DT 10_1101-2020_12_31_424971 296 4 689 689 CD 10_1101-2020_12_31_424971 296 5 essentiality essentiality NN 10_1101-2020_12_31_424971 296 6 of of IN 10_1101-2020_12_31_424971 296 7 trmD trmd CD 10_1101-2020_12_31_424971 296 8 for for IN 10_1101-2020_12_31_424971 296 9 cell cell NN 10_1101-2020_12_31_424971 296 10 growth growth NN 10_1101-2020_12_31_424971 296 11 , , , 10_1101-2020_12_31_424971 296 12 a a DT 10_1101-2020_12_31_424971 296 13 simple simple JJ 10_1101-2020_12_31_424971 296 14 knock knock NN 10_1101-2020_12_31_424971 296 15 - - HYPH 10_1101-2020_12_31_424971 296 16 out out NN 10_1101-2020_12_31_424971 296 17 can can MD 10_1101-2020_12_31_424971 296 18 not not RB 10_1101-2020_12_31_424971 296 19 be be VB 10_1101-2020_12_31_424971 296 20 made make VBN 10_1101-2020_12_31_424971 296 21 . . . 10_1101-2020_12_31_424971 297 1 We -PRON- PRP 10_1101-2020_12_31_424971 297 2 chose choose VBD 10_1101-2020_12_31_424971 297 3 human human JJ 10_1101-2020_12_31_424971 297 4 Trm5 Trm5 NNP 10_1101-2020_12_31_424971 297 5 690 690 CD 10_1101-2020_12_31_424971 297 6 as as IN 10_1101-2020_12_31_424971 297 7 the the DT 10_1101-2020_12_31_424971 297 8 maintenance maintenance NN 10_1101-2020_12_31_424971 297 9 protein protein NN 10_1101-2020_12_31_424971 297 10 in in IN 10_1101-2020_12_31_424971 297 11 the the DT 10_1101-2020_12_31_424971 297 12 trmD trmd CD 10_1101-2020_12_31_424971 297 13 - - HYPH 10_1101-2020_12_31_424971 297 14 KD KD NNP 10_1101-2020_12_31_424971 297 15 background background NN 10_1101-2020_12_31_424971 297 16 , , , 10_1101-2020_12_31_424971 297 17 because because IN 10_1101-2020_12_31_424971 297 18 this this DT 10_1101-2020_12_31_424971 297 19 enzyme enzyme NNS 10_1101-2020_12_31_424971 297 20 is be VBZ 10_1101-2020_12_31_424971 297 21 rapidly rapidly RB 10_1101-2020_12_31_424971 297 22 degraded degrade VBN 10_1101-2020_12_31_424971 297 23 691 691 CD 10_1101-2020_12_31_424971 297 24 in in IN 10_1101-2020_12_31_424971 297 25 E. E. NNP 10_1101-2020_12_31_424971 297 26 coli coli NNS 10_1101-2020_12_31_424971 297 27 once once IN 10_1101-2020_12_31_424971 297 28 its -PRON- PRP$ 10_1101-2020_12_31_424971 297 29 expression expression NN 10_1101-2020_12_31_424971 297 30 is be VBZ 10_1101-2020_12_31_424971 297 31 turned turn VBN 10_1101-2020_12_31_424971 297 32 off off RP 10_1101-2020_12_31_424971 297 33 to to TO 10_1101-2020_12_31_424971 297 34 allow allow VB 10_1101-2020_12_31_424971 297 35 immediate immediate JJ 10_1101-2020_12_31_424971 297 36 arrest arrest NN 10_1101-2020_12_31_424971 297 37 of of IN 10_1101-2020_12_31_424971 297 38 m1G37 m1g37 JJ 10_1101-2020_12_31_424971 297 39 synthesis synthesis NN 10_1101-2020_12_31_424971 297 40 . . . 10_1101-2020_12_31_424971 298 1 In in IN 10_1101-2020_12_31_424971 298 2 the the DT 10_1101-2020_12_31_424971 298 3 692 692 CD 10_1101-2020_12_31_424971 298 4 isogenic isogenic JJ 10_1101-2020_12_31_424971 298 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 298 6 and and CC 10_1101-2020_12_31_424971 298 7 ProL ProL NNP 10_1101-2020_12_31_424971 298 8 strains strain NNS 10_1101-2020_12_31_424971 298 9 , , , 10_1101-2020_12_31_424971 298 10 the the DT 10_1101-2020_12_31_424971 298 11 level level NN 10_1101-2020_12_31_424971 298 12 of of IN 10_1101-2020_12_31_424971 298 13 m1G37 m1g37 NN 10_1101-2020_12_31_424971 298 14 is be VBZ 10_1101-2020_12_31_424971 298 15 determined determine VBN 10_1101-2020_12_31_424971 298 16 by by IN 10_1101-2020_12_31_424971 298 17 the the DT 10_1101-2020_12_31_424971 298 18 concentration concentration NN 10_1101-2020_12_31_424971 298 19 of of IN 10_1101-2020_12_31_424971 298 20 the the DT 10_1101-2020_12_31_424971 298 21 693 693 CD 10_1101-2020_12_31_424971 298 22 added add VBN 10_1101-2020_12_31_424971 298 23 arabinose arabinose RB 10_1101-2020_12_31_424971 298 24 in in IN 10_1101-2020_12_31_424971 298 25 a a DT 10_1101-2020_12_31_424971 298 26 cellular cellular JJ 10_1101-2020_12_31_424971 298 27 context context NN 10_1101-2020_12_31_424971 298 28 that that WDT 10_1101-2020_12_31_424971 298 29 expresses express VBZ 10_1101-2020_12_31_424971 298 30 ProM ProM NNP 10_1101-2020_12_31_424971 298 31 as as IN 10_1101-2020_12_31_424971 298 32 the the DT 10_1101-2020_12_31_424971 298 33 only only JJ 10_1101-2020_12_31_424971 298 34 competing compete VBG 10_1101-2020_12_31_424971 298 35 tRNAPro tRNAPro NNP 10_1101-2020_12_31_424971 298 36 species species NN 10_1101-2020_12_31_424971 298 37 . . . 10_1101-2020_12_31_424971 299 1 694 694 CD 10_1101-2020_12_31_424971 299 2 In in IN 10_1101-2020_12_31_424971 299 3 the the DT 10_1101-2020_12_31_424971 299 4 m1G37 m1g37 NN 10_1101-2020_12_31_424971 299 5 + + CC 10_1101-2020_12_31_424971 299 6 condition condition NN 10_1101-2020_12_31_424971 299 7 , , , 10_1101-2020_12_31_424971 299 8 where where WRB 10_1101-2020_12_31_424971 299 9 arabinose arabinose DT 10_1101-2020_12_31_424971 299 10 was be VBD 10_1101-2020_12_31_424971 299 11 added add VBN 10_1101-2020_12_31_424971 299 12 to to IN 10_1101-2020_12_31_424971 299 13 0.2 0.2 CD 10_1101-2020_12_31_424971 299 14 % % NN 10_1101-2020_12_31_424971 299 15 in in IN 10_1101-2020_12_31_424971 299 16 the the DT 10_1101-2020_12_31_424971 299 17 medium medium NN 10_1101-2020_12_31_424971 299 18 , , , 10_1101-2020_12_31_424971 299 19 tRNA tRNA NNP 10_1101-2020_12_31_424971 299 20 substrates substrate VBZ 10_1101-2020_12_31_424971 299 21 695 695 CD 10_1101-2020_12_31_424971 299 22 of of IN 10_1101-2020_12_31_424971 299 23 N1-methylation N1-methylation NNP 10_1101-2020_12_31_424971 299 24 were be VBD 10_1101-2020_12_31_424971 299 25 confirmed confirm VBN 10_1101-2020_12_31_424971 299 26 to to TO 10_1101-2020_12_31_424971 299 27 be be VB 10_1101-2020_12_31_424971 299 28 100 100 CD 10_1101-2020_12_31_424971 299 29 % % NN 10_1101-2020_12_31_424971 299 30 methylated methylate VBN 10_1101-2020_12_31_424971 299 31 by by IN 10_1101-2020_12_31_424971 299 32 mass mass NNP 10_1101-2020_12_31_424971 299 33 spectrometry spectrometry NNP 10_1101-2020_12_31_424971 299 34 , , , 10_1101-2020_12_31_424971 299 35 whereas whereas IN 10_1101-2020_12_31_424971 299 36 in in IN 10_1101-2020_12_31_424971 299 37 the the DT 10_1101-2020_12_31_424971 299 38 696 696 CD 10_1101-2020_12_31_424971 299 39 m1G37 m1g37 NN 10_1101-2020_12_31_424971 299 40 – – : 10_1101-2020_12_31_424971 299 41 condition condition NN 10_1101-2020_12_31_424971 299 42 , , , 10_1101-2020_12_31_424971 299 43 where where WRB 10_1101-2020_12_31_424971 299 44 arabinose arabinose DT 10_1101-2020_12_31_424971 299 45 was be VBD 10_1101-2020_12_31_424971 299 46 not not RB 10_1101-2020_12_31_424971 299 47 added add VBN 10_1101-2020_12_31_424971 299 48 to to IN 10_1101-2020_12_31_424971 299 49 the the DT 10_1101-2020_12_31_424971 299 50 medium medium NN 10_1101-2020_12_31_424971 299 51 , , , 10_1101-2020_12_31_424971 299 52 tRNA trna JJ 10_1101-2020_12_31_424971 299 53 substrates substrate NNS 10_1101-2020_12_31_424971 299 54 of of IN 10_1101-2020_12_31_424971 299 55 N1 n1 NN 10_1101-2020_12_31_424971 299 56 - - HYPH 10_1101-2020_12_31_424971 299 57 697 697 CD 10_1101-2020_12_31_424971 299 58 methylation methylation NN 10_1101-2020_12_31_424971 299 59 were be VBD 10_1101-2020_12_31_424971 299 60 confirmed confirm VBN 10_1101-2020_12_31_424971 299 61 to to TO 10_1101-2020_12_31_424971 299 62 be be VB 10_1101-2020_12_31_424971 299 63 20 20 CD 10_1101-2020_12_31_424971 299 64 % % NN 10_1101-2020_12_31_424971 299 65 methylated methylate VBN 10_1101-2020_12_31_424971 299 66 by by IN 10_1101-2020_12_31_424971 299 67 mass mass NNP 10_1101-2020_12_31_424971 299 68 spectrometry30 spectrometry30 NNP 10_1101-2020_12_31_424971 299 69 . . . 10_1101-2020_12_31_424971 300 1 Each each DT 10_1101-2020_12_31_424971 300 2 strain strain NN 10_1101-2020_12_31_424971 300 3 was be VBD 10_1101-2020_12_31_424971 300 4 698 698 CD 10_1101-2020_12_31_424971 300 5 transformed transform VBN 10_1101-2020_12_31_424971 300 6 with with IN 10_1101-2020_12_31_424971 300 7 the the DT 10_1101-2020_12_31_424971 300 8 pKK223 pKK223 NNP 10_1101-2020_12_31_424971 300 9 - - HYPH 10_1101-2020_12_31_424971 300 10 3 3 CD 10_1101-2020_12_31_424971 300 11 plasmid plasmid NN 10_1101-2020_12_31_424971 300 12 expressing express VBG 10_1101-2020_12_31_424971 300 13 an an DT 10_1101-2020_12_31_424971 300 14 mRNA mrna NN 10_1101-2020_12_31_424971 300 15 with with IN 10_1101-2020_12_31_424971 300 16 a a DT 10_1101-2020_12_31_424971 300 17 CCC CCC NNP 10_1101-2020_12_31_424971 300 18 - - HYPH 10_1101-2020_12_31_424971 300 19 C C NNP 10_1101-2020_12_31_424971 300 20 motif motif NN 10_1101-2020_12_31_424971 300 21 at at IN 10_1101-2020_12_31_424971 300 22 the the DT 10_1101-2020_12_31_424971 300 23 2nd 2nd JJ 10_1101-2020_12_31_424971 300 24 codon codon NNP 10_1101-2020_12_31_424971 300 25 699 699 CD 10_1101-2020_12_31_424971 300 26 position position NN 10_1101-2020_12_31_424971 300 27 of of IN 10_1101-2020_12_31_424971 300 28 the the DT 10_1101-2020_12_31_424971 300 29 reporter reporter NN 10_1101-2020_12_31_424971 300 30 lacZ lacz CD 10_1101-2020_12_31_424971 300 31 gene gene NN 10_1101-2020_12_31_424971 300 32 . . . 10_1101-2020_12_31_424971 301 1 To to TO 10_1101-2020_12_31_424971 301 2 simplify simplify VB 10_1101-2020_12_31_424971 301 3 the the DT 10_1101-2020_12_31_424971 301 4 interpretation interpretation NN 10_1101-2020_12_31_424971 301 5 , , , 10_1101-2020_12_31_424971 301 6 the the DT 10_1101-2020_12_31_424971 301 7 natural natural JJ 10_1101-2020_12_31_424971 301 8 AUG AUG NNP 10_1101-2020_12_31_424971 301 9 codon codon NN 10_1101-2020_12_31_424971 301 10 at at IN 10_1101-2020_12_31_424971 301 11 the the DT 10_1101-2020_12_31_424971 301 12 5th 5th JJ 10_1101-2020_12_31_424971 301 13 700 700 CD 10_1101-2020_12_31_424971 301 14 position position NN 10_1101-2020_12_31_424971 301 15 of of IN 10_1101-2020_12_31_424971 301 16 lacZ lacz CD 10_1101-2020_12_31_424971 301 17 was be VBD 10_1101-2020_12_31_424971 301 18 removed remove VBN 10_1101-2020_12_31_424971 301 19 . . . 10_1101-2020_12_31_424971 302 1 A a DT 10_1101-2020_12_31_424971 302 2 +1 +1 : 10_1101-2020_12_31_424971 302 3 frameshift frameshift NN 10_1101-2020_12_31_424971 302 4 at at IN 10_1101-2020_12_31_424971 302 5 the the DT 10_1101-2020_12_31_424971 302 6 CCC CCC NNP 10_1101-2020_12_31_424971 302 7 - - HYPH 10_1101-2020_12_31_424971 302 8 C C NNP 10_1101-2020_12_31_424971 302 9 motif motif NN 10_1101-2020_12_31_424971 302 10 would would MD 10_1101-2020_12_31_424971 302 11 enable enable VB 10_1101-2020_12_31_424971 302 12 expression expression NN 10_1101-2020_12_31_424971 302 13 of of IN 10_1101-2020_12_31_424971 302 14 701 701 CD 10_1101-2020_12_31_424971 302 15 .CC .CC , 10_1101-2020_12_31_424971 302 16 - - HYPH 10_1101-2020_12_31_424971 302 17 BY by IN 10_1101-2020_12_31_424971 302 18 - - HYPH 10_1101-2020_12_31_424971 302 19 NC NC NNP 10_1101-2020_12_31_424971 302 20 - - HYPH 10_1101-2020_12_31_424971 302 21 ND ND NNP 10_1101-2020_12_31_424971 302 22 4.0 4.0 CD 10_1101-2020_12_31_424971 302 23 International International NNP 10_1101-2020_12_31_424971 302 24 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 302 25 under under IN 10_1101-2020_12_31_424971 302 26 a a DT 10_1101-2020_12_31_424971 302 27 ( ( -LRB- 10_1101-2020_12_31_424971 302 28 which which WDT 10_1101-2020_12_31_424971 302 29 was be VBD 10_1101-2020_12_31_424971 302 30 not not RB 10_1101-2020_12_31_424971 302 31 certified certify VBN 10_1101-2020_12_31_424971 302 32 by by IN 10_1101-2020_12_31_424971 302 33 peer peer NN 10_1101-2020_12_31_424971 302 34 review review NN 10_1101-2020_12_31_424971 302 35 ) ) -RRB- 10_1101-2020_12_31_424971 302 36 is be VBZ 10_1101-2020_12_31_424971 302 37 the the DT 10_1101-2020_12_31_424971 302 38 author author NN 10_1101-2020_12_31_424971 302 39 / / SYM 10_1101-2020_12_31_424971 302 40 funder funder NN 10_1101-2020_12_31_424971 302 41 , , , 10_1101-2020_12_31_424971 302 42 who who WP 10_1101-2020_12_31_424971 302 43 has have VBZ 10_1101-2020_12_31_424971 302 44 granted grant VBN 10_1101-2020_12_31_424971 302 45 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 302 46 a a DT 10_1101-2020_12_31_424971 302 47 license license NN 10_1101-2020_12_31_424971 302 48 to to TO 10_1101-2020_12_31_424971 302 49 display display VB 10_1101-2020_12_31_424971 302 50 the the DT 10_1101-2020_12_31_424971 302 51 preprint preprint NN 10_1101-2020_12_31_424971 302 52 in in IN 10_1101-2020_12_31_424971 302 53 perpetuity perpetuity NN 10_1101-2020_12_31_424971 302 54 . . . 10_1101-2020_12_31_424971 303 1 It -PRON- PRP 10_1101-2020_12_31_424971 303 2 is be VBZ 10_1101-2020_12_31_424971 303 3 made make VBN 10_1101-2020_12_31_424971 303 4 The the DT 10_1101-2020_12_31_424971 303 5 copyright copyright NN 10_1101-2020_12_31_424971 303 6 holder holder NN 10_1101-2020_12_31_424971 303 7 for for IN 10_1101-2020_12_31_424971 303 8 this this DT 10_1101-2020_12_31_424971 303 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 303 10 version version NN 10_1101-2020_12_31_424971 303 11 posted post VBD 10_1101-2020_12_31_424971 303 12 January January NNP 10_1101-2020_12_31_424971 303 13 2 2 CD 10_1101-2020_12_31_424971 303 14 , , , 10_1101-2020_12_31_424971 303 15 2021 2021 CD 10_1101-2020_12_31_424971 303 16 . . . 10_1101-2020_12_31_424971 303 17 ; ; : 10_1101-2020_12_31_424971 303 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 303 19 : : : 10_1101-2020_12_31_424971 303 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 303 21 preprint preprint NN 10_1101-2020_12_31_424971 303 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 303 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 303 24 29 29 CD 10_1101-2020_12_31_424971 303 25 lacZ. lacz. NN 10_1101-2020_12_31_424971 304 1 The the DT 10_1101-2020_12_31_424971 304 2 activity activity NN 10_1101-2020_12_31_424971 304 3 of of IN 10_1101-2020_12_31_424971 304 4 b b NN 10_1101-2020_12_31_424971 304 5 - - : 10_1101-2020_12_31_424971 304 6 Gal Gal NNP 10_1101-2020_12_31_424971 304 7 was be VBD 10_1101-2020_12_31_424971 304 8 directly directly RB 10_1101-2020_12_31_424971 304 9 measured measure VBN 10_1101-2020_12_31_424971 304 10 from from IN 10_1101-2020_12_31_424971 304 11 lysates lysate NNS 10_1101-2020_12_31_424971 304 12 of of IN 10_1101-2020_12_31_424971 304 13 cells cell NNS 10_1101-2020_12_31_424971 304 14 grown grow VBN 10_1101-2020_12_31_424971 304 15 in in IN 10_1101-2020_12_31_424971 304 16 the the DT 10_1101-2020_12_31_424971 304 17 presence presence NN 10_1101-2020_12_31_424971 304 18 or or CC 10_1101-2020_12_31_424971 304 19 702 702 CD 10_1101-2020_12_31_424971 304 20 absence absence NN 10_1101-2020_12_31_424971 304 21 of of IN 10_1101-2020_12_31_424971 304 22 0.2 0.2 CD 10_1101-2020_12_31_424971 304 23 % % NN 10_1101-2020_12_31_424971 304 24 arabinose arabinose RB 10_1101-2020_12_31_424971 304 25 to to TO 10_1101-2020_12_31_424971 304 26 induce induce VB 10_1101-2020_12_31_424971 304 27 or or CC 10_1101-2020_12_31_424971 304 28 not not RB 10_1101-2020_12_31_424971 304 29 induce induce VB 10_1101-2020_12_31_424971 304 30 , , , 10_1101-2020_12_31_424971 304 31 respectively respectively RB 10_1101-2020_12_31_424971 304 32 , , , 10_1101-2020_12_31_424971 304 33 the the DT 10_1101-2020_12_31_424971 304 34 plasmid plasmid NNP 10_1101-2020_12_31_424971 304 35 - - HYPH 10_1101-2020_12_31_424971 304 36 borne bear VBN 10_1101-2020_12_31_424971 304 37 human human JJ 10_1101-2020_12_31_424971 304 38 trm5 trm5 NN 10_1101-2020_12_31_424971 304 39 . . . 10_1101-2020_12_31_424971 305 1 703 703 CD 10_1101-2020_12_31_424971 305 2 In in IN 10_1101-2020_12_31_424971 305 3 these these DT 10_1101-2020_12_31_424971 305 4 assays assay NNS 10_1101-2020_12_31_424971 305 5 , , , 10_1101-2020_12_31_424971 305 6 decoding decode VBG 10_1101-2020_12_31_424971 305 7 of of IN 10_1101-2020_12_31_424971 305 8 the the DT 10_1101-2020_12_31_424971 305 9 CCC CCC NNP 10_1101-2020_12_31_424971 305 10 - - HYPH 10_1101-2020_12_31_424971 305 11 C C NNP 10_1101-2020_12_31_424971 305 12 codon codon NN 10_1101-2020_12_31_424971 305 13 motif motif NN 10_1101-2020_12_31_424971 305 14 would would MD 10_1101-2020_12_31_424971 305 15 be be VB 10_1101-2020_12_31_424971 305 16 mediated mediate VBN 10_1101-2020_12_31_424971 305 17 by by IN 10_1101-2020_12_31_424971 305 18 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 305 19 and and CC 10_1101-2020_12_31_424971 305 20 ProM ProM NNP 10_1101-2020_12_31_424971 305 21 in in IN 10_1101-2020_12_31_424971 305 22 704 704 CD 10_1101-2020_12_31_424971 305 23 the the DT 10_1101-2020_12_31_424971 305 24 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 305 25 strain strain NN 10_1101-2020_12_31_424971 305 26 , , , 10_1101-2020_12_31_424971 305 27 and and CC 10_1101-2020_12_31_424971 305 28 would would MD 10_1101-2020_12_31_424971 305 29 be be VB 10_1101-2020_12_31_424971 305 30 mediated mediate VBN 10_1101-2020_12_31_424971 305 31 by by IN 10_1101-2020_12_31_424971 305 32 ProL ProL NNP 10_1101-2020_12_31_424971 305 33 and and CC 10_1101-2020_12_31_424971 305 34 ProM ProM NNP 10_1101-2020_12_31_424971 305 35 in in IN 10_1101-2020_12_31_424971 305 36 the the DT 10_1101-2020_12_31_424971 305 37 ProL ProL NNP 10_1101-2020_12_31_424971 305 38 strain strain NN 10_1101-2020_12_31_424971 305 39 . . . 10_1101-2020_12_31_424971 306 1 Due due IN 10_1101-2020_12_31_424971 306 2 to to IN 10_1101-2020_12_31_424971 306 3 the the DT 10_1101-2020_12_31_424971 306 4 presence presence NN 10_1101-2020_12_31_424971 306 5 705 705 CD 10_1101-2020_12_31_424971 306 6 of of IN 10_1101-2020_12_31_424971 306 7 ProM ProM NNP 10_1101-2020_12_31_424971 306 8 in in IN 10_1101-2020_12_31_424971 306 9 both both DT 10_1101-2020_12_31_424971 306 10 strains strain NNS 10_1101-2020_12_31_424971 306 11 , , , 10_1101-2020_12_31_424971 306 12 there there EX 10_1101-2020_12_31_424971 306 13 would would MD 10_1101-2020_12_31_424971 306 14 be be VB 10_1101-2020_12_31_424971 306 15 no no DT 10_1101-2020_12_31_424971 306 16 vacancy vacancy NN 10_1101-2020_12_31_424971 306 17 at at IN 10_1101-2020_12_31_424971 306 18 the the DT 10_1101-2020_12_31_424971 306 19 CCC CCC NNP 10_1101-2020_12_31_424971 306 20 - - HYPH 10_1101-2020_12_31_424971 306 21 C C NNP 10_1101-2020_12_31_424971 306 22 codon codon NN 10_1101-2020_12_31_424971 306 23 motif motif NN 10_1101-2020_12_31_424971 306 24 . . . 10_1101-2020_12_31_424971 307 1 706 706 CD 10_1101-2020_12_31_424971 307 2 707 707 CD 10_1101-2020_12_31_424971 307 3 Cell cell NN 10_1101-2020_12_31_424971 307 4 - - HYPH 10_1101-2020_12_31_424971 307 5 based base VBN 10_1101-2020_12_31_424971 307 6 +1 +1 NNS 10_1101-2020_12_31_424971 307 7 frameshifting frameshifting NN 10_1101-2020_12_31_424971 307 8 lolB lolB NNS 10_1101-2020_12_31_424971 307 9 assays assays RB 10_1101-2020_12_31_424971 307 10 . . . 10_1101-2020_12_31_424971 308 1 To to TO 10_1101-2020_12_31_424971 308 2 quantify quantify VB 10_1101-2020_12_31_424971 308 3 the the DT 10_1101-2020_12_31_424971 308 4 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 308 5 efficiency efficiency NN 10_1101-2020_12_31_424971 308 6 at at IN 10_1101-2020_12_31_424971 308 7 the the DT 10_1101-2020_12_31_424971 308 8 708 708 CD 10_1101-2020_12_31_424971 308 9 CCC CCC NNP 10_1101-2020_12_31_424971 308 10 - - HYPH 10_1101-2020_12_31_424971 308 11 C C NNP 10_1101-2020_12_31_424971 308 12 motif motif NN 10_1101-2020_12_31_424971 308 13 at at IN 10_1101-2020_12_31_424971 308 14 the the DT 10_1101-2020_12_31_424971 308 15 2nd 2nd JJ 10_1101-2020_12_31_424971 308 16 codon codon NNP 10_1101-2020_12_31_424971 308 17 position position NN 10_1101-2020_12_31_424971 308 18 of of IN 10_1101-2020_12_31_424971 308 19 the the DT 10_1101-2020_12_31_424971 308 20 natural natural JJ 10_1101-2020_12_31_424971 308 21 lolB lolB NNP 10_1101-2020_12_31_424971 308 22 gene gene NN 10_1101-2020_12_31_424971 308 23 , , , 10_1101-2020_12_31_424971 308 24 the the DT 10_1101-2020_12_31_424971 308 25 ratio ratio NN 10_1101-2020_12_31_424971 308 26 of of IN 10_1101-2020_12_31_424971 308 27 protein protein NN 10_1101-2020_12_31_424971 308 28 synthesis synthesis NN 10_1101-2020_12_31_424971 308 29 of of IN 10_1101-2020_12_31_424971 308 30 709 709 CD 10_1101-2020_12_31_424971 308 31 lolB lolB NNS 10_1101-2020_12_31_424971 308 32 to to IN 10_1101-2020_12_31_424971 308 33 cysS cysS NNP 10_1101-2020_12_31_424971 308 34 was be VBD 10_1101-2020_12_31_424971 308 35 measured measure VBN 10_1101-2020_12_31_424971 308 36 by by IN 10_1101-2020_12_31_424971 308 37 Western western JJ 10_1101-2020_12_31_424971 308 38 blots blot NNS 10_1101-2020_12_31_424971 308 39 . . . 10_1101-2020_12_31_424971 309 1 Overnight overnight JJ 10_1101-2020_12_31_424971 309 2 cultures culture NNS 10_1101-2020_12_31_424971 309 3 of of IN 10_1101-2020_12_31_424971 309 4 the the DT 10_1101-2020_12_31_424971 309 5 isogenic isogenic JJ 10_1101-2020_12_31_424971 309 6 strains strain NNS 10_1101-2020_12_31_424971 309 7 expressing express VBG 10_1101-2020_12_31_424971 309 8 710 710 CD 10_1101-2020_12_31_424971 309 9 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 309 10 or or CC 10_1101-2020_12_31_424971 309 11 ProL ProL NNP 10_1101-2020_12_31_424971 309 12 were be VBD 10_1101-2020_12_31_424971 309 13 separately separately RB 10_1101-2020_12_31_424971 309 14 inoculated inoculate VBN 10_1101-2020_12_31_424971 309 15 into into IN 10_1101-2020_12_31_424971 309 16 fresh fresh JJ 10_1101-2020_12_31_424971 309 17 LB LB NNP 10_1101-2020_12_31_424971 309 18 media medium NNS 10_1101-2020_12_31_424971 309 19 in in IN 10_1101-2020_12_31_424971 309 20 the the DT 10_1101-2020_12_31_424971 309 21 presence presence NN 10_1101-2020_12_31_424971 309 22 or or CC 10_1101-2020_12_31_424971 309 23 absence absence NN 10_1101-2020_12_31_424971 309 24 of of IN 10_1101-2020_12_31_424971 309 25 0.2 0.2 CD 10_1101-2020_12_31_424971 309 26 % % NN 10_1101-2020_12_31_424971 309 27 711 711 CD 10_1101-2020_12_31_424971 309 28 arabinose arabinose RB 10_1101-2020_12_31_424971 309 29 and and CC 10_1101-2020_12_31_424971 309 30 were be VBD 10_1101-2020_12_31_424971 309 31 grown grow VBN 10_1101-2020_12_31_424971 309 32 for for IN 10_1101-2020_12_31_424971 309 33 4 4 CD 10_1101-2020_12_31_424971 309 34 h h NN 10_1101-2020_12_31_424971 309 35 to to TO 10_1101-2020_12_31_424971 309 36 produce produce VB 10_1101-2020_12_31_424971 309 37 the the DT 10_1101-2020_12_31_424971 309 38 m1G37 m1g37 NN 10_1101-2020_12_31_424971 309 39 + + SYM 10_1101-2020_12_31_424971 309 40 and and CC 10_1101-2020_12_31_424971 309 41 m1G37 m1g37 NN 10_1101-2020_12_31_424971 309 42 – – : 10_1101-2020_12_31_424971 309 43 conditions condition NNS 10_1101-2020_12_31_424971 309 44 , , , 10_1101-2020_12_31_424971 309 45 respectively respectively RB 10_1101-2020_12_31_424971 309 46 . . . 10_1101-2020_12_31_424971 310 1 712 712 CD 10_1101-2020_12_31_424971 310 2 Cultures culture NNS 10_1101-2020_12_31_424971 310 3 were be VBD 10_1101-2020_12_31_424971 310 4 diluted dilute VBN 10_1101-2020_12_31_424971 310 5 10- 10- CD 10_1101-2020_12_31_424971 310 6 to to IN 10_1101-2020_12_31_424971 310 7 16-fold 16-fold CD 10_1101-2020_12_31_424971 310 8 into into IN 10_1101-2020_12_31_424971 310 9 fresh fresh JJ 10_1101-2020_12_31_424971 310 10 media medium NNS 10_1101-2020_12_31_424971 310 11 to to IN 10_1101-2020_12_31_424971 310 12 an an DT 10_1101-2020_12_31_424971 310 13 optical optical JJ 10_1101-2020_12_31_424971 310 14 density density NN 10_1101-2020_12_31_424971 310 15 ( ( -LRB- 10_1101-2020_12_31_424971 310 16 OD OD NNP 10_1101-2020_12_31_424971 310 17 ) ) -RRB- 10_1101-2020_12_31_424971 310 18 of of IN 10_1101-2020_12_31_424971 310 19 ~0.1 ~0.1 CD 10_1101-2020_12_31_424971 310 20 and and CC 10_1101-2020_12_31_424971 310 21 grown grow VBD 10_1101-2020_12_31_424971 310 22 713 713 CD 10_1101-2020_12_31_424971 310 23 for for IN 10_1101-2020_12_31_424971 310 24 another another DT 10_1101-2020_12_31_424971 310 25 3 3 CD 10_1101-2020_12_31_424971 310 26 h. h. NN 10_1101-2020_12_31_424971 310 27 Cells cell NNS 10_1101-2020_12_31_424971 310 28 were be VBD 10_1101-2020_12_31_424971 310 29 harvested harvest VBN 10_1101-2020_12_31_424971 310 30 and and CC 10_1101-2020_12_31_424971 310 31 15 15 CD 10_1101-2020_12_31_424971 310 32 µg µg NN 10_1101-2020_12_31_424971 310 33 of of IN 10_1101-2020_12_31_424971 310 34 total total JJ 10_1101-2020_12_31_424971 310 35 protein protein NN 10_1101-2020_12_31_424971 310 36 from from IN 10_1101-2020_12_31_424971 310 37 cell cell NN 10_1101-2020_12_31_424971 310 38 lysates lysate NNS 10_1101-2020_12_31_424971 310 39 was be VBD 10_1101-2020_12_31_424971 310 40 separated separate VBN 10_1101-2020_12_31_424971 310 41 714 714 CD 10_1101-2020_12_31_424971 310 42 on on IN 10_1101-2020_12_31_424971 310 43 12 12 CD 10_1101-2020_12_31_424971 310 44 % % NN 10_1101-2020_12_31_424971 310 45 SDS SDS NNP 10_1101-2020_12_31_424971 310 46 - - HYPH 10_1101-2020_12_31_424971 310 47 PAGE PAGE NNP 10_1101-2020_12_31_424971 310 48 and and CC 10_1101-2020_12_31_424971 310 49 probed probe VBN 10_1101-2020_12_31_424971 310 50 with with IN 10_1101-2020_12_31_424971 310 51 rabbit rabbit NN 10_1101-2020_12_31_424971 310 52 polyclonal polyclonal JJ 10_1101-2020_12_31_424971 310 53 primary primary JJ 10_1101-2020_12_31_424971 310 54 antibodies antibody NNS 10_1101-2020_12_31_424971 310 55 against against IN 10_1101-2020_12_31_424971 310 56 LolB LolB NNP 10_1101-2020_12_31_424971 310 57 ( ( -LRB- 10_1101-2020_12_31_424971 310 58 at at IN 10_1101-2020_12_31_424971 310 59 a a DT 10_1101-2020_12_31_424971 310 60 715 715 CD 10_1101-2020_12_31_424971 310 61 10,000 10,000 CD 10_1101-2020_12_31_424971 310 62 dilution dilution NN 10_1101-2020_12_31_424971 310 63 ) ) -RRB- 10_1101-2020_12_31_424971 310 64 and and CC 10_1101-2020_12_31_424971 310 65 against against IN 10_1101-2020_12_31_424971 310 66 CysRS CysRS NNP 10_1101-2020_12_31_424971 310 67 ( ( -LRB- 10_1101-2020_12_31_424971 310 68 at at IN 10_1101-2020_12_31_424971 310 69 a a DT 10_1101-2020_12_31_424971 310 70 20,000 20,000 CD 10_1101-2020_12_31_424971 310 71 dilution dilution NN 10_1101-2020_12_31_424971 310 72 ) ) -RRB- 10_1101-2020_12_31_424971 310 73 , , , 10_1101-2020_12_31_424971 310 74 followed follow VBN 10_1101-2020_12_31_424971 310 75 by by IN 10_1101-2020_12_31_424971 310 76 goat goat NN 10_1101-2020_12_31_424971 310 77 polyclonal polyclonal JJ 10_1101-2020_12_31_424971 310 78 anti anti JJ 10_1101-2020_12_31_424971 310 79 - - JJ 10_1101-2020_12_31_424971 310 80 rabbit rabbit JJ 10_1101-2020_12_31_424971 310 81 716 716 CD 10_1101-2020_12_31_424971 310 82 IgG IgG NNP 10_1101-2020_12_31_424971 310 83 secondary secondary JJ 10_1101-2020_12_31_424971 310 84 antibody antibody NN 10_1101-2020_12_31_424971 310 85 ( ( -LRB- 10_1101-2020_12_31_424971 310 86 Sigma Sigma NNP 10_1101-2020_12_31_424971 310 87 - - HYPH 10_1101-2020_12_31_424971 310 88 Aldrich Aldrich NNP 10_1101-2020_12_31_424971 310 89 , , , 10_1101-2020_12_31_424971 310 90 # # NNP 10_1101-2020_12_31_424971 310 91 A0545 A0545 NNP 10_1101-2020_12_31_424971 310 92 ) ) -RRB- 10_1101-2020_12_31_424971 310 93 . . . 10_1101-2020_12_31_424971 311 1 The the DT 10_1101-2020_12_31_424971 311 2 ratio ratio NN 10_1101-2020_12_31_424971 311 3 of of IN 10_1101-2020_12_31_424971 311 4 protein protein NN 10_1101-2020_12_31_424971 311 5 synthesis synthesis NN 10_1101-2020_12_31_424971 311 6 of of IN 10_1101-2020_12_31_424971 311 7 lolB lolB NNP 10_1101-2020_12_31_424971 311 8 to to IN 10_1101-2020_12_31_424971 311 9 cysS cysS NNP 10_1101-2020_12_31_424971 311 10 717 717 CD 10_1101-2020_12_31_424971 311 11 was be VBD 10_1101-2020_12_31_424971 311 12 quantified quantify VBN 10_1101-2020_12_31_424971 311 13 using use VBG 10_1101-2020_12_31_424971 311 14 Super Super NNP 10_1101-2020_12_31_424971 311 15 Signal Signal NNP 10_1101-2020_12_31_424971 311 16 West West NNP 10_1101-2020_12_31_424971 311 17 Pico Pico NNP 10_1101-2020_12_31_424971 311 18 Chemiluminescent Chemiluminescent NNP 10_1101-2020_12_31_424971 311 19 substrate substrate NN 10_1101-2020_12_31_424971 311 20 ( ( -LRB- 10_1101-2020_12_31_424971 311 21 Thermo Thermo NNP 10_1101-2020_12_31_424971 311 22 Fischer Fischer NNP 10_1101-2020_12_31_424971 311 23 ) ) -RRB- 10_1101-2020_12_31_424971 311 24 in in IN 10_1101-2020_12_31_424971 311 25 a a DT 10_1101-2020_12_31_424971 311 26 718 718 CD 10_1101-2020_12_31_424971 311 27 Chemi Chemi NNP 10_1101-2020_12_31_424971 311 28 - - HYPH 10_1101-2020_12_31_424971 311 29 Doc Doc NNP 10_1101-2020_12_31_424971 311 30 XR XR NNP 10_1101-2020_12_31_424971 311 31 imager imager NN 10_1101-2020_12_31_424971 311 32 ( ( -LRB- 10_1101-2020_12_31_424971 311 33 Bio Bio NNP 10_1101-2020_12_31_424971 311 34 - - HYPH 10_1101-2020_12_31_424971 311 35 Rad Rad NNP 10_1101-2020_12_31_424971 311 36 ) ) -RRB- 10_1101-2020_12_31_424971 311 37 and and CC 10_1101-2020_12_31_424971 311 38 analyzed analyze VBN 10_1101-2020_12_31_424971 311 39 by by IN 10_1101-2020_12_31_424971 311 40 Image Image NNP 10_1101-2020_12_31_424971 311 41 Lab Lab NNP 10_1101-2020_12_31_424971 311 42 software software NN 10_1101-2020_12_31_424971 311 43 ( ( -LRB- 10_1101-2020_12_31_424971 311 44 Bio Bio NNP 10_1101-2020_12_31_424971 311 45 - - HYPH 10_1101-2020_12_31_424971 311 46 Rad Rad NNP 10_1101-2020_12_31_424971 311 47 , , , 10_1101-2020_12_31_424971 311 48 SOFT soft JJ 10_1101-2020_12_31_424971 311 49 - - HYPH 10_1101-2020_12_31_424971 311 50 LIT-170 lit-170 PRP 10_1101-2020_12_31_424971 311 51 - - HYPH 10_1101-2020_12_31_424971 311 52 719 719 CD 10_1101-2020_12_31_424971 311 53 9690-ILSPC 9690-ilspc CD 10_1101-2020_12_31_424971 311 54 - - HYPH 10_1101-2020_12_31_424971 311 55 V-6 V-6 NNP 10_1101-2020_12_31_424971 311 56 - - HYPH 10_1101-2020_12_31_424971 311 57 1 1 NNP 10_1101-2020_12_31_424971 311 58 ) ) -RRB- 10_1101-2020_12_31_424971 311 59 . . . 10_1101-2020_12_31_424971 312 1 To to TO 10_1101-2020_12_31_424971 312 2 measure measure VB 10_1101-2020_12_31_424971 312 3 the the DT 10_1101-2020_12_31_424971 312 4 +1-frameshifting +1-frameshifting JJ 10_1101-2020_12_31_424971 312 5 efficiency efficiency NN 10_1101-2020_12_31_424971 312 6 , , , 10_1101-2020_12_31_424971 312 7 we -PRON- PRP 10_1101-2020_12_31_424971 312 8 measured measure VBD 10_1101-2020_12_31_424971 312 9 the the DT 10_1101-2020_12_31_424971 312 10 ratio ratio NN 10_1101-2020_12_31_424971 312 11 of of IN 10_1101-2020_12_31_424971 312 12 protein protein NN 10_1101-2020_12_31_424971 312 13 720 720 CD 10_1101-2020_12_31_424971 312 14 synthesis synthesis NN 10_1101-2020_12_31_424971 312 15 of of IN 10_1101-2020_12_31_424971 312 16 lolB lolB NNS 10_1101-2020_12_31_424971 312 17 to to IN 10_1101-2020_12_31_424971 312 18 cysS cys NNS 10_1101-2020_12_31_424971 312 19 for for IN 10_1101-2020_12_31_424971 312 20 each each DT 10_1101-2020_12_31_424971 312 21 tRNA trna NN 10_1101-2020_12_31_424971 312 22 in in IN 10_1101-2020_12_31_424971 312 23 each each DT 10_1101-2020_12_31_424971 312 24 condition condition NN 10_1101-2020_12_31_424971 312 25 , , , 10_1101-2020_12_31_424971 312 26 and and CC 10_1101-2020_12_31_424971 312 27 we -PRON- PRP 10_1101-2020_12_31_424971 312 28 normalized normalize VBD 10_1101-2020_12_31_424971 312 29 the the DT 10_1101-2020_12_31_424971 312 30 observed observed JJ 10_1101-2020_12_31_424971 312 31 ratio ratio NN 10_1101-2020_12_31_424971 312 32 721 721 CD 10_1101-2020_12_31_424971 312 33 in in IN 10_1101-2020_12_31_424971 312 34 the the DT 10_1101-2020_12_31_424971 312 35 control control NN 10_1101-2020_12_31_424971 312 36 sample sample NN 10_1101-2020_12_31_424971 312 37 ( ( -LRB- 10_1101-2020_12_31_424971 312 38 i.e. i.e. FW 10_1101-2020_12_31_424971 312 39 , , , 10_1101-2020_12_31_424971 312 40 ProL ProL NNP 10_1101-2020_12_31_424971 312 41 in in IN 10_1101-2020_12_31_424971 312 42 the the DT 10_1101-2020_12_31_424971 312 43 m1G37 m1g37 NN 10_1101-2020_12_31_424971 312 44 + + SYM 10_1101-2020_12_31_424971 312 45 condition condition NN 10_1101-2020_12_31_424971 312 46 ) ) -RRB- 10_1101-2020_12_31_424971 312 47 to to IN 10_1101-2020_12_31_424971 312 48 1.0 1.0 CD 10_1101-2020_12_31_424971 312 49 , , , 10_1101-2020_12_31_424971 312 50 indicating indicate VBG 10_1101-2020_12_31_424971 312 51 that that IN 10_1101-2020_12_31_424971 312 52 protein protein NN 10_1101-2020_12_31_424971 312 53 synthesis synthesis NN 10_1101-2020_12_31_424971 312 54 722 722 CD 10_1101-2020_12_31_424971 312 55 of of IN 10_1101-2020_12_31_424971 312 56 these these DT 10_1101-2020_12_31_424971 312 57 two two CD 10_1101-2020_12_31_424971 312 58 genes gene NNS 10_1101-2020_12_31_424971 312 59 was be VBD 10_1101-2020_12_31_424971 312 60 in in IN 10_1101-2020_12_31_424971 312 61 the the DT 10_1101-2020_12_31_424971 312 62 0-frame 0-frame CD 10_1101-2020_12_31_424971 312 63 and and CC 10_1101-2020_12_31_424971 312 64 no no DT 10_1101-2020_12_31_424971 312 65 +1 +1 NNS 10_1101-2020_12_31_424971 312 66 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 312 67 . . . 10_1101-2020_12_31_424971 313 1 A a DT 10_1101-2020_12_31_424971 313 2 decrease decrease NN 10_1101-2020_12_31_424971 313 3 of of IN 10_1101-2020_12_31_424971 313 4 this this DT 10_1101-2020_12_31_424971 313 5 ratio ratio NN 10_1101-2020_12_31_424971 313 6 was be VBD 10_1101-2020_12_31_424971 313 7 723 723 CD 10_1101-2020_12_31_424971 313 8 interpreted interpret VBN 10_1101-2020_12_31_424971 313 9 as as IN 10_1101-2020_12_31_424971 313 10 a a DT 10_1101-2020_12_31_424971 313 11 proxy proxy NN 10_1101-2020_12_31_424971 313 12 of of IN 10_1101-2020_12_31_424971 313 13 +1 +1 NNS 10_1101-2020_12_31_424971 313 14 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 313 15 at at IN 10_1101-2020_12_31_424971 313 16 the the DT 10_1101-2020_12_31_424971 313 17 CCC CCC NNP 10_1101-2020_12_31_424971 313 18 - - HYPH 10_1101-2020_12_31_424971 313 19 C C NNP 10_1101-2020_12_31_424971 313 20 motif motif NN 10_1101-2020_12_31_424971 313 21 at at IN 10_1101-2020_12_31_424971 313 22 the the DT 10_1101-2020_12_31_424971 313 23 2nd 2nd JJ 10_1101-2020_12_31_424971 313 24 codon codon NN 10_1101-2020_12_31_424971 313 25 position position NN 10_1101-2020_12_31_424971 313 26 of of IN 10_1101-2020_12_31_424971 313 27 lolB. lolb. NN 10_1101-2020_12_31_424971 314 1 724 724 CD 10_1101-2020_12_31_424971 314 2 From from IN 10_1101-2020_12_31_424971 314 3 the the DT 10_1101-2020_12_31_424971 314 4 observed observed JJ 10_1101-2020_12_31_424971 314 5 ratio ratio NN 10_1101-2020_12_31_424971 314 6 of of IN 10_1101-2020_12_31_424971 314 7 each each DT 10_1101-2020_12_31_424971 314 8 sample sample NN 10_1101-2020_12_31_424971 314 9 in in IN 10_1101-2020_12_31_424971 314 10 each each DT 10_1101-2020_12_31_424971 314 11 condition condition NN 10_1101-2020_12_31_424971 314 12 , , , 10_1101-2020_12_31_424971 314 13 we -PRON- PRP 10_1101-2020_12_31_424971 314 14 calculated calculate VBD 10_1101-2020_12_31_424971 314 15 the the DT 10_1101-2020_12_31_424971 314 16 +1 +1 NNS 10_1101-2020_12_31_424971 314 17 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 314 18 725 725 CD 10_1101-2020_12_31_424971 314 19 efficiency efficiency NN 10_1101-2020_12_31_424971 314 20 relative relative JJ 10_1101-2020_12_31_424971 314 21 to to IN 10_1101-2020_12_31_424971 314 22 the the DT 10_1101-2020_12_31_424971 314 23 control control NN 10_1101-2020_12_31_424971 314 24 sample sample NN 10_1101-2020_12_31_424971 314 25 . . . 10_1101-2020_12_31_424971 315 1 726 726 CD 10_1101-2020_12_31_424971 315 2 727 727 CD 10_1101-2020_12_31_424971 315 3 .CC .CC : 10_1101-2020_12_31_424971 315 4 - - HYPH 10_1101-2020_12_31_424971 315 5 BY by IN 10_1101-2020_12_31_424971 315 6 - - HYPH 10_1101-2020_12_31_424971 315 7 NC NC NNP 10_1101-2020_12_31_424971 315 8 - - HYPH 10_1101-2020_12_31_424971 315 9 ND ND NNP 10_1101-2020_12_31_424971 315 10 4.0 4.0 CD 10_1101-2020_12_31_424971 315 11 International International NNP 10_1101-2020_12_31_424971 315 12 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 315 13 under under IN 10_1101-2020_12_31_424971 315 14 a a DT 10_1101-2020_12_31_424971 315 15 ( ( -LRB- 10_1101-2020_12_31_424971 315 16 which which WDT 10_1101-2020_12_31_424971 315 17 was be VBD 10_1101-2020_12_31_424971 315 18 not not RB 10_1101-2020_12_31_424971 315 19 certified certify VBN 10_1101-2020_12_31_424971 315 20 by by IN 10_1101-2020_12_31_424971 315 21 peer peer NN 10_1101-2020_12_31_424971 315 22 review review NN 10_1101-2020_12_31_424971 315 23 ) ) -RRB- 10_1101-2020_12_31_424971 315 24 is be VBZ 10_1101-2020_12_31_424971 315 25 the the DT 10_1101-2020_12_31_424971 315 26 author author NN 10_1101-2020_12_31_424971 315 27 / / SYM 10_1101-2020_12_31_424971 315 28 funder funder NN 10_1101-2020_12_31_424971 315 29 , , , 10_1101-2020_12_31_424971 315 30 who who WP 10_1101-2020_12_31_424971 315 31 has have VBZ 10_1101-2020_12_31_424971 315 32 granted grant VBN 10_1101-2020_12_31_424971 315 33 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 315 34 a a DT 10_1101-2020_12_31_424971 315 35 license license NN 10_1101-2020_12_31_424971 315 36 to to TO 10_1101-2020_12_31_424971 315 37 display display VB 10_1101-2020_12_31_424971 315 38 the the DT 10_1101-2020_12_31_424971 315 39 preprint preprint NN 10_1101-2020_12_31_424971 315 40 in in IN 10_1101-2020_12_31_424971 315 41 perpetuity perpetuity NN 10_1101-2020_12_31_424971 315 42 . . . 10_1101-2020_12_31_424971 316 1 It -PRON- PRP 10_1101-2020_12_31_424971 316 2 is be VBZ 10_1101-2020_12_31_424971 316 3 made make VBN 10_1101-2020_12_31_424971 316 4 The the DT 10_1101-2020_12_31_424971 316 5 copyright copyright NN 10_1101-2020_12_31_424971 316 6 holder holder NN 10_1101-2020_12_31_424971 316 7 for for IN 10_1101-2020_12_31_424971 316 8 this this DT 10_1101-2020_12_31_424971 316 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 316 10 version version NN 10_1101-2020_12_31_424971 316 11 posted post VBD 10_1101-2020_12_31_424971 316 12 January January NNP 10_1101-2020_12_31_424971 316 13 2 2 CD 10_1101-2020_12_31_424971 316 14 , , , 10_1101-2020_12_31_424971 316 15 2021 2021 CD 10_1101-2020_12_31_424971 316 16 . . . 10_1101-2020_12_31_424971 316 17 ; ; : 10_1101-2020_12_31_424971 316 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 316 19 : : : 10_1101-2020_12_31_424971 316 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 316 21 preprint preprint NN 10_1101-2020_12_31_424971 316 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 316 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 316 24 30 30 CD 10_1101-2020_12_31_424971 316 25 Cell cell NN 10_1101-2020_12_31_424971 316 26 - - HYPH 10_1101-2020_12_31_424971 316 27 free free JJ 10_1101-2020_12_31_424971 316 28 PURExpress purexpress NN 10_1101-2020_12_31_424971 316 29 in in IN 10_1101-2020_12_31_424971 316 30 vitro vitro FW 10_1101-2020_12_31_424971 316 31 translation translation NN 10_1101-2020_12_31_424971 316 32 assays assays RB 10_1101-2020_12_31_424971 316 33 . . . 10_1101-2020_12_31_424971 317 1 The the DT 10_1101-2020_12_31_424971 317 2 folA fola JJ 10_1101-2020_12_31_424971 317 3 gene gene NN 10_1101-2020_12_31_424971 317 4 , , , 10_1101-2020_12_31_424971 317 5 provided provide VBN 10_1101-2020_12_31_424971 317 6 as as IN 10_1101-2020_12_31_424971 317 7 part part NN 10_1101-2020_12_31_424971 317 8 of of IN 10_1101-2020_12_31_424971 317 9 the the DT 10_1101-2020_12_31_424971 317 10 E. E. NNP 10_1101-2020_12_31_424971 317 11 728 728 CD 10_1101-2020_12_31_424971 317 12 coli coli NNS 10_1101-2020_12_31_424971 317 13 PURExpress purexpress NN 10_1101-2020_12_31_424971 317 14 ( ( -LRB- 10_1101-2020_12_31_424971 317 15 New New NNP 10_1101-2020_12_31_424971 317 16 England England NNP 10_1101-2020_12_31_424971 317 17 BioLabs BioLabs NNP 10_1101-2020_12_31_424971 317 18 ) ) -RRB- 10_1101-2020_12_31_424971 317 19 in in IN 10_1101-2020_12_31_424971 317 20 vitro vitro FW 10_1101-2020_12_31_424971 317 21 translation translation NN 10_1101-2020_12_31_424971 317 22 system system NN 10_1101-2020_12_31_424971 317 23 , , , 10_1101-2020_12_31_424971 317 24 was be VBD 10_1101-2020_12_31_424971 317 25 modified modify VBN 10_1101-2020_12_31_424971 317 26 by by IN 10_1101-2020_12_31_424971 317 27 site site NN 10_1101-2020_12_31_424971 317 28 - - HYPH 10_1101-2020_12_31_424971 317 29 directed direct VBN 10_1101-2020_12_31_424971 317 30 729 729 CD 10_1101-2020_12_31_424971 317 31 mutagenesis mutagenesis NN 10_1101-2020_12_31_424971 317 32 to to TO 10_1101-2020_12_31_424971 317 33 introduce introduce VB 10_1101-2020_12_31_424971 317 34 a a DT 10_1101-2020_12_31_424971 317 35 CCC CCC NNP 10_1101-2020_12_31_424971 317 36 - - HYPH 10_1101-2020_12_31_424971 317 37 C C NNP 10_1101-2020_12_31_424971 317 38 motif motif NN 10_1101-2020_12_31_424971 317 39 into into IN 10_1101-2020_12_31_424971 317 40 the the DT 10_1101-2020_12_31_424971 317 41 5th 5th JJ 10_1101-2020_12_31_424971 317 42 codon codon NN 10_1101-2020_12_31_424971 317 43 position position NN 10_1101-2020_12_31_424971 317 44 . . . 10_1101-2020_12_31_424971 318 1 If if IN 10_1101-2020_12_31_424971 318 2 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 318 3 induced induce VBN 10_1101-2020_12_31_424971 318 4 +1 +1 NNS 10_1101-2020_12_31_424971 318 5 730 730 CD 10_1101-2020_12_31_424971 318 6 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 318 7 at at IN 10_1101-2020_12_31_424971 318 8 this this DT 10_1101-2020_12_31_424971 318 9 motif motif NN 10_1101-2020_12_31_424971 318 10 , , , 10_1101-2020_12_31_424971 318 11 a a DT 10_1101-2020_12_31_424971 318 12 full full JJ 10_1101-2020_12_31_424971 318 13 - - HYPH 10_1101-2020_12_31_424971 318 14 length length NN 10_1101-2020_12_31_424971 318 15 DHFR DHFR NNP 10_1101-2020_12_31_424971 318 16 would would MD 10_1101-2020_12_31_424971 318 17 be be VB 10_1101-2020_12_31_424971 318 18 made make VBN 10_1101-2020_12_31_424971 318 19 , , , 10_1101-2020_12_31_424971 318 20 whereas whereas IN 10_1101-2020_12_31_424971 318 21 if if IN 10_1101-2020_12_31_424971 318 22 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 318 23 failed fail VBD 10_1101-2020_12_31_424971 318 24 to to TO 10_1101-2020_12_31_424971 318 25 do do VB 10_1101-2020_12_31_424971 318 26 so so RB 10_1101-2020_12_31_424971 318 27 , , , 10_1101-2020_12_31_424971 318 28 a a DT 10_1101-2020_12_31_424971 318 29 731 731 CD 10_1101-2020_12_31_424971 318 30 C c NN 10_1101-2020_12_31_424971 318 31 - - HYPH 10_1101-2020_12_31_424971 318 32 terminal terminal JJ 10_1101-2020_12_31_424971 318 33 truncated truncate VBN 10_1101-2020_12_31_424971 318 34 fragment fragment NN 10_1101-2020_12_31_424971 318 35 ( ( -LRB- 10_1101-2020_12_31_424971 318 36 DC DC NNP 10_1101-2020_12_31_424971 318 37 ) ) -RRB- 10_1101-2020_12_31_424971 318 38 would would MD 10_1101-2020_12_31_424971 318 39 be be VB 10_1101-2020_12_31_424971 318 40 made make VBN 10_1101-2020_12_31_424971 318 41 due due JJ 10_1101-2020_12_31_424971 318 42 to to IN 10_1101-2020_12_31_424971 318 43 premature premature JJ 10_1101-2020_12_31_424971 318 44 termination termination NN 10_1101-2020_12_31_424971 318 45 of of IN 10_1101-2020_12_31_424971 318 46 protein protein NN 10_1101-2020_12_31_424971 318 47 732 732 CD 10_1101-2020_12_31_424971 318 48 synthesis synthesis NN 10_1101-2020_12_31_424971 318 49 . . . 10_1101-2020_12_31_424971 319 1 Because because IN 10_1101-2020_12_31_424971 319 2 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 319 3 has have VBZ 10_1101-2020_12_31_424971 319 4 no no DT 10_1101-2020_12_31_424971 319 5 orthogonal orthogonal JJ 10_1101-2020_12_31_424971 319 6 tRNA trna NN 10_1101-2020_12_31_424971 319 7 synthetase synthetase NN 10_1101-2020_12_31_424971 319 8 for for IN 10_1101-2020_12_31_424971 319 9 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 319 10 with with IN 10_1101-2020_12_31_424971 319 11 a a DT 10_1101-2020_12_31_424971 319 12 non-733 non-733 NNP 10_1101-2020_12_31_424971 319 13 proteinogenic proteinogenic NN 10_1101-2020_12_31_424971 319 14 amino amino NN 10_1101-2020_12_31_424971 319 15 acid acid NN 10_1101-2020_12_31_424971 319 16 , , , 10_1101-2020_12_31_424971 319 17 we -PRON- PRP 10_1101-2020_12_31_424971 319 18 used use VBD 10_1101-2020_12_31_424971 319 19 the the DT 10_1101-2020_12_31_424971 319 20 Flexizyme Flexizyme NNP 10_1101-2020_12_31_424971 319 21 ribozyme ribozyme NNS 10_1101-2020_12_31_424971 319 22 technology32 technology32 VB 10_1101-2020_12_31_424971 319 23 for for IN 10_1101-2020_12_31_424971 319 24 this this DT 10_1101-2020_12_31_424971 319 25 purpose purpose NN 10_1101-2020_12_31_424971 319 26 . . . 10_1101-2020_12_31_424971 320 1 734 734 CD 10_1101-2020_12_31_424971 320 2 Coupled couple VBN 10_1101-2020_12_31_424971 320 3 in in IN 10_1101-2020_12_31_424971 320 4 vitro vitro FW 10_1101-2020_12_31_424971 320 5 transcription transcription NN 10_1101-2020_12_31_424971 320 6 - - HYPH 10_1101-2020_12_31_424971 320 7 translation translation NN 10_1101-2020_12_31_424971 320 8 of of IN 10_1101-2020_12_31_424971 320 9 the the DT 10_1101-2020_12_31_424971 320 10 modified modify VBN 10_1101-2020_12_31_424971 320 11 E. E. NNP 10_1101-2020_12_31_424971 320 12 coli coli NNS 10_1101-2020_12_31_424971 320 13 folA fola JJ 10_1101-2020_12_31_424971 320 14 gene gene NN 10_1101-2020_12_31_424971 320 15 containing contain VBG 10_1101-2020_12_31_424971 320 16 the the DT 10_1101-2020_12_31_424971 320 17 CCC CCC NNP 10_1101-2020_12_31_424971 320 18 - - HYPH 10_1101-2020_12_31_424971 320 19 C C NNP 10_1101-2020_12_31_424971 320 20 735 735 CD 10_1101-2020_12_31_424971 320 21 motif motif NN 10_1101-2020_12_31_424971 320 22 at at IN 10_1101-2020_12_31_424971 320 23 the the DT 10_1101-2020_12_31_424971 320 24 5th 5th JJ 10_1101-2020_12_31_424971 320 25 codon codon NN 10_1101-2020_12_31_424971 320 26 position position NN 10_1101-2020_12_31_424971 320 27 was be VBD 10_1101-2020_12_31_424971 320 28 conducted conduct VBN 10_1101-2020_12_31_424971 320 29 in in IN 10_1101-2020_12_31_424971 320 30 the the DT 10_1101-2020_12_31_424971 320 31 presence presence NN 10_1101-2020_12_31_424971 320 32 of of IN 10_1101-2020_12_31_424971 320 33 [ [ -LRB- 10_1101-2020_12_31_424971 320 34 35S]-Met 35s]-met CD 10_1101-2020_12_31_424971 320 35 using use VBG 10_1101-2020_12_31_424971 320 36 the the DT 10_1101-2020_12_31_424971 320 37 PURExpress purexpress NN 10_1101-2020_12_31_424971 320 38 736 736 CD 10_1101-2020_12_31_424971 320 39 system system NN 10_1101-2020_12_31_424971 320 40 . . . 10_1101-2020_12_31_424971 321 1 SDS SDS NNP 10_1101-2020_12_31_424971 321 2 - - HYPH 10_1101-2020_12_31_424971 321 3 PAGE PAGE NNP 10_1101-2020_12_31_424971 321 4 analysis analysis NN 10_1101-2020_12_31_424971 321 5 was be VBD 10_1101-2020_12_31_424971 321 6 used use VBN 10_1101-2020_12_31_424971 321 7 to to TO 10_1101-2020_12_31_424971 321 8 detect detect VB 10_1101-2020_12_31_424971 321 9 [ [ -LRB- 10_1101-2020_12_31_424971 321 10 35S]-Met 35s]-met CD 10_1101-2020_12_31_424971 321 11 - - HYPH 10_1101-2020_12_31_424971 321 12 labeled label VBN 10_1101-2020_12_31_424971 321 13 polypeptides polypeptide NNS 10_1101-2020_12_31_424971 321 14 , , , 10_1101-2020_12_31_424971 321 15 which which WDT 10_1101-2020_12_31_424971 321 16 included include VBD 10_1101-2020_12_31_424971 321 17 737 737 CD 10_1101-2020_12_31_424971 321 18 the the DT 10_1101-2020_12_31_424971 321 19 full full JJ 10_1101-2020_12_31_424971 321 20 - - HYPH 10_1101-2020_12_31_424971 321 21 length length NN 10_1101-2020_12_31_424971 321 22 DHFR DHFR NNP 10_1101-2020_12_31_424971 321 23 , , , 10_1101-2020_12_31_424971 321 24 the the DT 10_1101-2020_12_31_424971 321 25 DC DC NNP 10_1101-2020_12_31_424971 321 26 fragment fragment NN 10_1101-2020_12_31_424971 321 27 , , , 10_1101-2020_12_31_424971 321 28 and and CC 10_1101-2020_12_31_424971 321 29 a a DT 10_1101-2020_12_31_424971 321 30 DN DN NNP 10_1101-2020_12_31_424971 321 31 fragment fragment NN 10_1101-2020_12_31_424971 321 32 that that WDT 10_1101-2020_12_31_424971 321 33 likely likely RB 10_1101-2020_12_31_424971 321 34 resulted result VBD 10_1101-2020_12_31_424971 321 35 from from IN 10_1101-2020_12_31_424971 321 36 initiation initiation NN 10_1101-2020_12_31_424971 321 37 of of IN 10_1101-2020_12_31_424971 321 38 738 738 CD 10_1101-2020_12_31_424971 321 39 translation translation NN 10_1101-2020_12_31_424971 321 40 at at IN 10_1101-2020_12_31_424971 321 41 a a DT 10_1101-2020_12_31_424971 321 42 cryptic cryptic JJ 10_1101-2020_12_31_424971 321 43 site site NN 10_1101-2020_12_31_424971 321 44 downstream downstream JJ 10_1101-2020_12_31_424971 321 45 from from IN 10_1101-2020_12_31_424971 321 46 the the DT 10_1101-2020_12_31_424971 321 47 CCC CCC NNP 10_1101-2020_12_31_424971 321 48 - - HYPH 10_1101-2020_12_31_424971 321 49 C C NNP 10_1101-2020_12_31_424971 321 50 motif motif NN 10_1101-2020_12_31_424971 321 51 ( ( -LRB- 10_1101-2020_12_31_424971 321 52 Figure Figure NNP 10_1101-2020_12_31_424971 321 53 2d 2d NNP 10_1101-2020_12_31_424971 321 54 ) ) -RRB- 10_1101-2020_12_31_424971 321 55 . . . 10_1101-2020_12_31_424971 322 1 The the DT 10_1101-2020_12_31_424971 322 2 fraction fraction NN 10_1101-2020_12_31_424971 322 3 of of IN 10_1101-2020_12_31_424971 322 4 the the DT 10_1101-2020_12_31_424971 322 5 full-739 full-739 NNP 10_1101-2020_12_31_424971 322 6 length length NN 10_1101-2020_12_31_424971 322 7 folA fola JJ 10_1101-2020_12_31_424971 322 8 gene gene NN 10_1101-2020_12_31_424971 322 9 product product NN 10_1101-2020_12_31_424971 322 10 , , , 10_1101-2020_12_31_424971 322 11 the the DT 10_1101-2020_12_31_424971 322 12 DC DC NNP 10_1101-2020_12_31_424971 322 13 fragment fragment NN 10_1101-2020_12_31_424971 322 14 , , , 10_1101-2020_12_31_424971 322 15 and and CC 10_1101-2020_12_31_424971 322 16 the the DT 10_1101-2020_12_31_424971 322 17 DN DN NNP 10_1101-2020_12_31_424971 322 18 fragment fragment NN 10_1101-2020_12_31_424971 322 19 was be VBD 10_1101-2020_12_31_424971 322 20 calculated calculate VBN 10_1101-2020_12_31_424971 322 21 from from IN 10_1101-2020_12_31_424971 322 22 the the DT 10_1101-2020_12_31_424971 322 23 amount amount NN 10_1101-2020_12_31_424971 322 24 740 740 CD 10_1101-2020_12_31_424971 322 25 of of IN 10_1101-2020_12_31_424971 322 26 each each DT 10_1101-2020_12_31_424971 322 27 in in IN 10_1101-2020_12_31_424971 322 28 the the DT 10_1101-2020_12_31_424971 322 29 sum sum NN 10_1101-2020_12_31_424971 322 30 of of IN 10_1101-2020_12_31_424971 322 31 all all DT 10_1101-2020_12_31_424971 322 32 three three CD 10_1101-2020_12_31_424971 322 33 products product NNS 10_1101-2020_12_31_424971 322 34 . . . 10_1101-2020_12_31_424971 323 1 We -PRON- PRP 10_1101-2020_12_31_424971 323 2 attribute attribute VBP 10_1101-2020_12_31_424971 323 3 the the DT 10_1101-2020_12_31_424971 323 4 overall overall JJ 10_1101-2020_12_31_424971 323 5 low low JJ 10_1101-2020_12_31_424971 323 6 recoding recode VBG 10_1101-2020_12_31_424971 323 7 efficiency efficiency NN 10_1101-2020_12_31_424971 323 8 ( ( -LRB- 10_1101-2020_12_31_424971 323 9 0.5 0.5 CD 10_1101-2020_12_31_424971 323 10 – – : 10_1101-2020_12_31_424971 323 11 741 741 CD 10_1101-2020_12_31_424971 323 12 5.0 5.0 CD 10_1101-2020_12_31_424971 323 13 % % NN 10_1101-2020_12_31_424971 323 14 ) ) -RRB- 10_1101-2020_12_31_424971 323 15 as as IN 10_1101-2020_12_31_424971 323 16 arising arise VBG 10_1101-2020_12_31_424971 323 17 from from IN 10_1101-2020_12_31_424971 323 18 a a DT 10_1101-2020_12_31_424971 323 19 combination combination NN 10_1101-2020_12_31_424971 323 20 of of IN 10_1101-2020_12_31_424971 323 21 the the DT 10_1101-2020_12_31_424971 323 22 rapid rapid JJ 10_1101-2020_12_31_424971 323 23 hydrolysis hydrolysis NN 10_1101-2020_12_31_424971 323 24 of of IN 10_1101-2020_12_31_424971 323 25 the the DT 10_1101-2020_12_31_424971 323 26 prolyl prolyl NNP 10_1101-2020_12_31_424971 323 27 linkage linkage NN 10_1101-2020_12_31_424971 323 28 , , , 10_1101-2020_12_31_424971 323 29 which which WDT 10_1101-2020_12_31_424971 323 30 is be VBZ 10_1101-2020_12_31_424971 323 31 the the DT 10_1101-2020_12_31_424971 323 32 least least JJS 10_1101-2020_12_31_424971 323 33 742 742 CD 10_1101-2020_12_31_424971 323 34 stable stable JJ 10_1101-2020_12_31_424971 323 35 among among IN 10_1101-2020_12_31_424971 323 36 aminoacyl aminoacyl NNP 10_1101-2020_12_31_424971 323 37 linkages63 linkages63 NNP 10_1101-2020_12_31_424971 323 38 , , , 10_1101-2020_12_31_424971 323 39 and and CC 10_1101-2020_12_31_424971 323 40 the the DT 10_1101-2020_12_31_424971 323 41 lack lack NN 10_1101-2020_12_31_424971 323 42 of of IN 10_1101-2020_12_31_424971 323 43 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 323 44 re re NN 10_1101-2020_12_31_424971 323 45 - - NN 10_1101-2020_12_31_424971 323 46 acylation acylation NN 10_1101-2020_12_31_424971 323 47 in in IN 10_1101-2020_12_31_424971 323 48 the the DT 10_1101-2020_12_31_424971 323 49 PURExpress purexpress NN 10_1101-2020_12_31_424971 323 50 system system NN 10_1101-2020_12_31_424971 323 51 . . . 10_1101-2020_12_31_424971 324 1 743 743 CD 10_1101-2020_12_31_424971 324 2 In in IN 10_1101-2020_12_31_424971 324 3 these these DT 10_1101-2020_12_31_424971 324 4 assays assay NNS 10_1101-2020_12_31_424971 324 5 , , , 10_1101-2020_12_31_424971 324 6 each each DT 10_1101-2020_12_31_424971 324 7 tRNA trna NN 10_1101-2020_12_31_424971 324 8 was be VBD 10_1101-2020_12_31_424971 324 9 tested test VBN 10_1101-2020_12_31_424971 324 10 in in IN 10_1101-2020_12_31_424971 324 11 the the DT 10_1101-2020_12_31_424971 324 12 G37-state G37-state NNP 10_1101-2020_12_31_424971 324 13 and and CC 10_1101-2020_12_31_424971 324 14 each each DT 10_1101-2020_12_31_424971 324 15 was be VBD 10_1101-2020_12_31_424971 324 16 normalized normalize VBN 10_1101-2020_12_31_424971 324 17 by by IN 10_1101-2020_12_31_424971 324 18 the the DT 10_1101-2020_12_31_424971 324 19 744 744 CD 10_1101-2020_12_31_424971 324 20 flexizyme flexizyme NNS 10_1101-2020_12_31_424971 324 21 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 324 22 efficiency efficiency NN 10_1101-2020_12_31_424971 324 23 , , , 10_1101-2020_12_31_424971 324 24 which which WDT 10_1101-2020_12_31_424971 324 25 was be VBD 10_1101-2020_12_31_424971 324 26 ~30 ~30 . 10_1101-2020_12_31_424971 324 27 % % NN 10_1101-2020_12_31_424971 324 28 for for IN 10_1101-2020_12_31_424971 324 29 Pro pro JJ 10_1101-2020_12_31_424971 324 30 and and CC 10_1101-2020_12_31_424971 324 31 Pro pro JJ 10_1101-2020_12_31_424971 324 32 analogues analogue NNS 10_1101-2020_12_31_424971 324 33 . . . 10_1101-2020_12_31_424971 325 1 The the DT 10_1101-2020_12_31_424971 325 2 745 745 CD 10_1101-2020_12_31_424971 325 3 PURExpress purexpress NN 10_1101-2020_12_31_424971 325 4 contained contain VBD 10_1101-2020_12_31_424971 325 5 all all DT 10_1101-2020_12_31_424971 325 6 natural natural JJ 10_1101-2020_12_31_424971 325 7 E. E. NNP 10_1101-2020_12_31_424971 325 8 coli coli NNS 10_1101-2020_12_31_424971 325 9 tRNAs tRNAs NNP 10_1101-2020_12_31_424971 325 10 , , , 10_1101-2020_12_31_424971 325 11 such such JJ 10_1101-2020_12_31_424971 325 12 that that IN 10_1101-2020_12_31_424971 325 13 the the DT 10_1101-2020_12_31_424971 325 14 CCC CCC NNP 10_1101-2020_12_31_424971 325 15 - - HYPH 10_1101-2020_12_31_424971 325 16 C C NNP 10_1101-2020_12_31_424971 325 17 codon codon NN 10_1101-2020_12_31_424971 325 18 motif motif NN 10_1101-2020_12_31_424971 325 19 would would MD 10_1101-2020_12_31_424971 325 20 not not RB 10_1101-2020_12_31_424971 325 21 have have VB 10_1101-2020_12_31_424971 325 22 746 746 CD 10_1101-2020_12_31_424971 325 23 a a DT 10_1101-2020_12_31_424971 325 24 chance chance NN 10_1101-2020_12_31_424971 325 25 of of IN 10_1101-2020_12_31_424971 325 26 vacancy vacancy NN 10_1101-2020_12_31_424971 325 27 even even RB 10_1101-2020_12_31_424971 325 28 when when WRB 10_1101-2020_12_31_424971 325 29 a a DT 10_1101-2020_12_31_424971 325 30 specific specific JJ 10_1101-2020_12_31_424971 325 31 CCC ccc NN 10_1101-2020_12_31_424971 325 32 - - HYPH 10_1101-2020_12_31_424971 325 33 reading read VBG 10_1101-2020_12_31_424971 325 34 tRNA trna NN 10_1101-2020_12_31_424971 325 35 was be VBD 10_1101-2020_12_31_424971 325 36 absent absent JJ 10_1101-2020_12_31_424971 325 37 . . . 10_1101-2020_12_31_424971 326 1 747 747 CD 10_1101-2020_12_31_424971 326 2 748 748 CD 10_1101-2020_12_31_424971 326 3 Rapid Rapid NNP 10_1101-2020_12_31_424971 326 4 kinetic kinetic NN 10_1101-2020_12_31_424971 326 5 GTPase GTPase NNP 10_1101-2020_12_31_424971 326 6 assays assay NNS 10_1101-2020_12_31_424971 326 7 . . . 10_1101-2020_12_31_424971 327 1 Ensemble ensemble JJ 10_1101-2020_12_31_424971 327 2 GTPase GTPase NNP 10_1101-2020_12_31_424971 327 3 assays assay NNS 10_1101-2020_12_31_424971 327 4 were be VBD 10_1101-2020_12_31_424971 327 5 performed perform VBN 10_1101-2020_12_31_424971 327 6 using use VBG 10_1101-2020_12_31_424971 327 7 the the DT 10_1101-2020_12_31_424971 327 8 codon codon NN 10_1101-2020_12_31_424971 327 9 - - HYPH 10_1101-2020_12_31_424971 327 10 walk walk VB 10_1101-2020_12_31_424971 327 11 749 749 CD 10_1101-2020_12_31_424971 327 12 approach approach NN 10_1101-2020_12_31_424971 327 13 , , , 10_1101-2020_12_31_424971 327 14 in in IN 10_1101-2020_12_31_424971 327 15 which which WDT 10_1101-2020_12_31_424971 327 16 an an DT 10_1101-2020_12_31_424971 327 17 E. E. NNP 10_1101-2020_12_31_424971 327 18 coli coli NNS 10_1101-2020_12_31_424971 327 19 in in IN 10_1101-2020_12_31_424971 327 20 vitro vitro FW 10_1101-2020_12_31_424971 327 21 translation translation NN 10_1101-2020_12_31_424971 327 22 system system NN 10_1101-2020_12_31_424971 327 23 composed compose VBN 10_1101-2020_12_31_424971 327 24 of of IN 10_1101-2020_12_31_424971 327 25 purified purify VBN 10_1101-2020_12_31_424971 327 26 components component NNS 10_1101-2020_12_31_424971 327 27 is be VBZ 10_1101-2020_12_31_424971 327 28 750 750 CD 10_1101-2020_12_31_424971 327 29 supplemented supplement VBN 10_1101-2020_12_31_424971 327 30 with with IN 10_1101-2020_12_31_424971 327 31 the the DT 10_1101-2020_12_31_424971 327 32 requisite requisite JJ 10_1101-2020_12_31_424971 327 33 tRNAs trnas NN 10_1101-2020_12_31_424971 327 34 and and CC 10_1101-2020_12_31_424971 327 35 translation translation NN 10_1101-2020_12_31_424971 327 36 factors factor NNS 10_1101-2020_12_31_424971 327 37 to to TO 10_1101-2020_12_31_424971 327 38 interrogate interrogate VB 10_1101-2020_12_31_424971 327 39 individual individual JJ 10_1101-2020_12_31_424971 327 40 steps step NNS 10_1101-2020_12_31_424971 327 41 of of IN 10_1101-2020_12_31_424971 327 42 751 751 CD 10_1101-2020_12_31_424971 327 43 the the DT 10_1101-2020_12_31_424971 327 44 elongation elongation NN 10_1101-2020_12_31_424971 327 45 cycle cycle NN 10_1101-2020_12_31_424971 327 46 . . . 10_1101-2020_12_31_424971 328 1 Programmed program VBN 10_1101-2020_12_31_424971 328 2 with with IN 10_1101-2020_12_31_424971 328 3 a a DT 10_1101-2020_12_31_424971 328 4 previously previously RB 10_1101-2020_12_31_424971 328 5 validated validate VBN 10_1101-2020_12_31_424971 328 6 synthetic synthetic JJ 10_1101-2020_12_31_424971 328 7 AUG AUG NNP 10_1101-2020_12_31_424971 328 8 - - HYPH 10_1101-2020_12_31_424971 328 9 CCC CCC NNP 10_1101-2020_12_31_424971 328 10 - - HYPH 10_1101-2020_12_31_424971 328 11 CGU CGU NNP 10_1101-2020_12_31_424971 328 12 - - HYPH 10_1101-2020_12_31_424971 328 13 U u NN 10_1101-2020_12_31_424971 328 14 mRNA mrna NN 10_1101-2020_12_31_424971 328 15 752 752 CD 10_1101-2020_12_31_424971 328 16 .CC .CC NFP 10_1101-2020_12_31_424971 328 17 - - HYPH 10_1101-2020_12_31_424971 328 18 BY by IN 10_1101-2020_12_31_424971 328 19 - - HYPH 10_1101-2020_12_31_424971 328 20 NC NC NNP 10_1101-2020_12_31_424971 328 21 - - HYPH 10_1101-2020_12_31_424971 328 22 ND ND NNP 10_1101-2020_12_31_424971 328 23 4.0 4.0 CD 10_1101-2020_12_31_424971 328 24 International International NNP 10_1101-2020_12_31_424971 328 25 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 328 26 under under IN 10_1101-2020_12_31_424971 328 27 a a DT 10_1101-2020_12_31_424971 328 28 ( ( -LRB- 10_1101-2020_12_31_424971 328 29 which which WDT 10_1101-2020_12_31_424971 328 30 was be VBD 10_1101-2020_12_31_424971 328 31 not not RB 10_1101-2020_12_31_424971 328 32 certified certify VBN 10_1101-2020_12_31_424971 328 33 by by IN 10_1101-2020_12_31_424971 328 34 peer peer NN 10_1101-2020_12_31_424971 328 35 review review NN 10_1101-2020_12_31_424971 328 36 ) ) -RRB- 10_1101-2020_12_31_424971 328 37 is be VBZ 10_1101-2020_12_31_424971 328 38 the the DT 10_1101-2020_12_31_424971 328 39 author author NN 10_1101-2020_12_31_424971 328 40 / / SYM 10_1101-2020_12_31_424971 328 41 funder funder NN 10_1101-2020_12_31_424971 328 42 , , , 10_1101-2020_12_31_424971 328 43 who who WP 10_1101-2020_12_31_424971 328 44 has have VBZ 10_1101-2020_12_31_424971 328 45 granted grant VBN 10_1101-2020_12_31_424971 328 46 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 328 47 a a DT 10_1101-2020_12_31_424971 328 48 license license NN 10_1101-2020_12_31_424971 328 49 to to TO 10_1101-2020_12_31_424971 328 50 display display VB 10_1101-2020_12_31_424971 328 51 the the DT 10_1101-2020_12_31_424971 328 52 preprint preprint NN 10_1101-2020_12_31_424971 328 53 in in IN 10_1101-2020_12_31_424971 328 54 perpetuity perpetuity NN 10_1101-2020_12_31_424971 328 55 . . . 10_1101-2020_12_31_424971 329 1 It -PRON- PRP 10_1101-2020_12_31_424971 329 2 is be VBZ 10_1101-2020_12_31_424971 329 3 made make VBN 10_1101-2020_12_31_424971 329 4 The the DT 10_1101-2020_12_31_424971 329 5 copyright copyright NN 10_1101-2020_12_31_424971 329 6 holder holder NN 10_1101-2020_12_31_424971 329 7 for for IN 10_1101-2020_12_31_424971 329 8 this this DT 10_1101-2020_12_31_424971 329 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 329 10 version version NN 10_1101-2020_12_31_424971 329 11 posted post VBD 10_1101-2020_12_31_424971 329 12 January January NNP 10_1101-2020_12_31_424971 329 13 2 2 CD 10_1101-2020_12_31_424971 329 14 , , , 10_1101-2020_12_31_424971 329 15 2021 2021 CD 10_1101-2020_12_31_424971 329 16 . . . 10_1101-2020_12_31_424971 329 17 ; ; : 10_1101-2020_12_31_424971 329 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 329 19 : : : 10_1101-2020_12_31_424971 329 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 329 21 preprint preprint NN 10_1101-2020_12_31_424971 329 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 329 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 329 24 31 31 CD 10_1101-2020_12_31_424971 329 25 template29,34 template29,34 NNP 10_1101-2020_12_31_424971 329 26 , , , 10_1101-2020_12_31_424971 329 27 a a DT 10_1101-2020_12_31_424971 329 28 70S 70S NNP 10_1101-2020_12_31_424971 329 29 IC IC NNP 10_1101-2020_12_31_424971 329 30 was be VBD 10_1101-2020_12_31_424971 329 31 assembled assemble VBN 10_1101-2020_12_31_424971 329 32 that that IN 10_1101-2020_12_31_424971 329 33 positioned position VBD 10_1101-2020_12_31_424971 329 34 the the DT 10_1101-2020_12_31_424971 329 35 AUG AUG NNP 10_1101-2020_12_31_424971 329 36 start start VBP 10_1101-2020_12_31_424971 329 37 codon codon NNP 10_1101-2020_12_31_424971 329 38 and and CC 10_1101-2020_12_31_424971 329 39 an an DT 10_1101-2020_12_31_424971 329 40 initiator initiator NN 10_1101-2020_12_31_424971 329 41 fMet-753 fMet-753 , 10_1101-2020_12_31_424971 329 42 tRNAfMet trnafmet ADD 10_1101-2020_12_31_424971 329 43 at at IN 10_1101-2020_12_31_424971 329 44 the the DT 10_1101-2020_12_31_424971 329 45 P p NN 10_1101-2020_12_31_424971 329 46 site site NN 10_1101-2020_12_31_424971 329 47 and and CC 10_1101-2020_12_31_424971 329 48 the the DT 10_1101-2020_12_31_424971 329 49 CCC CCC NNP 10_1101-2020_12_31_424971 329 50 - - HYPH 10_1101-2020_12_31_424971 329 51 C C NNP 10_1101-2020_12_31_424971 329 52 motif motif NN 10_1101-2020_12_31_424971 329 53 at at IN 10_1101-2020_12_31_424971 329 54 the the DT 10_1101-2020_12_31_424971 329 55 A A NNP 10_1101-2020_12_31_424971 329 56 site site NN 10_1101-2020_12_31_424971 329 57 . . . 10_1101-2020_12_31_424971 330 1 Reactions reaction NNS 10_1101-2020_12_31_424971 330 2 to to TO 10_1101-2020_12_31_424971 330 3 monitor monitor VB 10_1101-2020_12_31_424971 330 4 the the DT 10_1101-2020_12_31_424971 330 5 EF EF NNP 10_1101-2020_12_31_424971 330 6 - - HYPH 10_1101-2020_12_31_424971 330 7 Tu-754 tu-754 CD 10_1101-2020_12_31_424971 330 8 dependent dependent JJ 10_1101-2020_12_31_424971 330 9 hydrolysis hydrolysis NN 10_1101-2020_12_31_424971 330 10 of of IN 10_1101-2020_12_31_424971 330 11 GTP GTP NNP 10_1101-2020_12_31_424971 330 12 during during IN 10_1101-2020_12_31_424971 330 13 delivery delivery NN 10_1101-2020_12_31_424971 330 14 and and CC 10_1101-2020_12_31_424971 330 15 accommodation accommodation NN 10_1101-2020_12_31_424971 330 16 of of IN 10_1101-2020_12_31_424971 330 17 a a DT 10_1101-2020_12_31_424971 330 18 TC tc NN 10_1101-2020_12_31_424971 330 19 to to IN 10_1101-2020_12_31_424971 330 20 the the DT 10_1101-2020_12_31_424971 330 21 A A NNP 10_1101-2020_12_31_424971 330 22 site site NN 10_1101-2020_12_31_424971 330 23 were be VBD 10_1101-2020_12_31_424971 330 24 755 755 CD 10_1101-2020_12_31_424971 330 25 conducted conduct VBN 10_1101-2020_12_31_424971 330 26 at at IN 10_1101-2020_12_31_424971 330 27 20 20 CD 10_1101-2020_12_31_424971 330 28 ° ° NNS 10_1101-2020_12_31_424971 330 29 C C NNP 10_1101-2020_12_31_424971 330 30 in in IN 10_1101-2020_12_31_424971 330 31 a a DT 10_1101-2020_12_31_424971 330 32 buffer buffer NN 10_1101-2020_12_31_424971 330 33 containing contain VBG 10_1101-2020_12_31_424971 330 34 50 50 CD 10_1101-2020_12_31_424971 330 35 mM mM NNP 10_1101-2020_12_31_424971 330 36 Tris Tris NNP 10_1101-2020_12_31_424971 330 37 - - HYPH 10_1101-2020_12_31_424971 330 38 HCl HCl NNP 10_1101-2020_12_31_424971 330 39 ( ( -LRB- 10_1101-2020_12_31_424971 330 40 pH pH NNP 10_1101-2020_12_31_424971 330 41 7.5 7.5 CD 10_1101-2020_12_31_424971 330 42 ) ) -RRB- 10_1101-2020_12_31_424971 330 43 , , , 10_1101-2020_12_31_424971 330 44 70 70 CD 10_1101-2020_12_31_424971 330 45 mM mm NN 10_1101-2020_12_31_424971 330 46 NH4Cl nh4cl CD 10_1101-2020_12_31_424971 330 47 , , , 10_1101-2020_12_31_424971 330 48 30 30 CD 10_1101-2020_12_31_424971 330 49 mM mm NN 10_1101-2020_12_31_424971 330 50 KCl KCl NNS 10_1101-2020_12_31_424971 330 51 , , , 10_1101-2020_12_31_424971 330 52 7 7 CD 10_1101-2020_12_31_424971 330 53 756 756 CD 10_1101-2020_12_31_424971 330 54 mM mM NNP 10_1101-2020_12_31_424971 330 55 MgCl2 mgcl2 NN 10_1101-2020_12_31_424971 330 56 , , , 10_1101-2020_12_31_424971 330 57 1 1 CD 10_1101-2020_12_31_424971 330 58 mM mM NNP 10_1101-2020_12_31_424971 330 59 DTT DTT NNP 10_1101-2020_12_31_424971 330 60 , , , 10_1101-2020_12_31_424971 330 61 and and CC 10_1101-2020_12_31_424971 330 62 0.5 0.5 CD 10_1101-2020_12_31_424971 330 63 mM mm NN 10_1101-2020_12_31_424971 330 64 spermidine29 spermidine29 CD 10_1101-2020_12_31_424971 330 65 . . . 10_1101-2020_12_31_424971 331 1 Each each DT 10_1101-2020_12_31_424971 331 2 TC TC NNP 10_1101-2020_12_31_424971 331 3 was be VBD 10_1101-2020_12_31_424971 331 4 formed form VBN 10_1101-2020_12_31_424971 331 5 by by IN 10_1101-2020_12_31_424971 331 6 incubating incubate VBG 10_1101-2020_12_31_424971 331 7 EF EF NNP 10_1101-2020_12_31_424971 331 8 - - HYPH 10_1101-2020_12_31_424971 331 9 Tu Tu NNP 10_1101-2020_12_31_424971 331 10 with with IN 10_1101-2020_12_31_424971 331 11 757 757 CD 10_1101-2020_12_31_424971 331 12 8 8 CD 10_1101-2020_12_31_424971 331 13 nM nM NNP 10_1101-2020_12_31_424971 331 14 [ [ -LRB- 10_1101-2020_12_31_424971 331 15 g-32P]-GTP g-32p]-gtp NN 10_1101-2020_12_31_424971 331 16 ( ( -LRB- 10_1101-2020_12_31_424971 331 17 6000 6000 CD 10_1101-2020_12_31_424971 331 18 Ci Ci NNP 10_1101-2020_12_31_424971 331 19 / / SYM 10_1101-2020_12_31_424971 331 20 mmole mmole NN 10_1101-2020_12_31_424971 331 21 ) ) -RRB- 10_1101-2020_12_31_424971 331 22 for for IN 10_1101-2020_12_31_424971 331 23 15 15 CD 10_1101-2020_12_31_424971 331 24 min min NN 10_1101-2020_12_31_424971 331 25 at at IN 10_1101-2020_12_31_424971 331 26 37 37 CD 10_1101-2020_12_31_424971 331 27 ºC ºC NNP 10_1101-2020_12_31_424971 331 28 , , , 10_1101-2020_12_31_424971 331 29 after after IN 10_1101-2020_12_31_424971 331 30 which which WDT 10_1101-2020_12_31_424971 331 31 aminoacylated aminoacylate VBD 10_1101-2020_12_31_424971 331 32 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 331 33 or or CC 10_1101-2020_12_31_424971 331 34 ProL ProL NNP 10_1101-2020_12_31_424971 331 35 758 758 CD 10_1101-2020_12_31_424971 331 36 was be VBD 10_1101-2020_12_31_424971 331 37 added add VBN 10_1101-2020_12_31_424971 331 38 and and CC 10_1101-2020_12_31_424971 331 39 the the DT 10_1101-2020_12_31_424971 331 40 incubation incubation NN 10_1101-2020_12_31_424971 331 41 continued continue VBD 10_1101-2020_12_31_424971 331 42 for for IN 10_1101-2020_12_31_424971 331 43 15 15 CD 10_1101-2020_12_31_424971 331 44 min min NN 10_1101-2020_12_31_424971 331 45 at at IN 10_1101-2020_12_31_424971 331 46 4 4 CD 10_1101-2020_12_31_424971 331 47 ºC. ºC. NNS 10_1101-2020_12_31_424971 332 1 Unbound Unbound NNP 10_1101-2020_12_31_424971 332 2 [ [ -LRB- 10_1101-2020_12_31_424971 332 3 g-32P]-GTP g-32P]-GTP NNP 10_1101-2020_12_31_424971 332 4 was be VBD 10_1101-2020_12_31_424971 332 5 removed remove VBN 10_1101-2020_12_31_424971 332 6 759 759 CD 10_1101-2020_12_31_424971 332 7 from from IN 10_1101-2020_12_31_424971 332 8 the the DT 10_1101-2020_12_31_424971 332 9 TC TC NNP 10_1101-2020_12_31_424971 332 10 solution solution NN 10_1101-2020_12_31_424971 332 11 by by IN 10_1101-2020_12_31_424971 332 12 gel gel NN 10_1101-2020_12_31_424971 332 13 filtration filtration NN 10_1101-2020_12_31_424971 332 14 through through IN 10_1101-2020_12_31_424971 332 15 a a DT 10_1101-2020_12_31_424971 332 16 spin spin NN 10_1101-2020_12_31_424971 332 17 cartridge cartridge NN 10_1101-2020_12_31_424971 332 18 ( ( -LRB- 10_1101-2020_12_31_424971 332 19 CentriSpin-20 CentriSpin-20 NNP 10_1101-2020_12_31_424971 332 20 ; ; : 10_1101-2020_12_31_424971 332 21 Princeton Princeton NNP 10_1101-2020_12_31_424971 332 22 760 760 CD 10_1101-2020_12_31_424971 332 23 Separations Separations NNPS 10_1101-2020_12_31_424971 332 24 ) ) -RRB- 10_1101-2020_12_31_424971 332 25 . . . 10_1101-2020_12_31_424971 333 1 Equal equal JJ 10_1101-2020_12_31_424971 333 2 volumes volume NNS 10_1101-2020_12_31_424971 333 3 of of IN 10_1101-2020_12_31_424971 333 4 each each DT 10_1101-2020_12_31_424971 333 5 purified purify VBN 10_1101-2020_12_31_424971 333 6 TC TC NNP 10_1101-2020_12_31_424971 333 7 and and CC 10_1101-2020_12_31_424971 333 8 a a DT 10_1101-2020_12_31_424971 333 9 solution solution NN 10_1101-2020_12_31_424971 333 10 of of IN 10_1101-2020_12_31_424971 333 11 70S 70S NNP 10_1101-2020_12_31_424971 333 12 ICs ic NNS 10_1101-2020_12_31_424971 333 13 were be VBD 10_1101-2020_12_31_424971 333 14 rapidly rapidly RB 10_1101-2020_12_31_424971 333 15 mixed mix VBN 10_1101-2020_12_31_424971 333 16 in in IN 10_1101-2020_12_31_424971 333 17 761 761 CD 10_1101-2020_12_31_424971 333 18 the the DT 10_1101-2020_12_31_424971 333 19 RQF-3 RQF-3 NNP 10_1101-2020_12_31_424971 333 20 Kintek Kintek NNPS 10_1101-2020_12_31_424971 333 21 chemical chemical NN 10_1101-2020_12_31_424971 333 22 quench quench NN 10_1101-2020_12_31_424971 333 23 apparatus29 apparatus29 NNP 10_1101-2020_12_31_424971 333 24 . . . 10_1101-2020_12_31_424971 334 1 Final final JJ 10_1101-2020_12_31_424971 334 2 concentrations concentration NNS 10_1101-2020_12_31_424971 334 3 in in IN 10_1101-2020_12_31_424971 334 4 these these DT 10_1101-2020_12_31_424971 334 5 reactions reaction NNS 10_1101-2020_12_31_424971 334 6 were be VBD 10_1101-2020_12_31_424971 334 7 0.5 0.5 CD 10_1101-2020_12_31_424971 334 8 762 762 CD 10_1101-2020_12_31_424971 334 9 µM µm NN 10_1101-2020_12_31_424971 334 10 for for IN 10_1101-2020_12_31_424971 334 11 the the DT 10_1101-2020_12_31_424971 334 12 70S 70S NNP 10_1101-2020_12_31_424971 334 13 IC IC NNP 10_1101-2020_12_31_424971 334 14 ; ; : 10_1101-2020_12_31_424971 334 15 0.8 0.8 CD 10_1101-2020_12_31_424971 334 16 µM µm NN 10_1101-2020_12_31_424971 334 17 for for IN 10_1101-2020_12_31_424971 334 18 mRNA mRNA NNP 10_1101-2020_12_31_424971 334 19 ; ; : 10_1101-2020_12_31_424971 334 20 0.65 0.65 CD 10_1101-2020_12_31_424971 334 21 µM µM NNP 10_1101-2020_12_31_424971 334 22 each each DT 10_1101-2020_12_31_424971 334 23 for for IN 10_1101-2020_12_31_424971 334 24 IFs IFs NNP 10_1101-2020_12_31_424971 334 25 1 1 CD 10_1101-2020_12_31_424971 334 26 , , , 10_1101-2020_12_31_424971 334 27 2 2 CD 10_1101-2020_12_31_424971 334 28 , , , 10_1101-2020_12_31_424971 334 29 and and CC 10_1101-2020_12_31_424971 334 30 3 3 CD 10_1101-2020_12_31_424971 334 31 ; ; : 10_1101-2020_12_31_424971 334 32 0.65 0.65 CD 10_1101-2020_12_31_424971 334 33 µM µM NNP 10_1101-2020_12_31_424971 334 34 for for IN 10_1101-2020_12_31_424971 334 35 fMet fMet NNP 10_1101-2020_12_31_424971 334 36 - - HYPH 10_1101-2020_12_31_424971 334 37 tRNAMet tRNAMet NNP 10_1101-2020_12_31_424971 334 38 ; ; : 10_1101-2020_12_31_424971 334 39 763 763 CD 10_1101-2020_12_31_424971 334 40 1.8 1.8 CD 10_1101-2020_12_31_424971 334 41 µM µm NN 10_1101-2020_12_31_424971 334 42 for for IN 10_1101-2020_12_31_424971 334 43 EF EF NNP 10_1101-2020_12_31_424971 334 44 - - HYPH 10_1101-2020_12_31_424971 334 45 Tu Tu NNP 10_1101-2020_12_31_424971 334 46 ; ; : 10_1101-2020_12_31_424971 334 47 0.4 0.4 CD 10_1101-2020_12_31_424971 334 48 µM µM NNP 10_1101-2020_12_31_424971 334 49 for for IN 10_1101-2020_12_31_424971 334 50 aminoacylated aminoacylate VBN 10_1101-2020_12_31_424971 334 51 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 334 52 or or CC 10_1101-2020_12_31_424971 334 53 ProL ProL NNP 10_1101-2020_12_31_424971 334 54 ; ; : 10_1101-2020_12_31_424971 334 55 and and CC 10_1101-2020_12_31_424971 334 56 0.5 0.5 CD 10_1101-2020_12_31_424971 334 57 mM mm NN 10_1101-2020_12_31_424971 334 58 for for IN 10_1101-2020_12_31_424971 334 59 cold cold JJ 10_1101-2020_12_31_424971 334 60 GTP GTP NNP 10_1101-2020_12_31_424971 334 61 . . . 10_1101-2020_12_31_424971 335 1 The the DT 10_1101-2020_12_31_424971 335 2 yield yield NN 10_1101-2020_12_31_424971 335 3 764 764 CD 10_1101-2020_12_31_424971 335 4 of of IN 10_1101-2020_12_31_424971 335 5 GTP GTP NNP 10_1101-2020_12_31_424971 335 6 hydrolysis hydrolysis NN 10_1101-2020_12_31_424971 335 7 and and CC 10_1101-2020_12_31_424971 335 8 kGTP kgtp NN 10_1101-2020_12_31_424971 335 9 , , , 10_1101-2020_12_31_424971 335 10 obs obs NN 10_1101-2020_12_31_424971 335 11 upon upon IN 10_1101-2020_12_31_424971 335 12 rapid rapid JJ 10_1101-2020_12_31_424971 335 13 mixing mixing NN 10_1101-2020_12_31_424971 335 14 of of IN 10_1101-2020_12_31_424971 335 15 each each DT 10_1101-2020_12_31_424971 335 16 TC TC NNP 10_1101-2020_12_31_424971 335 17 with with IN 10_1101-2020_12_31_424971 335 18 excess excess JJ 10_1101-2020_12_31_424971 335 19 70S 70S NNP 10_1101-2020_12_31_424971 335 20 ICs ic NNS 10_1101-2020_12_31_424971 335 21 were be VBD 10_1101-2020_12_31_424971 335 22 measured measure VBN 10_1101-2020_12_31_424971 335 23 765 765 CD 10_1101-2020_12_31_424971 335 24 by by IN 10_1101-2020_12_31_424971 335 25 removing remove VBG 10_1101-2020_12_31_424971 335 26 aliquots aliquot NNS 10_1101-2020_12_31_424971 335 27 of of IN 10_1101-2020_12_31_424971 335 28 the the DT 10_1101-2020_12_31_424971 335 29 reaction reaction NN 10_1101-2020_12_31_424971 335 30 at at IN 10_1101-2020_12_31_424971 335 31 defined define VBN 10_1101-2020_12_31_424971 335 32 time time NN 10_1101-2020_12_31_424971 335 33 points point NNS 10_1101-2020_12_31_424971 335 34 , , , 10_1101-2020_12_31_424971 335 35 quenching quench VBG 10_1101-2020_12_31_424971 335 36 the the DT 10_1101-2020_12_31_424971 335 37 aliquots aliquot NNS 10_1101-2020_12_31_424971 335 38 with with IN 10_1101-2020_12_31_424971 335 39 40 40 CD 10_1101-2020_12_31_424971 335 40 % % NN 10_1101-2020_12_31_424971 335 41 formic formic JJ 10_1101-2020_12_31_424971 335 42 766 766 CD 10_1101-2020_12_31_424971 335 43 acid acid NN 10_1101-2020_12_31_424971 335 44 , , , 10_1101-2020_12_31_424971 335 45 separating separate VBG 10_1101-2020_12_31_424971 335 46 [ [ -LRB- 10_1101-2020_12_31_424971 335 47 g-32P g-32p NN 10_1101-2020_12_31_424971 335 48 ] ] -RRB- 10_1101-2020_12_31_424971 335 49 from from IN 10_1101-2020_12_31_424971 335 50 [ [ -LRB- 10_1101-2020_12_31_424971 335 51 g-32P]-GTP g-32p]-gtp NN 10_1101-2020_12_31_424971 335 52 using use VBG 10_1101-2020_12_31_424971 335 53 thin thin JJ 10_1101-2020_12_31_424971 335 54 layer layer NN 10_1101-2020_12_31_424971 335 55 chromatography chromatography NN 10_1101-2020_12_31_424971 335 56 ( ( -LRB- 10_1101-2020_12_31_424971 335 57 TLC TLC NNP 10_1101-2020_12_31_424971 335 58 ) ) -RRB- 10_1101-2020_12_31_424971 335 59 , , , 10_1101-2020_12_31_424971 335 60 and and CC 10_1101-2020_12_31_424971 335 61 quantifying quantify VBG 10_1101-2020_12_31_424971 335 62 767 767 CD 10_1101-2020_12_31_424971 335 63 the the DT 10_1101-2020_12_31_424971 335 64 amount amount NN 10_1101-2020_12_31_424971 335 65 of of IN 10_1101-2020_12_31_424971 335 66 each each DT 10_1101-2020_12_31_424971 335 67 as as IN 10_1101-2020_12_31_424971 335 68 a a DT 10_1101-2020_12_31_424971 335 69 function function NN 10_1101-2020_12_31_424971 335 70 of of IN 10_1101-2020_12_31_424971 335 71 time time NN 10_1101-2020_12_31_424971 335 72 using use VBG 10_1101-2020_12_31_424971 335 73 phosphorimaging29 phosphorimaging29 NNP 10_1101-2020_12_31_424971 335 74 . . . 10_1101-2020_12_31_424971 336 1 We -PRON- PRP 10_1101-2020_12_31_424971 336 2 adjusted adjust VBD 10_1101-2020_12_31_424971 336 3 reaction reaction NN 10_1101-2020_12_31_424971 336 4 768 768 CD 10_1101-2020_12_31_424971 336 5 conditions condition NNS 10_1101-2020_12_31_424971 336 6 such such JJ 10_1101-2020_12_31_424971 336 7 that that IN 10_1101-2020_12_31_424971 336 8 the the DT 10_1101-2020_12_31_424971 336 9 kGTP kgtp NN 10_1101-2020_12_31_424971 336 10 , , , 10_1101-2020_12_31_424971 336 11 obs obs NNP 10_1101-2020_12_31_424971 336 12 increased increase VBD 10_1101-2020_12_31_424971 336 13 linearly linearly RB 10_1101-2020_12_31_424971 336 14 as as IN 10_1101-2020_12_31_424971 336 15 a a DT 10_1101-2020_12_31_424971 336 16 function function NN 10_1101-2020_12_31_424971 336 17 of of IN 10_1101-2020_12_31_424971 336 18 70S 70S NNP 10_1101-2020_12_31_424971 336 19 IC IC NNP 10_1101-2020_12_31_424971 336 20 concentration concentration NN 10_1101-2020_12_31_424971 336 21 . . . 10_1101-2020_12_31_424971 337 1 769 769 CD 10_1101-2020_12_31_424971 337 2 770 770 CD 10_1101-2020_12_31_424971 337 3 Rapid Rapid NNP 10_1101-2020_12_31_424971 337 4 kinetic kinetic NN 10_1101-2020_12_31_424971 337 5 di- di- XX 10_1101-2020_12_31_424971 337 6 and and CC 10_1101-2020_12_31_424971 337 7 tripeptide tripeptide NN 10_1101-2020_12_31_424971 337 8 formation formation NN 10_1101-2020_12_31_424971 337 9 assays assays RB 10_1101-2020_12_31_424971 337 10 . . . 10_1101-2020_12_31_424971 338 1 Di- di- RB 10_1101-2020_12_31_424971 338 2 and and CC 10_1101-2020_12_31_424971 338 3 tripeptide tripeptide NN 10_1101-2020_12_31_424971 338 4 formation formation NN 10_1101-2020_12_31_424971 338 5 assays assays RB 10_1101-2020_12_31_424971 338 6 were be VBD 10_1101-2020_12_31_424971 338 7 771 771 CD 10_1101-2020_12_31_424971 338 8 performed perform VBN 10_1101-2020_12_31_424971 338 9 using use VBG 10_1101-2020_12_31_424971 338 10 the the DT 10_1101-2020_12_31_424971 338 11 codon codon NN 10_1101-2020_12_31_424971 338 12 - - HYPH 10_1101-2020_12_31_424971 338 13 walk walk NN 10_1101-2020_12_31_424971 338 14 approach approach NN 10_1101-2020_12_31_424971 338 15 described describe VBN 10_1101-2020_12_31_424971 338 16 above above RB 10_1101-2020_12_31_424971 338 17 in in IN 10_1101-2020_12_31_424971 338 18 50 50 CD 10_1101-2020_12_31_424971 338 19 mM mM NNP 10_1101-2020_12_31_424971 338 20 Tris Tris NNP 10_1101-2020_12_31_424971 338 21 - - HYPH 10_1101-2020_12_31_424971 338 22 HCl HCl NNP 10_1101-2020_12_31_424971 338 23 ( ( -LRB- 10_1101-2020_12_31_424971 338 24 pH pH NNP 10_1101-2020_12_31_424971 338 25 7.5 7.5 CD 10_1101-2020_12_31_424971 338 26 ) ) -RRB- 10_1101-2020_12_31_424971 338 27 , , , 10_1101-2020_12_31_424971 338 28 70 70 CD 10_1101-2020_12_31_424971 338 29 mM mM NNP 10_1101-2020_12_31_424971 338 30 772 772 CD 10_1101-2020_12_31_424971 338 31 NH4Cl nh4cl CD 10_1101-2020_12_31_424971 338 32 , , , 10_1101-2020_12_31_424971 338 33 30 30 CD 10_1101-2020_12_31_424971 338 34 mM mm NN 10_1101-2020_12_31_424971 338 35 KCl KCl NNS 10_1101-2020_12_31_424971 338 36 , , , 10_1101-2020_12_31_424971 338 37 3.5 3.5 CD 10_1101-2020_12_31_424971 338 38 mM mM NNP 10_1101-2020_12_31_424971 338 39 MgCl2 mgcl2 NN 10_1101-2020_12_31_424971 338 40 , , , 10_1101-2020_12_31_424971 338 41 1 1 CD 10_1101-2020_12_31_424971 338 42 mM mM NNP 10_1101-2020_12_31_424971 338 43 DTT DTT NNP 10_1101-2020_12_31_424971 338 44 , , , 10_1101-2020_12_31_424971 338 45 0.5 0.5 CD 10_1101-2020_12_31_424971 338 46 mM mm NN 10_1101-2020_12_31_424971 338 47 spermidine spermidine NN 10_1101-2020_12_31_424971 338 48 , , , 10_1101-2020_12_31_424971 338 49 at at IN 10_1101-2020_12_31_424971 338 50 20 20 CD 10_1101-2020_12_31_424971 338 51 ° ° NNS 10_1101-2020_12_31_424971 338 52 C C NNP 10_1101-2020_12_31_424971 338 53 unless unless IN 10_1101-2020_12_31_424971 338 54 otherwise otherwise RB 10_1101-2020_12_31_424971 338 55 773 773 CD 10_1101-2020_12_31_424971 338 56 indicated29 indicated29 NNPS 10_1101-2020_12_31_424971 338 57 . . . 10_1101-2020_12_31_424971 339 1 70S 70s CD 10_1101-2020_12_31_424971 339 2 ICs ic NNS 10_1101-2020_12_31_424971 339 3 were be VBD 10_1101-2020_12_31_424971 339 4 formed form VBN 10_1101-2020_12_31_424971 339 5 by by IN 10_1101-2020_12_31_424971 339 6 incubating incubate VBG 10_1101-2020_12_31_424971 339 7 70S 70S NNP 10_1101-2020_12_31_424971 339 8 ribosomes ribosome NNS 10_1101-2020_12_31_424971 339 9 , , , 10_1101-2020_12_31_424971 339 10 mRNA mRNA NNS 10_1101-2020_12_31_424971 339 11 , , , 10_1101-2020_12_31_424971 339 12 [ [ -LRB- 10_1101-2020_12_31_424971 339 13 35S]-fMet 35s]-fmet CD 10_1101-2020_12_31_424971 339 14 - - HYPH 10_1101-2020_12_31_424971 339 15 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 339 16 , , , 10_1101-2020_12_31_424971 339 17 and and CC 10_1101-2020_12_31_424971 339 18 774 774 CD 10_1101-2020_12_31_424971 339 19 IFs if NNS 10_1101-2020_12_31_424971 339 20 1 1 CD 10_1101-2020_12_31_424971 339 21 , , , 10_1101-2020_12_31_424971 339 22 2 2 CD 10_1101-2020_12_31_424971 339 23 , , , 10_1101-2020_12_31_424971 339 24 and and CC 10_1101-2020_12_31_424971 339 25 3 3 CD 10_1101-2020_12_31_424971 339 26 , , , 10_1101-2020_12_31_424971 339 27 and and CC 10_1101-2020_12_31_424971 339 28 GTP GTP NNP 10_1101-2020_12_31_424971 339 29 , , , 10_1101-2020_12_31_424971 339 30 for for IN 10_1101-2020_12_31_424971 339 31 25 25 CD 10_1101-2020_12_31_424971 339 32 min min NN 10_1101-2020_12_31_424971 339 33 at at IN 10_1101-2020_12_31_424971 339 34 37 37 CD 10_1101-2020_12_31_424971 339 35 ° ° NNS 10_1101-2020_12_31_424971 339 36 C C NNP 10_1101-2020_12_31_424971 339 37 in in IN 10_1101-2020_12_31_424971 339 38 the the DT 10_1101-2020_12_31_424971 339 39 reaction reaction NN 10_1101-2020_12_31_424971 339 40 buffer buffer NN 10_1101-2020_12_31_424971 339 41 . . . 10_1101-2020_12_31_424971 340 1 Separately separately RB 10_1101-2020_12_31_424971 340 2 , , , 10_1101-2020_12_31_424971 340 3 TCs tc NNS 10_1101-2020_12_31_424971 340 4 were be VBD 10_1101-2020_12_31_424971 340 5 formed form VBN 10_1101-2020_12_31_424971 340 6 775 775 CD 10_1101-2020_12_31_424971 340 7 in in IN 10_1101-2020_12_31_424971 340 8 the the DT 10_1101-2020_12_31_424971 340 9 reaction reaction NN 10_1101-2020_12_31_424971 340 10 buffer buffer NN 10_1101-2020_12_31_424971 340 11 by by IN 10_1101-2020_12_31_424971 340 12 incubating incubate VBG 10_1101-2020_12_31_424971 340 13 EF EF NNP 10_1101-2020_12_31_424971 340 14 - - HYPH 10_1101-2020_12_31_424971 340 15 Tu Tu NNP 10_1101-2020_12_31_424971 340 16 and and CC 10_1101-2020_12_31_424971 340 17 GTP GTP NNP 10_1101-2020_12_31_424971 340 18 for for IN 10_1101-2020_12_31_424971 340 19 15 15 CD 10_1101-2020_12_31_424971 340 20 min min NN 10_1101-2020_12_31_424971 340 21 at at IN 10_1101-2020_12_31_424971 340 22 37 37 CD 10_1101-2020_12_31_424971 340 23 ° ° NNS 10_1101-2020_12_31_424971 340 24 C C NNP 10_1101-2020_12_31_424971 340 25 followed follow VBN 10_1101-2020_12_31_424971 340 26 by by IN 10_1101-2020_12_31_424971 340 27 adding add VBG 10_1101-2020_12_31_424971 340 28 the the DT 10_1101-2020_12_31_424971 340 29 776 776 CD 10_1101-2020_12_31_424971 340 30 requisite requisite JJ 10_1101-2020_12_31_424971 340 31 aa aa NNP 10_1101-2020_12_31_424971 340 32 - - HYPH 10_1101-2020_12_31_424971 340 33 tRNAs tRNAs NNP 10_1101-2020_12_31_424971 340 34 and and CC 10_1101-2020_12_31_424971 340 35 incubating incubate VBG 10_1101-2020_12_31_424971 340 36 in in IN 10_1101-2020_12_31_424971 340 37 an an DT 10_1101-2020_12_31_424971 340 38 ice ice NN 10_1101-2020_12_31_424971 340 39 bath bath NN 10_1101-2020_12_31_424971 340 40 for for IN 10_1101-2020_12_31_424971 340 41 15 15 CD 10_1101-2020_12_31_424971 340 42 min min NN 10_1101-2020_12_31_424971 340 43 . . . 10_1101-2020_12_31_424971 341 1 In in IN 10_1101-2020_12_31_424971 341 2 dipeptide dipeptide NN 10_1101-2020_12_31_424971 341 3 formation formation NN 10_1101-2020_12_31_424971 341 4 assays assays RB 10_1101-2020_12_31_424971 341 5 , , , 10_1101-2020_12_31_424971 341 6 70S 70S NNP 10_1101-2020_12_31_424971 341 7 777 777 CD 10_1101-2020_12_31_424971 341 8 .CC .CC : 10_1101-2020_12_31_424971 341 9 - - HYPH 10_1101-2020_12_31_424971 341 10 BY by IN 10_1101-2020_12_31_424971 341 11 - - HYPH 10_1101-2020_12_31_424971 341 12 NC NC NNP 10_1101-2020_12_31_424971 341 13 - - HYPH 10_1101-2020_12_31_424971 341 14 ND ND NNP 10_1101-2020_12_31_424971 341 15 4.0 4.0 CD 10_1101-2020_12_31_424971 341 16 International International NNP 10_1101-2020_12_31_424971 341 17 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 341 18 under under IN 10_1101-2020_12_31_424971 341 19 a a DT 10_1101-2020_12_31_424971 341 20 ( ( -LRB- 10_1101-2020_12_31_424971 341 21 which which WDT 10_1101-2020_12_31_424971 341 22 was be VBD 10_1101-2020_12_31_424971 341 23 not not RB 10_1101-2020_12_31_424971 341 24 certified certify VBN 10_1101-2020_12_31_424971 341 25 by by IN 10_1101-2020_12_31_424971 341 26 peer peer NN 10_1101-2020_12_31_424971 341 27 review review NN 10_1101-2020_12_31_424971 341 28 ) ) -RRB- 10_1101-2020_12_31_424971 341 29 is be VBZ 10_1101-2020_12_31_424971 341 30 the the DT 10_1101-2020_12_31_424971 341 31 author author NN 10_1101-2020_12_31_424971 341 32 / / SYM 10_1101-2020_12_31_424971 341 33 funder funder NN 10_1101-2020_12_31_424971 341 34 , , , 10_1101-2020_12_31_424971 341 35 who who WP 10_1101-2020_12_31_424971 341 36 has have VBZ 10_1101-2020_12_31_424971 341 37 granted grant VBN 10_1101-2020_12_31_424971 341 38 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 341 39 a a DT 10_1101-2020_12_31_424971 341 40 license license NN 10_1101-2020_12_31_424971 341 41 to to TO 10_1101-2020_12_31_424971 341 42 display display VB 10_1101-2020_12_31_424971 341 43 the the DT 10_1101-2020_12_31_424971 341 44 preprint preprint NN 10_1101-2020_12_31_424971 341 45 in in IN 10_1101-2020_12_31_424971 341 46 perpetuity perpetuity NN 10_1101-2020_12_31_424971 341 47 . . . 10_1101-2020_12_31_424971 342 1 It -PRON- PRP 10_1101-2020_12_31_424971 342 2 is be VBZ 10_1101-2020_12_31_424971 342 3 made make VBN 10_1101-2020_12_31_424971 342 4 The the DT 10_1101-2020_12_31_424971 342 5 copyright copyright NN 10_1101-2020_12_31_424971 342 6 holder holder NN 10_1101-2020_12_31_424971 342 7 for for IN 10_1101-2020_12_31_424971 342 8 this this DT 10_1101-2020_12_31_424971 342 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 342 10 version version NN 10_1101-2020_12_31_424971 342 11 posted post VBD 10_1101-2020_12_31_424971 342 12 January January NNP 10_1101-2020_12_31_424971 342 13 2 2 CD 10_1101-2020_12_31_424971 342 14 , , , 10_1101-2020_12_31_424971 342 15 2021 2021 CD 10_1101-2020_12_31_424971 342 16 . . . 10_1101-2020_12_31_424971 342 17 ; ; : 10_1101-2020_12_31_424971 342 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 342 19 : : : 10_1101-2020_12_31_424971 342 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 342 21 preprint preprint NN 10_1101-2020_12_31_424971 342 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 342 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 342 24 32 32 CD 10_1101-2020_12_31_424971 342 25 ICs ic NNS 10_1101-2020_12_31_424971 342 26 templated template VBN 10_1101-2020_12_31_424971 342 27 with with IN 10_1101-2020_12_31_424971 342 28 the the DT 10_1101-2020_12_31_424971 342 29 specified specify VBN 10_1101-2020_12_31_424971 342 30 variants variant NNS 10_1101-2020_12_31_424971 342 31 of of IN 10_1101-2020_12_31_424971 342 32 an an DT 10_1101-2020_12_31_424971 342 33 AUG AUG NNP 10_1101-2020_12_31_424971 342 34 - - HYPH 10_1101-2020_12_31_424971 342 35 NNN NNN NNP 10_1101-2020_12_31_424971 342 36 - - HYPH 10_1101-2020_12_31_424971 342 37 NGU NGU NNP 10_1101-2020_12_31_424971 342 38 - - HYPH 10_1101-2020_12_31_424971 342 39 U U NNP 10_1101-2020_12_31_424971 342 40 mRNA mrna NN 10_1101-2020_12_31_424971 342 41 were be VBD 10_1101-2020_12_31_424971 342 42 mixed mixed JJ 10_1101-2020_12_31_424971 342 43 with with IN 10_1101-2020_12_31_424971 342 44 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 342 45 - - HYPH 10_1101-2020_12_31_424971 342 46 778 778 CD 10_1101-2020_12_31_424971 342 47 TC TC NNP 10_1101-2020_12_31_424971 342 48 or or CC 10_1101-2020_12_31_424971 342 49 ProL ProL NNP 10_1101-2020_12_31_424971 342 50 - - HYPH 10_1101-2020_12_31_424971 342 51 TC TC NNP 10_1101-2020_12_31_424971 342 52 . . . 10_1101-2020_12_31_424971 343 1 fMP fMP NNP 10_1101-2020_12_31_424971 343 2 formation formation NN 10_1101-2020_12_31_424971 343 3 was be VBD 10_1101-2020_12_31_424971 343 4 monitored monitor VBN 10_1101-2020_12_31_424971 343 5 in in IN 10_1101-2020_12_31_424971 343 6 an an DT 10_1101-2020_12_31_424971 343 7 RQF-3 RQF-3 NNP 10_1101-2020_12_31_424971 343 8 Kintek Kintek NNPS 10_1101-2020_12_31_424971 343 9 chemical chemical NN 10_1101-2020_12_31_424971 343 10 quench quench NN 10_1101-2020_12_31_424971 343 11 apparatus apparatus NN 10_1101-2020_12_31_424971 343 12 . . . 10_1101-2020_12_31_424971 344 1 In in IN 10_1101-2020_12_31_424971 344 2 779 779 CD 10_1101-2020_12_31_424971 344 3 tripeptide tripeptide NN 10_1101-2020_12_31_424971 344 4 formation formation NN 10_1101-2020_12_31_424971 344 5 assays assays RB 10_1101-2020_12_31_424971 344 6 , , , 10_1101-2020_12_31_424971 344 7 70S 70S NNP 10_1101-2020_12_31_424971 344 8 ICs ic NNS 10_1101-2020_12_31_424971 344 9 templated template VBD 10_1101-2020_12_31_424971 344 10 with with IN 10_1101-2020_12_31_424971 344 11 the the DT 10_1101-2020_12_31_424971 344 12 specified specify VBN 10_1101-2020_12_31_424971 344 13 variants variant NNS 10_1101-2020_12_31_424971 344 14 of of IN 10_1101-2020_12_31_424971 344 15 the the DT 10_1101-2020_12_31_424971 344 16 AUG AUG NNP 10_1101-2020_12_31_424971 344 17 - - HYPH 10_1101-2020_12_31_424971 344 18 NCC NCC NNP 10_1101-2020_12_31_424971 344 19 - - HYPH 10_1101-2020_12_31_424971 344 20 NGU-780 NGU-780 NNP 10_1101-2020_12_31_424971 344 21 U U NNP 10_1101-2020_12_31_424971 344 22 mRNA mRNA NNS 10_1101-2020_12_31_424971 344 23 were be VBD 10_1101-2020_12_31_424971 344 24 mixed mixed JJ 10_1101-2020_12_31_424971 344 25 , , , 10_1101-2020_12_31_424971 344 26 either either CC 10_1101-2020_12_31_424971 344 27 in in IN 10_1101-2020_12_31_424971 344 28 one one CD 10_1101-2020_12_31_424971 344 29 step step NN 10_1101-2020_12_31_424971 344 30 or or CC 10_1101-2020_12_31_424971 344 31 in in IN 10_1101-2020_12_31_424971 344 32 two two CD 10_1101-2020_12_31_424971 344 33 steps step NNS 10_1101-2020_12_31_424971 344 34 , , , 10_1101-2020_12_31_424971 344 35 with with IN 10_1101-2020_12_31_424971 344 36 equimolar equimolar JJ 10_1101-2020_12_31_424971 344 37 mixtures mixture NNS 10_1101-2020_12_31_424971 344 38 of of IN 10_1101-2020_12_31_424971 344 39 SufB2- sufb2- NN 10_1101-2020_12_31_424971 344 40 , , , 10_1101-2020_12_31_424971 344 41 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 344 42 781 781 CD 10_1101-2020_12_31_424971 344 43 ( ( -LRB- 10_1101-2020_12_31_424971 344 44 anticodon anticodon NNP 10_1101-2020_12_31_424971 344 45 U*AC U*AC NNP 10_1101-2020_12_31_424971 344 46 , , , 10_1101-2020_12_31_424971 344 47 where where WRB 10_1101-2020_12_31_424971 344 48 U u NN 10_1101-2020_12_31_424971 344 49 * * NFP 10_1101-2020_12_31_424971 344 50 = = SYM 10_1101-2020_12_31_424971 344 51 cmo5U)- cmo5U)- NNS 10_1101-2020_12_31_424971 344 52 , , , 10_1101-2020_12_31_424971 344 53 and and CC 10_1101-2020_12_31_424971 344 54 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 344 55 ( ( -LRB- 10_1101-2020_12_31_424971 344 56 anticodon anticodon NNP 10_1101-2020_12_31_424971 344 57 ICG ICG NNP 10_1101-2020_12_31_424971 344 58 , , , 10_1101-2020_12_31_424971 344 59 where where WRB 10_1101-2020_12_31_424971 344 60 I -PRON- PRP 10_1101-2020_12_31_424971 344 61 = = SYM 10_1101-2020_12_31_424971 344 62 inosine)-TCs inosine)-TCs NNP 10_1101-2020_12_31_424971 344 63 and and CC 10_1101-2020_12_31_424971 344 64 782 782 CD 10_1101-2020_12_31_424971 344 65 EF EF NNP 10_1101-2020_12_31_424971 344 66 - - HYPH 10_1101-2020_12_31_424971 344 67 G. G. NNP 10_1101-2020_12_31_424971 345 1 Formation formation NN 10_1101-2020_12_31_424971 345 2 of of IN 10_1101-2020_12_31_424971 345 3 fMPV fmpv NN 10_1101-2020_12_31_424971 345 4 and and CC 10_1101-2020_12_31_424971 345 5 fMPR fmpr NN 10_1101-2020_12_31_424971 345 6 were be VBD 10_1101-2020_12_31_424971 345 7 monitored monitor VBN 10_1101-2020_12_31_424971 345 8 in in IN 10_1101-2020_12_31_424971 345 9 an an DT 10_1101-2020_12_31_424971 345 10 RQF-3 RQF-3 NNP 10_1101-2020_12_31_424971 345 11 Kintek Kintek NNPS 10_1101-2020_12_31_424971 345 12 chemical chemical NN 10_1101-2020_12_31_424971 345 13 quench quench NN 10_1101-2020_12_31_424971 345 14 783 783 CD 10_1101-2020_12_31_424971 345 15 apparatus apparatus NN 10_1101-2020_12_31_424971 345 16 . . . 10_1101-2020_12_31_424971 346 1 Tripeptide tripeptide JJ 10_1101-2020_12_31_424971 346 2 formation formation NN 10_1101-2020_12_31_424971 346 3 assays assay VBZ 10_1101-2020_12_31_424971 346 4 with with IN 10_1101-2020_12_31_424971 346 5 one one CD 10_1101-2020_12_31_424971 346 6 - - HYPH 10_1101-2020_12_31_424971 346 7 step step NN 10_1101-2020_12_31_424971 346 8 delivery delivery NN 10_1101-2020_12_31_424971 346 9 of of IN 10_1101-2020_12_31_424971 346 10 TCs tc NNS 10_1101-2020_12_31_424971 346 11 were be VBD 10_1101-2020_12_31_424971 346 12 initiated initiate VBN 10_1101-2020_12_31_424971 346 13 by by IN 10_1101-2020_12_31_424971 346 14 rapidly rapidly RB 10_1101-2020_12_31_424971 346 15 784 784 CD 10_1101-2020_12_31_424971 346 16 mixing mix VBG 10_1101-2020_12_31_424971 346 17 the the DT 10_1101-2020_12_31_424971 346 18 70S 70S NNP 10_1101-2020_12_31_424971 346 19 IC IC NNP 10_1101-2020_12_31_424971 346 20 with with IN 10_1101-2020_12_31_424971 346 21 two two CD 10_1101-2020_12_31_424971 346 22 or or CC 10_1101-2020_12_31_424971 346 23 more more JJR 10_1101-2020_12_31_424971 346 24 of of IN 10_1101-2020_12_31_424971 346 25 the the DT 10_1101-2020_12_31_424971 346 26 TCs tc NNS 10_1101-2020_12_31_424971 346 27 in in IN 10_1101-2020_12_31_424971 346 28 the the DT 10_1101-2020_12_31_424971 346 29 RQF-3 RQF-3 NNP 10_1101-2020_12_31_424971 346 30 Kintek Kintek NNPS 10_1101-2020_12_31_424971 346 31 chemical chemical NN 10_1101-2020_12_31_424971 346 32 quench quench NN 10_1101-2020_12_31_424971 346 33 apparatus apparatus NN 10_1101-2020_12_31_424971 346 34 . . . 10_1101-2020_12_31_424971 347 1 785 785 CD 10_1101-2020_12_31_424971 347 2 Final final JJ 10_1101-2020_12_31_424971 347 3 concentrations concentration NNS 10_1101-2020_12_31_424971 347 4 in in IN 10_1101-2020_12_31_424971 347 5 these these DT 10_1101-2020_12_31_424971 347 6 reactions reaction NNS 10_1101-2020_12_31_424971 347 7 were be VBD 10_1101-2020_12_31_424971 347 8 0.37 0.37 CD 10_1101-2020_12_31_424971 347 9 µM µm NN 10_1101-2020_12_31_424971 347 10 for for IN 10_1101-2020_12_31_424971 347 11 the the DT 10_1101-2020_12_31_424971 347 12 70S 70S NNP 10_1101-2020_12_31_424971 347 13 IC IC NNP 10_1101-2020_12_31_424971 347 14 ; ; : 10_1101-2020_12_31_424971 347 15 0.5 0.5 CD 10_1101-2020_12_31_424971 347 16 µM µm NN 10_1101-2020_12_31_424971 347 17 for for IN 10_1101-2020_12_31_424971 347 18 mRNA mRNA NNP 10_1101-2020_12_31_424971 347 19 ; ; : 10_1101-2020_12_31_424971 347 20 0.5 0.5 CD 10_1101-2020_12_31_424971 347 21 µM µM NNP 10_1101-2020_12_31_424971 347 22 786 786 CD 10_1101-2020_12_31_424971 347 23 each each DT 10_1101-2020_12_31_424971 347 24 for for IN 10_1101-2020_12_31_424971 347 25 IFs if NNS 10_1101-2020_12_31_424971 347 26 1 1 CD 10_1101-2020_12_31_424971 347 27 , , , 10_1101-2020_12_31_424971 347 28 2 2 CD 10_1101-2020_12_31_424971 347 29 , , , 10_1101-2020_12_31_424971 347 30 and and CC 10_1101-2020_12_31_424971 347 31 3 3 CD 10_1101-2020_12_31_424971 347 32 ; ; : 10_1101-2020_12_31_424971 347 33 0.25 0.25 CD 10_1101-2020_12_31_424971 347 34 µM µm NN 10_1101-2020_12_31_424971 347 35 for for IN 10_1101-2020_12_31_424971 347 36 [ [ -LRB- 10_1101-2020_12_31_424971 347 37 35S]-fMet 35s]-fmet CD 10_1101-2020_12_31_424971 347 38 - - HYPH 10_1101-2020_12_31_424971 347 39 tRNAfMet trnafmet CD 10_1101-2020_12_31_424971 347 40 ; ; : 10_1101-2020_12_31_424971 347 41 2.0 2.0 CD 10_1101-2020_12_31_424971 347 42 µM µm NN 10_1101-2020_12_31_424971 347 43 for for IN 10_1101-2020_12_31_424971 347 44 EF EF NNP 10_1101-2020_12_31_424971 347 45 - - HYPH 10_1101-2020_12_31_424971 347 46 G G NNP 10_1101-2020_12_31_424971 347 47 ; ; : 10_1101-2020_12_31_424971 347 48 0.75 0.75 CD 10_1101-2020_12_31_424971 347 49 µM µm NN 10_1101-2020_12_31_424971 347 50 for for IN 10_1101-2020_12_31_424971 347 51 EF EF NNP 10_1101-2020_12_31_424971 347 52 - - HYPH 10_1101-2020_12_31_424971 347 53 Tu Tu NNP 10_1101-2020_12_31_424971 347 54 for for IN 10_1101-2020_12_31_424971 347 55 787 787 CD 10_1101-2020_12_31_424971 347 56 each each DT 10_1101-2020_12_31_424971 347 57 aa aa NNP 10_1101-2020_12_31_424971 347 58 - - HYPH 10_1101-2020_12_31_424971 347 59 tRNA tRNA NNP 10_1101-2020_12_31_424971 347 60 ; ; : 10_1101-2020_12_31_424971 347 61 0.5 0.5 CD 10_1101-2020_12_31_424971 347 62 µM µm NN 10_1101-2020_12_31_424971 347 63 each each DT 10_1101-2020_12_31_424971 347 64 for for IN 10_1101-2020_12_31_424971 347 65 the the DT 10_1101-2020_12_31_424971 347 66 aa aa NNP 10_1101-2020_12_31_424971 347 67 - - HYPH 10_1101-2020_12_31_424971 347 68 tRNAs tRNAs NNP 10_1101-2020_12_31_424971 347 69 ; ; : 10_1101-2020_12_31_424971 347 70 and and CC 10_1101-2020_12_31_424971 347 71 1 1 CD 10_1101-2020_12_31_424971 347 72 mM mM NNP 10_1101-2020_12_31_424971 347 73 for for IN 10_1101-2020_12_31_424971 347 74 GTP GTP NNP 10_1101-2020_12_31_424971 347 75 . . . 10_1101-2020_12_31_424971 348 1 For for IN 10_1101-2020_12_31_424971 348 2 tripeptide tripeptide NN 10_1101-2020_12_31_424971 348 3 formation formation NN 10_1101-2020_12_31_424971 348 4 assays assays RB 10_1101-2020_12_31_424971 348 5 788 788 CD 10_1101-2020_12_31_424971 348 6 with with IN 10_1101-2020_12_31_424971 348 7 one one CD 10_1101-2020_12_31_424971 348 8 - - HYPH 10_1101-2020_12_31_424971 348 9 step step NN 10_1101-2020_12_31_424971 348 10 delivery delivery NN 10_1101-2020_12_31_424971 348 11 of of IN 10_1101-2020_12_31_424971 348 12 G37-state G37-state NNP 10_1101-2020_12_31_424971 348 13 SufB2- sufb2- NN 10_1101-2020_12_31_424971 348 14 , , , 10_1101-2020_12_31_424971 348 15 tRNAVal- tRNAVal- NNP 10_1101-2020_12_31_424971 348 16 , , , 10_1101-2020_12_31_424971 348 17 and and CC 10_1101-2020_12_31_424971 348 18 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 348 19 - - HYPH 10_1101-2020_12_31_424971 348 20 TCs TCs NNP 10_1101-2020_12_31_424971 348 21 to to IN 10_1101-2020_12_31_424971 348 22 the the DT 10_1101-2020_12_31_424971 348 23 70S 70S NNP 10_1101-2020_12_31_424971 348 24 ICs ic NNS 10_1101-2020_12_31_424971 348 25 , , , 10_1101-2020_12_31_424971 348 26 the the DT 10_1101-2020_12_31_424971 348 27 yield yield NN 10_1101-2020_12_31_424971 348 28 789 789 CD 10_1101-2020_12_31_424971 348 29 of of IN 10_1101-2020_12_31_424971 348 30 fMPV fMPV NNP 10_1101-2020_12_31_424971 348 31 and and CC 10_1101-2020_12_31_424971 348 32 kfMPV kfMPV NNP 10_1101-2020_12_31_424971 348 33 , , , 10_1101-2020_12_31_424971 348 34 obs obs NN 10_1101-2020_12_31_424971 348 35 report report NN 10_1101-2020_12_31_424971 348 36 on on IN 10_1101-2020_12_31_424971 348 37 the the DT 10_1101-2020_12_31_424971 348 38 activity activity NN 10_1101-2020_12_31_424971 348 39 of of IN 10_1101-2020_12_31_424971 348 40 ribosomes ribosome NNS 10_1101-2020_12_31_424971 348 41 that that WDT 10_1101-2020_12_31_424971 348 42 shifted shift VBD 10_1101-2020_12_31_424971 348 43 to to IN 10_1101-2020_12_31_424971 348 44 the the DT 10_1101-2020_12_31_424971 348 45 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 348 46 , , , 10_1101-2020_12_31_424971 348 47 whereas whereas IN 10_1101-2020_12_31_424971 348 48 the the DT 10_1101-2020_12_31_424971 348 49 790 790 CD 10_1101-2020_12_31_424971 348 50 yield yield NN 10_1101-2020_12_31_424971 348 51 of of IN 10_1101-2020_12_31_424971 348 52 fMPR fmpr NN 10_1101-2020_12_31_424971 348 53 and and CC 10_1101-2020_12_31_424971 348 54 kfMPR kfMPR NNP 10_1101-2020_12_31_424971 348 55 , , , 10_1101-2020_12_31_424971 348 56 obs obs NN 10_1101-2020_12_31_424971 348 57 report report NN 10_1101-2020_12_31_424971 348 58 on on IN 10_1101-2020_12_31_424971 348 59 the the DT 10_1101-2020_12_31_424971 348 60 activity activity NN 10_1101-2020_12_31_424971 348 61 of of IN 10_1101-2020_12_31_424971 348 62 ribosomes ribosome NNS 10_1101-2020_12_31_424971 348 63 that that WDT 10_1101-2020_12_31_424971 348 64 remained remain VBD 10_1101-2020_12_31_424971 348 65 in in IN 10_1101-2020_12_31_424971 348 66 the the DT 10_1101-2020_12_31_424971 348 67 0-frame29,34 0-frame29,34 CD 10_1101-2020_12_31_424971 348 68 . . . 10_1101-2020_12_31_424971 349 1 791 791 CD 10_1101-2020_12_31_424971 349 2 We -PRON- PRP 10_1101-2020_12_31_424971 349 3 chose choose VBD 10_1101-2020_12_31_424971 349 4 G37-state G37-state NNP 10_1101-2020_12_31_424971 349 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 349 6 to to TO 10_1101-2020_12_31_424971 349 7 maximize maximize VB 10_1101-2020_12_31_424971 349 8 its -PRON- PRP$ 10_1101-2020_12_31_424971 349 9 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 349 10 efficiency efficiency NN 10_1101-2020_12_31_424971 349 11 but but CC 10_1101-2020_12_31_424971 349 12 native native JJ 10_1101-2020_12_31_424971 349 13 - - HYPH 10_1101-2020_12_31_424971 349 14 state state NN 10_1101-2020_12_31_424971 349 15 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 349 16 792 792 CD 10_1101-2020_12_31_424971 349 17 and and CC 10_1101-2020_12_31_424971 349 18 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 349 19 to to TO 10_1101-2020_12_31_424971 349 20 prevent prevent VB 10_1101-2020_12_31_424971 349 21 them -PRON- PRP 10_1101-2020_12_31_424971 349 22 from from IN 10_1101-2020_12_31_424971 349 23 undergoing undergo VBG 10_1101-2020_12_31_424971 349 24 unwanted unwanted JJ 10_1101-2020_12_31_424971 349 25 frameshifting frameshifting NN 10_1101-2020_12_31_424971 349 26 ( ( -LRB- 10_1101-2020_12_31_424971 349 27 note note VB 10_1101-2020_12_31_424971 349 28 that that IN 10_1101-2020_12_31_424971 349 29 , , , 10_1101-2020_12_31_424971 349 30 for for IN 10_1101-2020_12_31_424971 349 31 simplicity simplicity NN 10_1101-2020_12_31_424971 349 32 , , , 10_1101-2020_12_31_424971 349 33 793 793 CD 10_1101-2020_12_31_424971 349 34 we -PRON- PRP 10_1101-2020_12_31_424971 349 35 have have VBP 10_1101-2020_12_31_424971 349 36 not not RB 10_1101-2020_12_31_424971 349 37 denoted denote VBN 10_1101-2020_12_31_424971 349 38 the the DT 10_1101-2020_12_31_424971 349 39 aminoacyl aminoacyl JJ 10_1101-2020_12_31_424971 349 40 or or CC 10_1101-2020_12_31_424971 349 41 dipeptidyl dipeptidyl JJ 10_1101-2020_12_31_424971 349 42 moieties moiety NNS 10_1101-2020_12_31_424971 349 43 of of IN 10_1101-2020_12_31_424971 349 44 the the DT 10_1101-2020_12_31_424971 349 45 tRNAs trnas CD 10_1101-2020_12_31_424971 349 46 ) ) -RRB- 10_1101-2020_12_31_424971 349 47 . . . 10_1101-2020_12_31_424971 350 1 Tripeptide tripeptide NN 10_1101-2020_12_31_424971 350 2 formation formation NN 10_1101-2020_12_31_424971 350 3 794 794 CD 10_1101-2020_12_31_424971 350 4 assays assay NNS 10_1101-2020_12_31_424971 350 5 with with IN 10_1101-2020_12_31_424971 350 6 two two CD 10_1101-2020_12_31_424971 350 7 - - HYPH 10_1101-2020_12_31_424971 350 8 step step NN 10_1101-2020_12_31_424971 350 9 delivery delivery NN 10_1101-2020_12_31_424971 350 10 of of IN 10_1101-2020_12_31_424971 350 11 TCs29 TCs29 NNS 10_1101-2020_12_31_424971 350 12 were be VBD 10_1101-2020_12_31_424971 350 13 performed perform VBN 10_1101-2020_12_31_424971 350 14 in in IN 10_1101-2020_12_31_424971 350 15 a a DT 10_1101-2020_12_31_424971 350 16 manner manner NN 10_1101-2020_12_31_424971 350 17 similar similar JJ 10_1101-2020_12_31_424971 350 18 to to IN 10_1101-2020_12_31_424971 350 19 those those DT 10_1101-2020_12_31_424971 350 20 with with IN 10_1101-2020_12_31_424971 350 21 one one CD 10_1101-2020_12_31_424971 350 22 - - HYPH 10_1101-2020_12_31_424971 350 23 step step NN 10_1101-2020_12_31_424971 350 24 795 795 CD 10_1101-2020_12_31_424971 350 25 delivery delivery NN 10_1101-2020_12_31_424971 350 26 of of IN 10_1101-2020_12_31_424971 350 27 TCs tc NNS 10_1101-2020_12_31_424971 350 28 , , , 10_1101-2020_12_31_424971 350 29 except except IN 10_1101-2020_12_31_424971 350 30 that that IN 10_1101-2020_12_31_424971 350 31 the the DT 10_1101-2020_12_31_424971 350 32 70S 70S NNP 10_1101-2020_12_31_424971 350 33 ICs ic NNS 10_1101-2020_12_31_424971 350 34 were be VBD 10_1101-2020_12_31_424971 350 35 incubated incubate VBN 10_1101-2020_12_31_424971 350 36 with with IN 10_1101-2020_12_31_424971 350 37 a a DT 10_1101-2020_12_31_424971 350 38 SufB2- sufb2- NN 10_1101-2020_12_31_424971 350 39 or or CC 10_1101-2020_12_31_424971 350 40 ProL ProL NNP 10_1101-2020_12_31_424971 350 41 - - HYPH 10_1101-2020_12_31_424971 350 42 TC TC NNP 10_1101-2020_12_31_424971 350 43 and and CC 10_1101-2020_12_31_424971 350 44 2.0 2.0 CD 10_1101-2020_12_31_424971 350 45 µM µM NNP 10_1101-2020_12_31_424971 350 46 796 796 CD 10_1101-2020_12_31_424971 350 47 EF EF NNP 10_1101-2020_12_31_424971 350 48 - - HYPH 10_1101-2020_12_31_424971 350 49 G G NNP 10_1101-2020_12_31_424971 350 50 for for IN 10_1101-2020_12_31_424971 350 51 0.5 0.5 CD 10_1101-2020_12_31_424971 350 52 - - SYM 10_1101-2020_12_31_424971 350 53 10 10 CD 10_1101-2020_12_31_424971 350 54 min min NN 10_1101-2020_12_31_424971 350 55 , , , 10_1101-2020_12_31_424971 350 56 as as IN 10_1101-2020_12_31_424971 350 57 specified specify VBN 10_1101-2020_12_31_424971 350 58 , , , 10_1101-2020_12_31_424971 350 59 followed follow VBN 10_1101-2020_12_31_424971 350 60 by by IN 10_1101-2020_12_31_424971 350 61 manual manual JJ 10_1101-2020_12_31_424971 350 62 addition addition NN 10_1101-2020_12_31_424971 350 63 of of IN 10_1101-2020_12_31_424971 350 64 an an DT 10_1101-2020_12_31_424971 350 65 equimolar equimolar JJ 10_1101-2020_12_31_424971 350 66 mixture mixture NN 10_1101-2020_12_31_424971 350 67 of of IN 10_1101-2020_12_31_424971 350 68 tRNAArg- trnaarg- NN 10_1101-2020_12_31_424971 350 69 797 797 CD 10_1101-2020_12_31_424971 350 70 and and CC 10_1101-2020_12_31_424971 350 71 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 350 72 - - HYPH 10_1101-2020_12_31_424971 350 73 TCs TCs NNP 10_1101-2020_12_31_424971 350 74 . . . 10_1101-2020_12_31_424971 351 1 Reactions reaction NNS 10_1101-2020_12_31_424971 351 2 were be VBD 10_1101-2020_12_31_424971 351 3 conducted conduct VBN 10_1101-2020_12_31_424971 351 4 at at IN 10_1101-2020_12_31_424971 351 5 20 20 CD 10_1101-2020_12_31_424971 351 6 ° ° NNS 10_1101-2020_12_31_424971 351 7 C C NNP 10_1101-2020_12_31_424971 351 8 unless unless IN 10_1101-2020_12_31_424971 351 9 otherwise otherwise RB 10_1101-2020_12_31_424971 351 10 specified specify VBN 10_1101-2020_12_31_424971 351 11 , , , 10_1101-2020_12_31_424971 351 12 and and CC 10_1101-2020_12_31_424971 351 13 were be VBD 10_1101-2020_12_31_424971 351 14 798 798 CD 10_1101-2020_12_31_424971 351 15 quenched quench VBN 10_1101-2020_12_31_424971 351 16 by by IN 10_1101-2020_12_31_424971 351 17 adding add VBG 10_1101-2020_12_31_424971 351 18 concentrated concentrate VBD 10_1101-2020_12_31_424971 351 19 KOH KOH NNP 10_1101-2020_12_31_424971 351 20 to to IN 10_1101-2020_12_31_424971 351 21 0.5 0.5 CD 10_1101-2020_12_31_424971 351 22 M. M. NNP 10_1101-2020_12_31_424971 351 23 After after IN 10_1101-2020_12_31_424971 351 24 a a DT 10_1101-2020_12_31_424971 351 25 brief brief JJ 10_1101-2020_12_31_424971 351 26 incubation incubation NN 10_1101-2020_12_31_424971 351 27 at at IN 10_1101-2020_12_31_424971 351 28 37 37 CD 10_1101-2020_12_31_424971 351 29 ° ° NNS 10_1101-2020_12_31_424971 351 30 C C NNP 10_1101-2020_12_31_424971 351 31 , , , 10_1101-2020_12_31_424971 351 32 aliquots aliquot NNS 10_1101-2020_12_31_424971 351 33 of of IN 10_1101-2020_12_31_424971 351 34 0.65 0.65 CD 10_1101-2020_12_31_424971 351 35 799 799 CD 10_1101-2020_12_31_424971 351 36 µL µl CD 10_1101-2020_12_31_424971 351 37 were be VBD 10_1101-2020_12_31_424971 351 38 spotted spot VBN 10_1101-2020_12_31_424971 351 39 onto onto IN 10_1101-2020_12_31_424971 351 40 a a DT 10_1101-2020_12_31_424971 351 41 cellulose cellulose NN 10_1101-2020_12_31_424971 351 42 - - HYPH 10_1101-2020_12_31_424971 351 43 backed back VBN 10_1101-2020_12_31_424971 351 44 plastic plastic NN 10_1101-2020_12_31_424971 351 45 TLC tlc NN 10_1101-2020_12_31_424971 351 46 sheet sheet NN 10_1101-2020_12_31_424971 351 47 and and CC 10_1101-2020_12_31_424971 351 48 electrophoresed electrophorese VBD 10_1101-2020_12_31_424971 351 49 at at IN 10_1101-2020_12_31_424971 351 50 1000 1000 CD 10_1101-2020_12_31_424971 351 51 V v NN 10_1101-2020_12_31_424971 351 52 in in IN 10_1101-2020_12_31_424971 351 53 800 800 CD 10_1101-2020_12_31_424971 351 54 PYRAC pyrac NN 10_1101-2020_12_31_424971 351 55 buffer buffer NN 10_1101-2020_12_31_424971 351 56 ( ( -LRB- 10_1101-2020_12_31_424971 351 57 62 62 CD 10_1101-2020_12_31_424971 351 58 mM mM NNP 10_1101-2020_12_31_424971 351 59 pyridine pyridine JJ 10_1101-2020_12_31_424971 351 60 , , , 10_1101-2020_12_31_424971 351 61 3.48 3.48 CD 10_1101-2020_12_31_424971 351 62 M M NNP 10_1101-2020_12_31_424971 351 63 acetic acetic JJ 10_1101-2020_12_31_424971 351 64 acid acid NN 10_1101-2020_12_31_424971 351 65 , , , 10_1101-2020_12_31_424971 351 66 pH pH NNP 10_1101-2020_12_31_424971 351 67 2.7 2.7 CD 10_1101-2020_12_31_424971 351 68 ) ) -RRB- 10_1101-2020_12_31_424971 351 69 until until IN 10_1101-2020_12_31_424971 351 70 the the DT 10_1101-2020_12_31_424971 351 71 marker marker NN 10_1101-2020_12_31_424971 351 72 dye dye NN 10_1101-2020_12_31_424971 351 73 bromophenol bromophenol NN 10_1101-2020_12_31_424971 351 74 801 801 CD 10_1101-2020_12_31_424971 351 75 blue blue JJ 10_1101-2020_12_31_424971 351 76 reached reach VBD 10_1101-2020_12_31_424971 351 77 the the DT 10_1101-2020_12_31_424971 351 78 water water NN 10_1101-2020_12_31_424971 351 79 - - HYPH 10_1101-2020_12_31_424971 351 80 oil oil NN 10_1101-2020_12_31_424971 351 81 interface interface NN 10_1101-2020_12_31_424971 351 82 at at IN 10_1101-2020_12_31_424971 351 83 the the DT 10_1101-2020_12_31_424971 351 84 anode29 anode29 NNS 10_1101-2020_12_31_424971 351 85 . . . 10_1101-2020_12_31_424971 352 1 The the DT 10_1101-2020_12_31_424971 352 2 position position NN 10_1101-2020_12_31_424971 352 3 of of IN 10_1101-2020_12_31_424971 352 4 the the DT 10_1101-2020_12_31_424971 352 5 origin origin NN 10_1101-2020_12_31_424971 352 6 was be VBD 10_1101-2020_12_31_424971 352 7 adjusted adjust VBN 10_1101-2020_12_31_424971 352 8 to to IN 10_1101-2020_12_31_424971 352 9 802 802 CD 10_1101-2020_12_31_424971 352 10 maximize maximize NN 10_1101-2020_12_31_424971 352 11 separation separation NN 10_1101-2020_12_31_424971 352 12 of of IN 10_1101-2020_12_31_424971 352 13 the the DT 10_1101-2020_12_31_424971 352 14 expected expect VBN 10_1101-2020_12_31_424971 352 15 oligopeptide oligopeptide IN 10_1101-2020_12_31_424971 352 16 products product NNS 10_1101-2020_12_31_424971 352 17 . . . 10_1101-2020_12_31_424971 353 1 The the DT 10_1101-2020_12_31_424971 353 2 separation separation NN 10_1101-2020_12_31_424971 353 3 of of IN 10_1101-2020_12_31_424971 353 4 unreacted unreacted JJ 10_1101-2020_12_31_424971 353 5 [ [ -LRB- 10_1101-2020_12_31_424971 353 6 35S]-803 35s]-803 CD 10_1101-2020_12_31_424971 353 7 .CC .CC NFP 10_1101-2020_12_31_424971 353 8 - - HYPH 10_1101-2020_12_31_424971 353 9 BY by IN 10_1101-2020_12_31_424971 353 10 - - HYPH 10_1101-2020_12_31_424971 353 11 NC NC NNP 10_1101-2020_12_31_424971 353 12 - - HYPH 10_1101-2020_12_31_424971 353 13 ND ND NNP 10_1101-2020_12_31_424971 353 14 4.0 4.0 CD 10_1101-2020_12_31_424971 353 15 International International NNP 10_1101-2020_12_31_424971 353 16 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 353 17 under under IN 10_1101-2020_12_31_424971 353 18 a a DT 10_1101-2020_12_31_424971 353 19 ( ( -LRB- 10_1101-2020_12_31_424971 353 20 which which WDT 10_1101-2020_12_31_424971 353 21 was be VBD 10_1101-2020_12_31_424971 353 22 not not RB 10_1101-2020_12_31_424971 353 23 certified certify VBN 10_1101-2020_12_31_424971 353 24 by by IN 10_1101-2020_12_31_424971 353 25 peer peer NN 10_1101-2020_12_31_424971 353 26 review review NN 10_1101-2020_12_31_424971 353 27 ) ) -RRB- 10_1101-2020_12_31_424971 353 28 is be VBZ 10_1101-2020_12_31_424971 353 29 the the DT 10_1101-2020_12_31_424971 353 30 author author NN 10_1101-2020_12_31_424971 353 31 / / SYM 10_1101-2020_12_31_424971 353 32 funder funder NN 10_1101-2020_12_31_424971 353 33 , , , 10_1101-2020_12_31_424971 353 34 who who WP 10_1101-2020_12_31_424971 353 35 has have VBZ 10_1101-2020_12_31_424971 353 36 granted grant VBN 10_1101-2020_12_31_424971 353 37 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 353 38 a a DT 10_1101-2020_12_31_424971 353 39 license license NN 10_1101-2020_12_31_424971 353 40 to to TO 10_1101-2020_12_31_424971 353 41 display display VB 10_1101-2020_12_31_424971 353 42 the the DT 10_1101-2020_12_31_424971 353 43 preprint preprint NN 10_1101-2020_12_31_424971 353 44 in in IN 10_1101-2020_12_31_424971 353 45 perpetuity perpetuity NN 10_1101-2020_12_31_424971 353 46 . . . 10_1101-2020_12_31_424971 354 1 It -PRON- PRP 10_1101-2020_12_31_424971 354 2 is be VBZ 10_1101-2020_12_31_424971 354 3 made make VBN 10_1101-2020_12_31_424971 354 4 The the DT 10_1101-2020_12_31_424971 354 5 copyright copyright NN 10_1101-2020_12_31_424971 354 6 holder holder NN 10_1101-2020_12_31_424971 354 7 for for IN 10_1101-2020_12_31_424971 354 8 this this DT 10_1101-2020_12_31_424971 354 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 354 10 version version NN 10_1101-2020_12_31_424971 354 11 posted post VBD 10_1101-2020_12_31_424971 354 12 January January NNP 10_1101-2020_12_31_424971 354 13 2 2 CD 10_1101-2020_12_31_424971 354 14 , , , 10_1101-2020_12_31_424971 354 15 2021 2021 CD 10_1101-2020_12_31_424971 354 16 . . . 10_1101-2020_12_31_424971 354 17 ; ; : 10_1101-2020_12_31_424971 354 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 354 19 : : : 10_1101-2020_12_31_424971 354 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 354 21 preprint preprint NN 10_1101-2020_12_31_424971 354 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 354 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 354 24 33 33 CD 10_1101-2020_12_31_424971 354 25 fMet fMet NNP 10_1101-2020_12_31_424971 354 26 and and CC 10_1101-2020_12_31_424971 354 27 each each DT 10_1101-2020_12_31_424971 354 28 of of IN 10_1101-2020_12_31_424971 354 29 the the DT 10_1101-2020_12_31_424971 354 30 [ [ -LRB- 10_1101-2020_12_31_424971 354 31 35S]-fMet 35s]-fmet CD 10_1101-2020_12_31_424971 354 32 - - HYPH 10_1101-2020_12_31_424971 354 33 peptide peptide NN 10_1101-2020_12_31_424971 354 34 products product NNS 10_1101-2020_12_31_424971 354 35 was be VBD 10_1101-2020_12_31_424971 354 36 visualized visualize VBN 10_1101-2020_12_31_424971 354 37 by by IN 10_1101-2020_12_31_424971 354 38 phosphorimaging phosphorimage VBG 10_1101-2020_12_31_424971 354 39 and and CC 10_1101-2020_12_31_424971 354 40 804 804 CD 10_1101-2020_12_31_424971 354 41 quantified quantify VBN 10_1101-2020_12_31_424971 354 42 using use VBG 10_1101-2020_12_31_424971 354 43 ImageQuant ImageQuant NNP 10_1101-2020_12_31_424971 354 44 ( ( -LRB- 10_1101-2020_12_31_424971 354 45 GE GE NNP 10_1101-2020_12_31_424971 354 46 Healthcare Healthcare NNP 10_1101-2020_12_31_424971 354 47 ) ) -RRB- 10_1101-2020_12_31_424971 354 48 and and CC 10_1101-2020_12_31_424971 354 49 kinetic kinetic JJ 10_1101-2020_12_31_424971 354 50 plots plot NNS 10_1101-2020_12_31_424971 354 51 were be VBD 10_1101-2020_12_31_424971 354 52 fitted fit VBN 10_1101-2020_12_31_424971 354 53 using use VBG 10_1101-2020_12_31_424971 354 54 Kaleidagraph Kaleidagraph NNP 10_1101-2020_12_31_424971 354 55 805 805 CD 10_1101-2020_12_31_424971 354 56 ( ( -LRB- 10_1101-2020_12_31_424971 354 57 Synergy Synergy NNP 10_1101-2020_12_31_424971 354 58 software software NN 10_1101-2020_12_31_424971 354 59 ) ) -RRB- 10_1101-2020_12_31_424971 354 60 . . . 10_1101-2020_12_31_424971 355 1 806 806 CD 10_1101-2020_12_31_424971 355 2 807 807 CD 10_1101-2020_12_31_424971 355 3 Assembly Assembly NNP 10_1101-2020_12_31_424971 355 4 and and CC 10_1101-2020_12_31_424971 355 5 purification purification NN 10_1101-2020_12_31_424971 355 6 of of IN 10_1101-2020_12_31_424971 355 7 70S 70S NNP 10_1101-2020_12_31_424971 355 8 ICs ic NNS 10_1101-2020_12_31_424971 355 9 , , , 10_1101-2020_12_31_424971 355 10 TCs tc NNS 10_1101-2020_12_31_424971 355 11 , , , 10_1101-2020_12_31_424971 355 12 POST POST NNP 10_1101-2020_12_31_424971 355 13 , , , 10_1101-2020_12_31_424971 355 14 and and CC 10_1101-2020_12_31_424971 355 15 PRE PRE NNP 10_1101-2020_12_31_424971 355 16 – – : 10_1101-2020_12_31_424971 355 17 A a DT 10_1101-2020_12_31_424971 355 18 complexes complex NNS 10_1101-2020_12_31_424971 355 19 for for IN 10_1101-2020_12_31_424971 355 20 use use NN 10_1101-2020_12_31_424971 355 21 in in IN 10_1101-2020_12_31_424971 355 22 smFRET smfret $ 10_1101-2020_12_31_424971 355 23 808 808 CD 10_1101-2020_12_31_424971 355 24 experiments experiment NNS 10_1101-2020_12_31_424971 355 25 . . . 10_1101-2020_12_31_424971 356 1 70S 70s CD 10_1101-2020_12_31_424971 356 2 ICs ic NNS 10_1101-2020_12_31_424971 356 3 were be VBD 10_1101-2020_12_31_424971 356 4 assembled assemble VBN 10_1101-2020_12_31_424971 356 5 in in IN 10_1101-2020_12_31_424971 356 6 a a DT 10_1101-2020_12_31_424971 356 7 manner manner NN 10_1101-2020_12_31_424971 356 8 analogous analogous JJ 10_1101-2020_12_31_424971 356 9 to to IN 10_1101-2020_12_31_424971 356 10 those those DT 10_1101-2020_12_31_424971 356 11 for for IN 10_1101-2020_12_31_424971 356 12 the the DT 10_1101-2020_12_31_424971 356 13 ensemble ensemble JJ 10_1101-2020_12_31_424971 356 14 rapid rapid JJ 10_1101-2020_12_31_424971 356 15 809 809 CD 10_1101-2020_12_31_424971 356 16 kinetic kinetic JJ 10_1101-2020_12_31_424971 356 17 studies study NNS 10_1101-2020_12_31_424971 356 18 described describe VBN 10_1101-2020_12_31_424971 356 19 above above RB 10_1101-2020_12_31_424971 356 20 , , , 10_1101-2020_12_31_424971 356 21 except except IN 10_1101-2020_12_31_424971 356 22 that that IN 10_1101-2020_12_31_424971 356 23 the the DT 10_1101-2020_12_31_424971 356 24 mRNA mrna NN 10_1101-2020_12_31_424971 356 25 containing contain VBG 10_1101-2020_12_31_424971 356 26 an an DT 10_1101-2020_12_31_424971 356 27 AUG AUG NNP 10_1101-2020_12_31_424971 356 28 - - HYPH 10_1101-2020_12_31_424971 356 29 CCC CCC NNP 10_1101-2020_12_31_424971 356 30 - - HYPH 10_1101-2020_12_31_424971 356 31 CGU CGU NNP 10_1101-2020_12_31_424971 356 32 - - HYPH 10_1101-2020_12_31_424971 356 33 U u NN 10_1101-2020_12_31_424971 356 34 coding code VBG 10_1101-2020_12_31_424971 356 35 810 810 CD 10_1101-2020_12_31_424971 356 36 sequence sequence NN 10_1101-2020_12_31_424971 356 37 was be VBD 10_1101-2020_12_31_424971 356 38 5'-biotinylated 5'-biotinylate VBN 10_1101-2020_12_31_424971 356 39 and and CC 10_1101-2020_12_31_424971 356 40 the the DT 10_1101-2020_12_31_424971 356 41 50S 50S NNP 10_1101-2020_12_31_424971 356 42 subunits subunit NNS 10_1101-2020_12_31_424971 356 43 were be VBD 10_1101-2020_12_31_424971 356 44 labeled label VBN 10_1101-2020_12_31_424971 356 45 with with IN 10_1101-2020_12_31_424971 356 46 bL9(Cy3 bl9(cy3 NN 10_1101-2020_12_31_424971 356 47 ) ) -RRB- 10_1101-2020_12_31_424971 356 48 and and CC 10_1101-2020_12_31_424971 356 49 uL1(Cy5 ul1(cy5 NN 10_1101-2020_12_31_424971 356 50 ) ) -RRB- 10_1101-2020_12_31_424971 356 51 . . . 10_1101-2020_12_31_424971 357 1 811 811 CD 10_1101-2020_12_31_424971 357 2 More more RBR 10_1101-2020_12_31_424971 357 3 specifically specifically RB 10_1101-2020_12_31_424971 357 4 , , , 10_1101-2020_12_31_424971 357 5 70S 70S NNP 10_1101-2020_12_31_424971 357 6 ICs ic NNS 10_1101-2020_12_31_424971 357 7 were be VBD 10_1101-2020_12_31_424971 357 8 assembled assemble VBN 10_1101-2020_12_31_424971 357 9 in in IN 10_1101-2020_12_31_424971 357 10 three three CD 10_1101-2020_12_31_424971 357 11 steps step NNS 10_1101-2020_12_31_424971 357 12 . . . 10_1101-2020_12_31_424971 358 1 First first RB 10_1101-2020_12_31_424971 358 2 , , , 10_1101-2020_12_31_424971 358 3 15 15 CD 10_1101-2020_12_31_424971 358 4 pmol pmol NN 10_1101-2020_12_31_424971 358 5 of of IN 10_1101-2020_12_31_424971 358 6 30S 30s CD 10_1101-2020_12_31_424971 358 7 subunits subunit NNS 10_1101-2020_12_31_424971 358 8 , , , 10_1101-2020_12_31_424971 358 9 27 27 CD 10_1101-2020_12_31_424971 358 10 pmol pmol NN 10_1101-2020_12_31_424971 358 11 812 812 CD 10_1101-2020_12_31_424971 358 12 of of IN 10_1101-2020_12_31_424971 358 13 IF1 IF1 NNP 10_1101-2020_12_31_424971 358 14 , , , 10_1101-2020_12_31_424971 358 15 27 27 CD 10_1101-2020_12_31_424971 358 16 pmol pmol NN 10_1101-2020_12_31_424971 358 17 of of IN 10_1101-2020_12_31_424971 358 18 IF2 IF2 NNP 10_1101-2020_12_31_424971 358 19 , , , 10_1101-2020_12_31_424971 358 20 27 27 CD 10_1101-2020_12_31_424971 358 21 pmol pmol NN 10_1101-2020_12_31_424971 358 22 of of IN 10_1101-2020_12_31_424971 358 23 IF3 IF3 NNP 10_1101-2020_12_31_424971 358 24 , , , 10_1101-2020_12_31_424971 358 25 18 18 CD 10_1101-2020_12_31_424971 358 26 nmol nmol NN 10_1101-2020_12_31_424971 358 27 of of IN 10_1101-2020_12_31_424971 358 28 GTP GTP NNP 10_1101-2020_12_31_424971 358 29 , , , 10_1101-2020_12_31_424971 358 30 and and CC 10_1101-2020_12_31_424971 358 31 25 25 CD 10_1101-2020_12_31_424971 358 32 pmol pmol NN 10_1101-2020_12_31_424971 358 33 of of IN 10_1101-2020_12_31_424971 358 34 biotin biotin NN 10_1101-2020_12_31_424971 358 35 - - HYPH 10_1101-2020_12_31_424971 358 36 mRNA mrna NN 10_1101-2020_12_31_424971 358 37 in in IN 10_1101-2020_12_31_424971 358 38 7 7 CD 10_1101-2020_12_31_424971 358 39 µL µL NNP 10_1101-2020_12_31_424971 358 40 of of IN 10_1101-2020_12_31_424971 358 41 813 813 CD 10_1101-2020_12_31_424971 358 42 Tris Tris NNP 10_1101-2020_12_31_424971 358 43 - - HYPH 10_1101-2020_12_31_424971 358 44 Polymix Polymix NNP 10_1101-2020_12_31_424971 358 45 Buffer Buffer NNP 10_1101-2020_12_31_424971 358 46 ( ( -LRB- 10_1101-2020_12_31_424971 358 47 50 50 CD 10_1101-2020_12_31_424971 358 48 mM mm NN 10_1101-2020_12_31_424971 358 49 Tris-(hydroxymethyl)-aminomethane tris-(hydroxymethyl)-aminomethane NN 10_1101-2020_12_31_424971 358 50 acetate acetate NN 10_1101-2020_12_31_424971 358 51 ( ( -LRB- 10_1101-2020_12_31_424971 358 52 Tris Tris NNP 10_1101-2020_12_31_424971 358 53 - - HYPH 10_1101-2020_12_31_424971 358 54 OAc OAc NNP 10_1101-2020_12_31_424971 358 55 ) ) -RRB- 10_1101-2020_12_31_424971 358 56 ( ( -LRB- 10_1101-2020_12_31_424971 358 57 pH25 pH25 JJR 10_1101-2020_12_31_424971 358 58 ° ° NNP 10_1101-2020_12_31_424971 358 59 C C NNP 10_1101-2020_12_31_424971 358 60 = = SYM 10_1101-2020_12_31_424971 358 61 814 814 CD 10_1101-2020_12_31_424971 358 62 7.0 7.0 CD 10_1101-2020_12_31_424971 358 63 ) ) -RRB- 10_1101-2020_12_31_424971 358 64 , , , 10_1101-2020_12_31_424971 358 65 100 100 CD 10_1101-2020_12_31_424971 358 66 mM mM NNP 10_1101-2020_12_31_424971 358 67 KCl KCl NNS 10_1101-2020_12_31_424971 358 68 , , , 10_1101-2020_12_31_424971 358 69 5 5 CD 10_1101-2020_12_31_424971 358 70 mM mm NN 10_1101-2020_12_31_424971 358 71 NH4OAc NH4OAc NNS 10_1101-2020_12_31_424971 358 72 , , , 10_1101-2020_12_31_424971 358 73 0.5 0.5 CD 10_1101-2020_12_31_424971 358 74 mM mm NN 10_1101-2020_12_31_424971 358 75 Ca(OAc)2 ca(oac)2 NN 10_1101-2020_12_31_424971 358 76 , , , 10_1101-2020_12_31_424971 358 77 0.1 0.1 CD 10_1101-2020_12_31_424971 358 78 mM mm NN 10_1101-2020_12_31_424971 358 79 EDTA edta NN 10_1101-2020_12_31_424971 358 80 , , , 10_1101-2020_12_31_424971 358 81 10 10 CD 10_1101-2020_12_31_424971 358 82 mM mm NN 10_1101-2020_12_31_424971 358 83 2-mercaptoethanol 2-mercaptoethanol CD 10_1101-2020_12_31_424971 358 84 815 815 CD 10_1101-2020_12_31_424971 358 85 ( ( -LRB- 10_1101-2020_12_31_424971 358 86 BME BME NNP 10_1101-2020_12_31_424971 358 87 ) ) -RRB- 10_1101-2020_12_31_424971 358 88 , , , 10_1101-2020_12_31_424971 358 89 5 5 CD 10_1101-2020_12_31_424971 358 90 mM mM NNP 10_1101-2020_12_31_424971 358 91 putrescine putrescine NN 10_1101-2020_12_31_424971 358 92 dihydrochloride dihydrochloride NN 10_1101-2020_12_31_424971 358 93 , , , 10_1101-2020_12_31_424971 358 94 and and CC 10_1101-2020_12_31_424971 358 95 1 1 CD 10_1101-2020_12_31_424971 358 96 mM mM NNP 10_1101-2020_12_31_424971 358 97 spermidine spermidine NN 10_1101-2020_12_31_424971 358 98 ( ( -LRB- 10_1101-2020_12_31_424971 358 99 free free JJ 10_1101-2020_12_31_424971 358 100 base base NN 10_1101-2020_12_31_424971 358 101 ) ) -RRB- 10_1101-2020_12_31_424971 358 102 ) ) -RRB- 10_1101-2020_12_31_424971 358 103 at at IN 10_1101-2020_12_31_424971 358 104 5 5 CD 10_1101-2020_12_31_424971 358 105 mM mm NN 10_1101-2020_12_31_424971 358 106 Mg(OAc)2 mg(oac)2 NN 10_1101-2020_12_31_424971 358 107 816 816 CD 10_1101-2020_12_31_424971 358 108 were be VBD 10_1101-2020_12_31_424971 358 109 incubated incubate VBN 10_1101-2020_12_31_424971 358 110 for for IN 10_1101-2020_12_31_424971 358 111 10 10 CD 10_1101-2020_12_31_424971 358 112 min min NN 10_1101-2020_12_31_424971 358 113 at at IN 10_1101-2020_12_31_424971 358 114 37 37 CD 10_1101-2020_12_31_424971 358 115 ºC. ºC. NNS 10_1101-2020_12_31_424971 359 1 Then then RB 10_1101-2020_12_31_424971 359 2 20 20 CD 10_1101-2020_12_31_424971 359 3 pmol pmol NN 10_1101-2020_12_31_424971 359 4 of of IN 10_1101-2020_12_31_424971 359 5 fMet fMet NNP 10_1101-2020_12_31_424971 359 6 - - HYPH 10_1101-2020_12_31_424971 359 7 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 359 8 in in IN 10_1101-2020_12_31_424971 359 9 2 2 CD 10_1101-2020_12_31_424971 359 10 µL µl CD 10_1101-2020_12_31_424971 359 11 of of IN 10_1101-2020_12_31_424971 359 12 10 10 CD 10_1101-2020_12_31_424971 359 13 mM mM NNP 10_1101-2020_12_31_424971 359 14 KOAc KOAc NNS 10_1101-2020_12_31_424971 359 15 ( ( -LRB- 10_1101-2020_12_31_424971 359 16 pH ph NN 10_1101-2020_12_31_424971 359 17 = = SYM 10_1101-2020_12_31_424971 359 18 817 817 CD 10_1101-2020_12_31_424971 359 19 5 5 CD 10_1101-2020_12_31_424971 359 20 ) ) -RRB- 10_1101-2020_12_31_424971 359 21 was be VBD 10_1101-2020_12_31_424971 359 22 added add VBN 10_1101-2020_12_31_424971 359 23 to to IN 10_1101-2020_12_31_424971 359 24 the the DT 10_1101-2020_12_31_424971 359 25 reaction reaction NN 10_1101-2020_12_31_424971 359 26 , , , 10_1101-2020_12_31_424971 359 27 followed follow VBN 10_1101-2020_12_31_424971 359 28 by by IN 10_1101-2020_12_31_424971 359 29 an an DT 10_1101-2020_12_31_424971 359 30 additional additional JJ 10_1101-2020_12_31_424971 359 31 incubation incubation NN 10_1101-2020_12_31_424971 359 32 of of IN 10_1101-2020_12_31_424971 359 33 10 10 CD 10_1101-2020_12_31_424971 359 34 min min NN 10_1101-2020_12_31_424971 359 35 at at IN 10_1101-2020_12_31_424971 359 36 37 37 CD 10_1101-2020_12_31_424971 359 37 ºC. ºC. NNS 10_1101-2020_12_31_424971 360 1 Finally finally RB 10_1101-2020_12_31_424971 360 2 , , , 10_1101-2020_12_31_424971 360 3 10 10 CD 10_1101-2020_12_31_424971 360 4 818 818 CD 10_1101-2020_12_31_424971 360 5 pmol pmol NN 10_1101-2020_12_31_424971 360 6 of of IN 10_1101-2020_12_31_424971 360 7 bL9(Cy3)- bL9(Cy3)- NNS 10_1101-2020_12_31_424971 360 8 and and CC 10_1101-2020_12_31_424971 360 9 uL1(Cy5)-labeled ul1(cy5)-labeled CD 10_1101-2020_12_31_424971 360 10 50S 50s CD 10_1101-2020_12_31_424971 360 11 subunits subunit NNS 10_1101-2020_12_31_424971 360 12 in in IN 10_1101-2020_12_31_424971 360 13 1 1 CD 10_1101-2020_12_31_424971 360 14 µL µl CD 10_1101-2020_12_31_424971 360 15 of of IN 10_1101-2020_12_31_424971 360 16 Reconstitution Reconstitution NNP 10_1101-2020_12_31_424971 360 17 Buffer Buffer NNP 10_1101-2020_12_31_424971 360 18 ( ( -LRB- 10_1101-2020_12_31_424971 360 19 20 20 CD 10_1101-2020_12_31_424971 360 20 mM mm NN 10_1101-2020_12_31_424971 360 21 819 819 CD 10_1101-2020_12_31_424971 360 22 Tris Tris NNP 10_1101-2020_12_31_424971 360 23 - - HYPH 10_1101-2020_12_31_424971 360 24 HCl HCl NNP 10_1101-2020_12_31_424971 360 25 ( ( -LRB- 10_1101-2020_12_31_424971 360 26 pH25 pH25 JJR 10_1101-2020_12_31_424971 360 27 ° ° NNP 10_1101-2020_12_31_424971 360 28 C C NNP 10_1101-2020_12_31_424971 360 29 = = SYM 10_1101-2020_12_31_424971 360 30 7.8 7.8 CD 10_1101-2020_12_31_424971 360 31 ) ) -RRB- 10_1101-2020_12_31_424971 360 32 , , , 10_1101-2020_12_31_424971 360 33 8 8 CD 10_1101-2020_12_31_424971 360 34 mM mm NN 10_1101-2020_12_31_424971 360 35 Mg(OAc)2 mg(oac)2 NN 10_1101-2020_12_31_424971 360 36 , , , 10_1101-2020_12_31_424971 360 37 150 150 CD 10_1101-2020_12_31_424971 360 38 mM mm NN 10_1101-2020_12_31_424971 360 39 NH4Cl nh4cl CD 10_1101-2020_12_31_424971 360 40 , , , 10_1101-2020_12_31_424971 360 41 0.2 0.2 CD 10_1101-2020_12_31_424971 360 42 mM mm NN 10_1101-2020_12_31_424971 360 43 EDTA edta NN 10_1101-2020_12_31_424971 360 44 , , , 10_1101-2020_12_31_424971 360 45 and and CC 10_1101-2020_12_31_424971 360 46 5 5 CD 10_1101-2020_12_31_424971 360 47 mM mm NN 10_1101-2020_12_31_424971 360 48 BME BME NNP 10_1101-2020_12_31_424971 360 49 ) ) -RRB- 10_1101-2020_12_31_424971 360 50 was be VBD 10_1101-2020_12_31_424971 360 51 820 820 CD 10_1101-2020_12_31_424971 360 52 added add VBN 10_1101-2020_12_31_424971 360 53 to to IN 10_1101-2020_12_31_424971 360 54 the the DT 10_1101-2020_12_31_424971 360 55 reaction reaction NN 10_1101-2020_12_31_424971 360 56 to to TO 10_1101-2020_12_31_424971 360 57 give give VB 10_1101-2020_12_31_424971 360 58 a a DT 10_1101-2020_12_31_424971 360 59 final final JJ 10_1101-2020_12_31_424971 360 60 volume volume NN 10_1101-2020_12_31_424971 360 61 of of IN 10_1101-2020_12_31_424971 360 62 10 10 CD 10_1101-2020_12_31_424971 360 63 µL µl CD 10_1101-2020_12_31_424971 360 64 , , , 10_1101-2020_12_31_424971 360 65 followed follow VBN 10_1101-2020_12_31_424971 360 66 by by IN 10_1101-2020_12_31_424971 360 67 a a DT 10_1101-2020_12_31_424971 360 68 final final JJ 10_1101-2020_12_31_424971 360 69 incubation incubation NN 10_1101-2020_12_31_424971 360 70 of of IN 10_1101-2020_12_31_424971 360 71 10 10 CD 10_1101-2020_12_31_424971 360 72 min min NN 10_1101-2020_12_31_424971 360 73 at at IN 10_1101-2020_12_31_424971 360 74 821 821 CD 10_1101-2020_12_31_424971 360 75 37 37 CD 10_1101-2020_12_31_424971 360 76 ºC. ºC. NNS 10_1101-2020_12_31_424971 361 1 The the DT 10_1101-2020_12_31_424971 361 2 reaction reaction NN 10_1101-2020_12_31_424971 361 3 was be VBD 10_1101-2020_12_31_424971 361 4 then then RB 10_1101-2020_12_31_424971 361 5 adjusted adjust VBN 10_1101-2020_12_31_424971 361 6 to to IN 10_1101-2020_12_31_424971 361 7 100 100 CD 10_1101-2020_12_31_424971 361 8 µL µL NNP 10_1101-2020_12_31_424971 361 9 with with IN 10_1101-2020_12_31_424971 361 10 Tris Tris NNP 10_1101-2020_12_31_424971 361 11 - - HYPH 10_1101-2020_12_31_424971 361 12 Polymix Polymix NNP 10_1101-2020_12_31_424971 361 13 Buffer Buffer NNP 10_1101-2020_12_31_424971 361 14 at at IN 10_1101-2020_12_31_424971 361 15 20 20 CD 10_1101-2020_12_31_424971 361 16 mM mm NN 10_1101-2020_12_31_424971 361 17 Mg(OAc)2 mg(oac)2 NN 10_1101-2020_12_31_424971 361 18 , , , 10_1101-2020_12_31_424971 361 19 822 822 CD 10_1101-2020_12_31_424971 361 20 loaded load VBN 10_1101-2020_12_31_424971 361 21 onto onto IN 10_1101-2020_12_31_424971 361 22 a a DT 10_1101-2020_12_31_424971 361 23 10 10 CD 10_1101-2020_12_31_424971 361 24 - - SYM 10_1101-2020_12_31_424971 361 25 40 40 CD 10_1101-2020_12_31_424971 361 26 % % NN 10_1101-2020_12_31_424971 361 27 ( ( -LRB- 10_1101-2020_12_31_424971 361 28 w w NNP 10_1101-2020_12_31_424971 361 29 / / SYM 10_1101-2020_12_31_424971 361 30 v v NNP 10_1101-2020_12_31_424971 361 31 ) ) -RRB- 10_1101-2020_12_31_424971 361 32 sucrose sucrose JJ 10_1101-2020_12_31_424971 361 33 gradient gradient NN 10_1101-2020_12_31_424971 361 34 prepared prepare VBD 10_1101-2020_12_31_424971 361 35 in in IN 10_1101-2020_12_31_424971 361 36 Tris Tris NNP 10_1101-2020_12_31_424971 361 37 - - HYPH 10_1101-2020_12_31_424971 361 38 Polymix Polymix NNP 10_1101-2020_12_31_424971 361 39 Buffer Buffer NNP 10_1101-2020_12_31_424971 361 40 at at IN 10_1101-2020_12_31_424971 361 41 20 20 CD 10_1101-2020_12_31_424971 361 42 mM mm NN 10_1101-2020_12_31_424971 361 43 Mg(OAc)2 Mg(OAc)2 NNS 10_1101-2020_12_31_424971 361 44 , , , 10_1101-2020_12_31_424971 361 45 823 823 CD 10_1101-2020_12_31_424971 361 46 and and CC 10_1101-2020_12_31_424971 361 47 purified purify VBN 10_1101-2020_12_31_424971 361 48 by by IN 10_1101-2020_12_31_424971 361 49 sucrose sucrose JJ 10_1101-2020_12_31_424971 361 50 density density NN 10_1101-2020_12_31_424971 361 51 gradient gradient NNP 10_1101-2020_12_31_424971 361 52 ultracentrifugation ultracentrifugation NN 10_1101-2020_12_31_424971 361 53 to to TO 10_1101-2020_12_31_424971 361 54 remove remove VB 10_1101-2020_12_31_424971 361 55 any any DT 10_1101-2020_12_31_424971 361 56 free free JJ 10_1101-2020_12_31_424971 361 57 mRNA mRNA NNS 10_1101-2020_12_31_424971 361 58 , , , 10_1101-2020_12_31_424971 361 59 IFs if NNS 10_1101-2020_12_31_424971 361 60 , , , 10_1101-2020_12_31_424971 361 61 and and CC 10_1101-2020_12_31_424971 361 62 824 824 CD 10_1101-2020_12_31_424971 361 63 fMet fMet NNP 10_1101-2020_12_31_424971 361 64 - - HYPH 10_1101-2020_12_31_424971 361 65 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 361 66 . . . 10_1101-2020_12_31_424971 362 1 Purified Purified NNP 10_1101-2020_12_31_424971 362 2 70S 70S NNP 10_1101-2020_12_31_424971 362 3 ICs ic NNS 10_1101-2020_12_31_424971 362 4 were be VBD 10_1101-2020_12_31_424971 362 5 aliquoted aliquote VBN 10_1101-2020_12_31_424971 362 6 , , , 10_1101-2020_12_31_424971 362 7 flash flash NNP 10_1101-2020_12_31_424971 362 8 frozen frozen JJ 10_1101-2020_12_31_424971 362 9 in in IN 10_1101-2020_12_31_424971 362 10 liquid liquid JJ 10_1101-2020_12_31_424971 362 11 nitrogen nitrogen NN 10_1101-2020_12_31_424971 362 12 , , , 10_1101-2020_12_31_424971 362 13 and and CC 10_1101-2020_12_31_424971 362 14 stored store VBD 10_1101-2020_12_31_424971 362 15 at at IN 10_1101-2020_12_31_424971 362 16 – – : 10_1101-2020_12_31_424971 362 17 80 80 CD 10_1101-2020_12_31_424971 362 18 825 825 CD 10_1101-2020_12_31_424971 362 19 ºC ºC NNP 10_1101-2020_12_31_424971 362 20 until until IN 10_1101-2020_12_31_424971 362 21 use use NN 10_1101-2020_12_31_424971 362 22 in in IN 10_1101-2020_12_31_424971 362 23 smFRET smfret NN 10_1101-2020_12_31_424971 362 24 experiments experiment NNS 10_1101-2020_12_31_424971 362 25 . . . 10_1101-2020_12_31_424971 363 1 826 826 CD 10_1101-2020_12_31_424971 363 2 TCs tc NNS 10_1101-2020_12_31_424971 363 3 were be VBD 10_1101-2020_12_31_424971 363 4 prepared prepare VBN 10_1101-2020_12_31_424971 363 5 in in IN 10_1101-2020_12_31_424971 363 6 two two CD 10_1101-2020_12_31_424971 363 7 steps step NNS 10_1101-2020_12_31_424971 363 8 . . . 10_1101-2020_12_31_424971 364 1 First first RB 10_1101-2020_12_31_424971 364 2 , , , 10_1101-2020_12_31_424971 364 3 300 300 CD 10_1101-2020_12_31_424971 364 4 pmol pmol NN 10_1101-2020_12_31_424971 364 5 of of IN 10_1101-2020_12_31_424971 364 6 EF EF NNP 10_1101-2020_12_31_424971 364 7 - - HYPH 10_1101-2020_12_31_424971 364 8 Tu Tu NNP 10_1101-2020_12_31_424971 364 9 and and CC 10_1101-2020_12_31_424971 364 10 200 200 CD 10_1101-2020_12_31_424971 364 11 pmol pmol NN 10_1101-2020_12_31_424971 364 12 of of IN 10_1101-2020_12_31_424971 364 13 EF EF NNP 10_1101-2020_12_31_424971 364 14 - - HYPH 10_1101-2020_12_31_424971 364 15 Ts Ts NNP 10_1101-2020_12_31_424971 364 16 in in IN 10_1101-2020_12_31_424971 364 17 8 8 CD 10_1101-2020_12_31_424971 364 18 µL µl CD 10_1101-2020_12_31_424971 364 19 of of IN 10_1101-2020_12_31_424971 364 20 827 827 CD 10_1101-2020_12_31_424971 364 21 Tris Tris NNP 10_1101-2020_12_31_424971 364 22 - - HYPH 10_1101-2020_12_31_424971 364 23 Polymix Polymix NNP 10_1101-2020_12_31_424971 364 24 Buffer Buffer NNP 10_1101-2020_12_31_424971 364 25 at at IN 10_1101-2020_12_31_424971 364 26 5 5 CD 10_1101-2020_12_31_424971 364 27 mM mm NN 10_1101-2020_12_31_424971 364 28 Mg(OAc)2 mg(oac)2 NN 10_1101-2020_12_31_424971 364 29 supplemented supplement VBN 10_1101-2020_12_31_424971 364 30 with with IN 10_1101-2020_12_31_424971 364 31 GTP GTP NNP 10_1101-2020_12_31_424971 364 32 Charging Charging NNP 10_1101-2020_12_31_424971 364 33 Components Components NNPS 10_1101-2020_12_31_424971 364 34 ( ( -LRB- 10_1101-2020_12_31_424971 364 35 1 1 CD 10_1101-2020_12_31_424971 364 36 mM mM NNP 10_1101-2020_12_31_424971 364 37 828 828 CD 10_1101-2020_12_31_424971 364 38 GTP GTP NNP 10_1101-2020_12_31_424971 364 39 , , , 10_1101-2020_12_31_424971 364 40 3 3 CD 10_1101-2020_12_31_424971 364 41 mM mM NNP 10_1101-2020_12_31_424971 364 42 phosphoenolpyruvate phosphoenolpyruvate NN 10_1101-2020_12_31_424971 364 43 , , , 10_1101-2020_12_31_424971 364 44 and and CC 10_1101-2020_12_31_424971 364 45 2 2 CD 10_1101-2020_12_31_424971 364 46 units unit NNS 10_1101-2020_12_31_424971 364 47 / / SYM 10_1101-2020_12_31_424971 364 48 mL mL NNP 10_1101-2020_12_31_424971 364 49 pyruvate pyruvate NNP 10_1101-2020_12_31_424971 364 50 kinase kinase NN 10_1101-2020_12_31_424971 364 51 ) ) -RRB- 10_1101-2020_12_31_424971 364 52 were be VBD 10_1101-2020_12_31_424971 364 53 incubated incubate VBN 10_1101-2020_12_31_424971 364 54 for for IN 10_1101-2020_12_31_424971 364 55 1 1 CD 10_1101-2020_12_31_424971 364 56 min min NN 10_1101-2020_12_31_424971 364 57 at at IN 10_1101-2020_12_31_424971 364 58 829 829 CD 10_1101-2020_12_31_424971 364 59 .CC .CC , 10_1101-2020_12_31_424971 364 60 - - HYPH 10_1101-2020_12_31_424971 364 61 BY by IN 10_1101-2020_12_31_424971 364 62 - - HYPH 10_1101-2020_12_31_424971 364 63 NC NC NNP 10_1101-2020_12_31_424971 364 64 - - HYPH 10_1101-2020_12_31_424971 364 65 ND ND NNP 10_1101-2020_12_31_424971 364 66 4.0 4.0 CD 10_1101-2020_12_31_424971 364 67 International International NNP 10_1101-2020_12_31_424971 364 68 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 364 69 under under IN 10_1101-2020_12_31_424971 364 70 a a DT 10_1101-2020_12_31_424971 364 71 ( ( -LRB- 10_1101-2020_12_31_424971 364 72 which which WDT 10_1101-2020_12_31_424971 364 73 was be VBD 10_1101-2020_12_31_424971 364 74 not not RB 10_1101-2020_12_31_424971 364 75 certified certify VBN 10_1101-2020_12_31_424971 364 76 by by IN 10_1101-2020_12_31_424971 364 77 peer peer NN 10_1101-2020_12_31_424971 364 78 review review NN 10_1101-2020_12_31_424971 364 79 ) ) -RRB- 10_1101-2020_12_31_424971 364 80 is be VBZ 10_1101-2020_12_31_424971 364 81 the the DT 10_1101-2020_12_31_424971 364 82 author author NN 10_1101-2020_12_31_424971 364 83 / / SYM 10_1101-2020_12_31_424971 364 84 funder funder NN 10_1101-2020_12_31_424971 364 85 , , , 10_1101-2020_12_31_424971 364 86 who who WP 10_1101-2020_12_31_424971 364 87 has have VBZ 10_1101-2020_12_31_424971 364 88 granted grant VBN 10_1101-2020_12_31_424971 364 89 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 364 90 a a DT 10_1101-2020_12_31_424971 364 91 license license NN 10_1101-2020_12_31_424971 364 92 to to TO 10_1101-2020_12_31_424971 364 93 display display VB 10_1101-2020_12_31_424971 364 94 the the DT 10_1101-2020_12_31_424971 364 95 preprint preprint NN 10_1101-2020_12_31_424971 364 96 in in IN 10_1101-2020_12_31_424971 364 97 perpetuity perpetuity NN 10_1101-2020_12_31_424971 364 98 . . . 10_1101-2020_12_31_424971 365 1 It -PRON- PRP 10_1101-2020_12_31_424971 365 2 is be VBZ 10_1101-2020_12_31_424971 365 3 made make VBN 10_1101-2020_12_31_424971 365 4 The the DT 10_1101-2020_12_31_424971 365 5 copyright copyright NN 10_1101-2020_12_31_424971 365 6 holder holder NN 10_1101-2020_12_31_424971 365 7 for for IN 10_1101-2020_12_31_424971 365 8 this this DT 10_1101-2020_12_31_424971 365 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 365 10 version version NN 10_1101-2020_12_31_424971 365 11 posted post VBD 10_1101-2020_12_31_424971 365 12 January January NNP 10_1101-2020_12_31_424971 365 13 2 2 CD 10_1101-2020_12_31_424971 365 14 , , , 10_1101-2020_12_31_424971 365 15 2021 2021 CD 10_1101-2020_12_31_424971 365 16 . . . 10_1101-2020_12_31_424971 365 17 ; ; : 10_1101-2020_12_31_424971 365 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 365 19 : : : 10_1101-2020_12_31_424971 365 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 365 21 preprint preprint NN 10_1101-2020_12_31_424971 365 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 365 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 365 24 34 34 CD 10_1101-2020_12_31_424971 365 25 37 37 CD 10_1101-2020_12_31_424971 365 26 ºC. ºC. NNS 10_1101-2020_12_31_424971 366 1 Then then RB 10_1101-2020_12_31_424971 366 2 , , , 10_1101-2020_12_31_424971 366 3 30 30 CD 10_1101-2020_12_31_424971 366 4 pmol pmol NN 10_1101-2020_12_31_424971 366 5 of of IN 10_1101-2020_12_31_424971 366 6 aa aa NNP 10_1101-2020_12_31_424971 366 7 - - HYPH 10_1101-2020_12_31_424971 366 8 tRNA tRNA NNP 10_1101-2020_12_31_424971 366 9 in in IN 10_1101-2020_12_31_424971 366 10 2 2 CD 10_1101-2020_12_31_424971 366 11 µL µl CD 10_1101-2020_12_31_424971 366 12 of of IN 10_1101-2020_12_31_424971 366 13 25 25 CD 10_1101-2020_12_31_424971 366 14 mM mM NNP 10_1101-2020_12_31_424971 366 15 NaOAc NaOAc NNS 10_1101-2020_12_31_424971 366 16 ( ( -LRB- 10_1101-2020_12_31_424971 366 17 pH pH NNP 10_1101-2020_12_31_424971 366 18 = = SYM 10_1101-2020_12_31_424971 366 19 5 5 CD 10_1101-2020_12_31_424971 366 20 ) ) -RRB- 10_1101-2020_12_31_424971 366 21 was be VBD 10_1101-2020_12_31_424971 366 22 added add VBN 10_1101-2020_12_31_424971 366 23 to to IN 10_1101-2020_12_31_424971 366 24 the the DT 10_1101-2020_12_31_424971 366 25 reaction reaction NN 10_1101-2020_12_31_424971 366 26 , , , 10_1101-2020_12_31_424971 366 27 830 830 CD 10_1101-2020_12_31_424971 366 28 followed follow VBN 10_1101-2020_12_31_424971 366 29 by by IN 10_1101-2020_12_31_424971 366 30 an an DT 10_1101-2020_12_31_424971 366 31 additional additional JJ 10_1101-2020_12_31_424971 366 32 incubation incubation NN 10_1101-2020_12_31_424971 366 33 of of IN 10_1101-2020_12_31_424971 366 34 1 1 CD 10_1101-2020_12_31_424971 366 35 min min NN 10_1101-2020_12_31_424971 366 36 at at IN 10_1101-2020_12_31_424971 366 37 37 37 CD 10_1101-2020_12_31_424971 366 38 ºC. ºC. NNS 10_1101-2020_12_31_424971 367 1 This this DT 10_1101-2020_12_31_424971 367 2 results result VBZ 10_1101-2020_12_31_424971 367 3 in in IN 10_1101-2020_12_31_424971 367 4 a a DT 10_1101-2020_12_31_424971 367 5 TC TC NNP 10_1101-2020_12_31_424971 367 6 solution solution NN 10_1101-2020_12_31_424971 367 7 with with IN 10_1101-2020_12_31_424971 367 8 a a DT 10_1101-2020_12_31_424971 367 9 final final JJ 10_1101-2020_12_31_424971 367 10 831 831 CD 10_1101-2020_12_31_424971 367 11 volume volume NN 10_1101-2020_12_31_424971 367 12 of of IN 10_1101-2020_12_31_424971 367 13 10 10 CD 10_1101-2020_12_31_424971 367 14 µL µL . 10_1101-2020_12_31_424971 367 15 that that WDT 10_1101-2020_12_31_424971 367 16 was be VBD 10_1101-2020_12_31_424971 367 17 then then RB 10_1101-2020_12_31_424971 367 18 stored store VBN 10_1101-2020_12_31_424971 367 19 on on IN 10_1101-2020_12_31_424971 367 20 ice ice NN 10_1101-2020_12_31_424971 367 21 until until IN 10_1101-2020_12_31_424971 367 22 used use VBN 10_1101-2020_12_31_424971 367 23 for for IN 10_1101-2020_12_31_424971 367 24 smFRET smfret NN 10_1101-2020_12_31_424971 367 25 experiments experiment NNS 10_1101-2020_12_31_424971 367 26 . . . 10_1101-2020_12_31_424971 368 1 832 832 CD 10_1101-2020_12_31_424971 368 2 To to TO 10_1101-2020_12_31_424971 368 3 prepare prepare VB 10_1101-2020_12_31_424971 368 4 PRE PRE NNP 10_1101-2020_12_31_424971 368 5 – – : 10_1101-2020_12_31_424971 368 6 A a DT 10_1101-2020_12_31_424971 368 7 complexes complex NNS 10_1101-2020_12_31_424971 368 8 , , , 10_1101-2020_12_31_424971 368 9 we -PRON- PRP 10_1101-2020_12_31_424971 368 10 first first RB 10_1101-2020_12_31_424971 368 11 needed need VBD 10_1101-2020_12_31_424971 368 12 to to TO 10_1101-2020_12_31_424971 368 13 assemble assemble VB 10_1101-2020_12_31_424971 368 14 POST POST NNP 10_1101-2020_12_31_424971 368 15 complexes complex NNS 10_1101-2020_12_31_424971 368 16 . . . 10_1101-2020_12_31_424971 369 1 POST POST NNP 10_1101-2020_12_31_424971 369 2 833 833 CD 10_1101-2020_12_31_424971 369 3 complexes complex NNS 10_1101-2020_12_31_424971 369 4 were be VBD 10_1101-2020_12_31_424971 369 5 assembled assemble VBN 10_1101-2020_12_31_424971 369 6 by by IN 10_1101-2020_12_31_424971 369 7 first first RB 10_1101-2020_12_31_424971 369 8 preparing prepare VBG 10_1101-2020_12_31_424971 369 9 a a DT 10_1101-2020_12_31_424971 369 10 10-µL 10-µL NNS 10_1101-2020_12_31_424971 369 11 solution solution NN 10_1101-2020_12_31_424971 369 12 of of IN 10_1101-2020_12_31_424971 369 13 70S 70S NNP 10_1101-2020_12_31_424971 369 14 IC IC NNP 10_1101-2020_12_31_424971 369 15 and and CC 10_1101-2020_12_31_424971 369 16 a a DT 10_1101-2020_12_31_424971 369 17 10-µL 10-µl JJ 10_1101-2020_12_31_424971 369 18 solution solution NN 10_1101-2020_12_31_424971 369 19 of of IN 10_1101-2020_12_31_424971 369 20 834 834 CD 10_1101-2020_12_31_424971 369 21 TC TC NNP 10_1101-2020_12_31_424971 369 22 as as IN 10_1101-2020_12_31_424971 369 23 described describe VBN 10_1101-2020_12_31_424971 369 24 above above RB 10_1101-2020_12_31_424971 369 25 . . . 10_1101-2020_12_31_424971 370 1 Separately separately RB 10_1101-2020_12_31_424971 370 2 , , , 10_1101-2020_12_31_424971 370 3 a a DT 10_1101-2020_12_31_424971 370 4 solution solution NN 10_1101-2020_12_31_424971 370 5 of of IN 10_1101-2020_12_31_424971 370 6 GTP GTP NNP 10_1101-2020_12_31_424971 370 7 - - HYPH 10_1101-2020_12_31_424971 370 8 bound bind VBN 10_1101-2020_12_31_424971 370 9 EF EF NNP 10_1101-2020_12_31_424971 370 10 - - HYPH 10_1101-2020_12_31_424971 370 11 G G NNP 10_1101-2020_12_31_424971 370 12 was be VBD 10_1101-2020_12_31_424971 370 13 prepared prepare VBN 10_1101-2020_12_31_424971 370 14 by by IN 10_1101-2020_12_31_424971 370 15 incubating incubate VBG 10_1101-2020_12_31_424971 370 16 835 835 CD 10_1101-2020_12_31_424971 370 17 120 120 CD 10_1101-2020_12_31_424971 370 18 pmol pmol NN 10_1101-2020_12_31_424971 370 19 EF EF NNP 10_1101-2020_12_31_424971 370 20 - - HYPH 10_1101-2020_12_31_424971 370 21 G G NNP 10_1101-2020_12_31_424971 370 22 in in IN 10_1101-2020_12_31_424971 370 23 5 5 CD 10_1101-2020_12_31_424971 370 24 µL µl CD 10_1101-2020_12_31_424971 370 25 of of IN 10_1101-2020_12_31_424971 370 26 Tris Tris NNP 10_1101-2020_12_31_424971 370 27 - - HYPH 10_1101-2020_12_31_424971 370 28 Polymix Polymix NNP 10_1101-2020_12_31_424971 370 29 Buffer Buffer NNP 10_1101-2020_12_31_424971 370 30 at at IN 10_1101-2020_12_31_424971 370 31 5 5 CD 10_1101-2020_12_31_424971 370 32 mM mm NN 10_1101-2020_12_31_424971 370 33 Mg(OAc)2 mg(oac)2 NN 10_1101-2020_12_31_424971 370 34 supplemented supplement VBN 10_1101-2020_12_31_424971 370 35 with with IN 10_1101-2020_12_31_424971 370 36 GTP GTP NNP 10_1101-2020_12_31_424971 370 37 836 836 CD 10_1101-2020_12_31_424971 370 38 Charging charge VBG 10_1101-2020_12_31_424971 370 39 Components Components NNPS 10_1101-2020_12_31_424971 370 40 for for IN 10_1101-2020_12_31_424971 370 41 2 2 CD 10_1101-2020_12_31_424971 370 42 min min NN 10_1101-2020_12_31_424971 370 43 at at IN 10_1101-2020_12_31_424971 370 44 room room NN 10_1101-2020_12_31_424971 370 45 temperature temperature NN 10_1101-2020_12_31_424971 370 46 . . . 10_1101-2020_12_31_424971 371 1 Then then RB 10_1101-2020_12_31_424971 371 2 10 10 CD 10_1101-2020_12_31_424971 371 3 µL µl CD 10_1101-2020_12_31_424971 371 4 of of IN 10_1101-2020_12_31_424971 371 5 the the DT 10_1101-2020_12_31_424971 371 6 70S 70S NNP 10_1101-2020_12_31_424971 371 7 IC IC NNP 10_1101-2020_12_31_424971 371 8 , , , 10_1101-2020_12_31_424971 371 9 10 10 CD 10_1101-2020_12_31_424971 371 10 µL µl CD 10_1101-2020_12_31_424971 371 11 of of IN 10_1101-2020_12_31_424971 371 12 the the DT 10_1101-2020_12_31_424971 371 13 TC TC NNP 10_1101-2020_12_31_424971 371 14 , , , 10_1101-2020_12_31_424971 371 15 837 837 CD 10_1101-2020_12_31_424971 371 16 and and CC 10_1101-2020_12_31_424971 371 17 2.5 2.5 CD 10_1101-2020_12_31_424971 371 18 µL µL NNP 10_1101-2020_12_31_424971 371 19 the the DT 10_1101-2020_12_31_424971 371 20 GTP GTP NNP 10_1101-2020_12_31_424971 371 21 - - HYPH 10_1101-2020_12_31_424971 371 22 bound bind VBN 10_1101-2020_12_31_424971 371 23 EF EF NNP 10_1101-2020_12_31_424971 371 24 - - HYPH 10_1101-2020_12_31_424971 371 25 G G NNP 10_1101-2020_12_31_424971 371 26 solution solution NN 10_1101-2020_12_31_424971 371 27 were be VBD 10_1101-2020_12_31_424971 371 28 mixed mixed JJ 10_1101-2020_12_31_424971 371 29 , , , 10_1101-2020_12_31_424971 371 30 and and CC 10_1101-2020_12_31_424971 371 31 incubated incubate VBD 10_1101-2020_12_31_424971 371 32 for for IN 10_1101-2020_12_31_424971 371 33 5 5 CD 10_1101-2020_12_31_424971 371 34 min min NN 10_1101-2020_12_31_424971 371 35 at at IN 10_1101-2020_12_31_424971 371 36 room room NN 10_1101-2020_12_31_424971 371 37 838 838 CD 10_1101-2020_12_31_424971 371 38 temperature temperature NN 10_1101-2020_12_31_424971 371 39 and and CC 10_1101-2020_12_31_424971 371 40 for for IN 10_1101-2020_12_31_424971 371 41 additional additional JJ 10_1101-2020_12_31_424971 371 42 5 5 CD 10_1101-2020_12_31_424971 371 43 min min NN 10_1101-2020_12_31_424971 371 44 on on IN 10_1101-2020_12_31_424971 371 45 ice ice NN 10_1101-2020_12_31_424971 371 46 . . . 10_1101-2020_12_31_424971 372 1 The the DT 10_1101-2020_12_31_424971 372 2 resulting result VBG 10_1101-2020_12_31_424971 372 3 POST POST NNP 10_1101-2020_12_31_424971 372 4 complex complex NN 10_1101-2020_12_31_424971 372 5 was be VBD 10_1101-2020_12_31_424971 372 6 diluted dilute VBN 10_1101-2020_12_31_424971 372 7 by by IN 10_1101-2020_12_31_424971 372 8 adjusting adjust VBG 10_1101-2020_12_31_424971 372 9 839 839 CD 10_1101-2020_12_31_424971 372 10 the the DT 10_1101-2020_12_31_424971 372 11 reaction reaction NN 10_1101-2020_12_31_424971 372 12 volume volume NN 10_1101-2020_12_31_424971 372 13 to to IN 10_1101-2020_12_31_424971 372 14 100 100 CD 10_1101-2020_12_31_424971 372 15 µL µL NNP 10_1101-2020_12_31_424971 372 16 with with IN 10_1101-2020_12_31_424971 372 17 Tris Tris NNP 10_1101-2020_12_31_424971 372 18 - - HYPH 10_1101-2020_12_31_424971 372 19 Polymix Polymix NNP 10_1101-2020_12_31_424971 372 20 Buffer Buffer NNP 10_1101-2020_12_31_424971 372 21 at at IN 10_1101-2020_12_31_424971 372 22 20 20 CD 10_1101-2020_12_31_424971 372 23 mM mm NN 10_1101-2020_12_31_424971 372 24 Mg(OAc)2 Mg(OAc)2 NNS 10_1101-2020_12_31_424971 372 25 and and CC 10_1101-2020_12_31_424971 372 26 purified purify VBD 10_1101-2020_12_31_424971 372 27 via via IN 10_1101-2020_12_31_424971 372 28 840 840 CD 10_1101-2020_12_31_424971 372 29 sucrose sucrose JJ 10_1101-2020_12_31_424971 372 30 density density NN 10_1101-2020_12_31_424971 372 31 gradient gradient NN 10_1101-2020_12_31_424971 372 32 ultracentrifugation ultracentrifugation NN 10_1101-2020_12_31_424971 372 33 as as IN 10_1101-2020_12_31_424971 372 34 described describe VBN 10_1101-2020_12_31_424971 372 35 above above RB 10_1101-2020_12_31_424971 372 36 for for IN 10_1101-2020_12_31_424971 372 37 the the DT 10_1101-2020_12_31_424971 372 38 70S 70S NNP 10_1101-2020_12_31_424971 372 39 ICs ic NNS 10_1101-2020_12_31_424971 372 40 . . . 10_1101-2020_12_31_424971 373 1 Purified Purified NNP 10_1101-2020_12_31_424971 373 2 POST POST NNP 10_1101-2020_12_31_424971 373 3 841 841 CD 10_1101-2020_12_31_424971 373 4 complexes complex NNS 10_1101-2020_12_31_424971 373 5 were be VBD 10_1101-2020_12_31_424971 373 6 aliquoted aliquote VBN 10_1101-2020_12_31_424971 373 7 , , , 10_1101-2020_12_31_424971 373 8 flash flash NNP 10_1101-2020_12_31_424971 373 9 frozen frozen JJ 10_1101-2020_12_31_424971 373 10 in in IN 10_1101-2020_12_31_424971 373 11 liquid liquid JJ 10_1101-2020_12_31_424971 373 12 nitrogen nitrogen NN 10_1101-2020_12_31_424971 373 13 , , , 10_1101-2020_12_31_424971 373 14 and and CC 10_1101-2020_12_31_424971 373 15 stored store VBD 10_1101-2020_12_31_424971 373 16 at at IN 10_1101-2020_12_31_424971 373 17 – – : 10_1101-2020_12_31_424971 373 18 80 80 CD 10_1101-2020_12_31_424971 373 19 ºC ºC NNP 10_1101-2020_12_31_424971 373 20 until until IN 10_1101-2020_12_31_424971 373 21 use use NN 10_1101-2020_12_31_424971 373 22 in in IN 10_1101-2020_12_31_424971 373 23 842 842 CD 10_1101-2020_12_31_424971 373 24 smFRET smfret NN 10_1101-2020_12_31_424971 373 25 experiments experiment NNS 10_1101-2020_12_31_424971 373 26 . . . 10_1101-2020_12_31_424971 374 1 PRE PRE NNP 10_1101-2020_12_31_424971 374 2 – – : 10_1101-2020_12_31_424971 374 3 A a DT 10_1101-2020_12_31_424971 374 4 complexes complex NNS 10_1101-2020_12_31_424971 374 5 were be VBD 10_1101-2020_12_31_424971 374 6 then then RB 10_1101-2020_12_31_424971 374 7 generated generate VBN 10_1101-2020_12_31_424971 374 8 by by IN 10_1101-2020_12_31_424971 374 9 mixing mix VBG 10_1101-2020_12_31_424971 374 10 3 3 CD 10_1101-2020_12_31_424971 374 11 µL µL NNP 10_1101-2020_12_31_424971 374 12 of of IN 10_1101-2020_12_31_424971 374 13 POST POST NNP 10_1101-2020_12_31_424971 374 14 complex complex NN 10_1101-2020_12_31_424971 374 15 , , , 10_1101-2020_12_31_424971 374 16 843 843 CD 10_1101-2020_12_31_424971 374 17 2 2 CD 10_1101-2020_12_31_424971 374 18 µL µl CD 10_1101-2020_12_31_424971 374 19 of of IN 10_1101-2020_12_31_424971 374 20 a a DT 10_1101-2020_12_31_424971 374 21 10 10 CD 10_1101-2020_12_31_424971 374 22 mM mm NN 10_1101-2020_12_31_424971 374 23 puromycin puromycin JJ 10_1101-2020_12_31_424971 374 24 solution solution NN 10_1101-2020_12_31_424971 374 25 ( ( -LRB- 10_1101-2020_12_31_424971 374 26 prepared prepare VBN 10_1101-2020_12_31_424971 374 27 in in IN 10_1101-2020_12_31_424971 374 28 Nanopure Nanopure NNP 10_1101-2020_12_31_424971 374 29 water water NN 10_1101-2020_12_31_424971 374 30 and and CC 10_1101-2020_12_31_424971 374 31 filtered filter VBD 10_1101-2020_12_31_424971 374 32 using use VBG 10_1101-2020_12_31_424971 374 33 a a DT 10_1101-2020_12_31_424971 374 34 0.22 0.22 CD 10_1101-2020_12_31_424971 374 35 µm µm NNP 10_1101-2020_12_31_424971 374 36 844 844 CD 10_1101-2020_12_31_424971 374 37 filter filter NN 10_1101-2020_12_31_424971 374 38 ) ) -RRB- 10_1101-2020_12_31_424971 374 39 , , , 10_1101-2020_12_31_424971 374 40 and and CC 10_1101-2020_12_31_424971 374 41 15 15 CD 10_1101-2020_12_31_424971 374 42 µL µl CD 10_1101-2020_12_31_424971 374 43 of of IN 10_1101-2020_12_31_424971 374 44 Tris Tris NNP 10_1101-2020_12_31_424971 374 45 - - HYPH 10_1101-2020_12_31_424971 374 46 Polymix Polymix NNP 10_1101-2020_12_31_424971 374 47 Buffer Buffer NNP 10_1101-2020_12_31_424971 374 48 at at IN 10_1101-2020_12_31_424971 374 49 15 15 CD 10_1101-2020_12_31_424971 374 50 mM mm NN 10_1101-2020_12_31_424971 374 51 Mg(OAc)2 Mg(OAc)2 NNS 10_1101-2020_12_31_424971 374 52 and and CC 10_1101-2020_12_31_424971 374 53 incubating incubate VBG 10_1101-2020_12_31_424971 374 54 the the DT 10_1101-2020_12_31_424971 374 55 mixture mixture NN 10_1101-2020_12_31_424971 374 56 for for IN 10_1101-2020_12_31_424971 374 57 10 10 CD 10_1101-2020_12_31_424971 374 58 min min NN 10_1101-2020_12_31_424971 374 59 845 845 CD 10_1101-2020_12_31_424971 374 60 at at IN 10_1101-2020_12_31_424971 374 61 room room NN 10_1101-2020_12_31_424971 374 62 temperature temperature NN 10_1101-2020_12_31_424971 374 63 . . . 10_1101-2020_12_31_424971 375 1 PRE PRE NNP 10_1101-2020_12_31_424971 375 2 – – : 10_1101-2020_12_31_424971 375 3 A a DT 10_1101-2020_12_31_424971 375 4 complexes complex NNS 10_1101-2020_12_31_424971 375 5 were be VBD 10_1101-2020_12_31_424971 375 6 used use VBN 10_1101-2020_12_31_424971 375 7 for for IN 10_1101-2020_12_31_424971 375 8 smFRET smfret NN 10_1101-2020_12_31_424971 375 9 experiments experiment NNS 10_1101-2020_12_31_424971 375 10 immediately immediately RB 10_1101-2020_12_31_424971 375 11 upon upon IN 10_1101-2020_12_31_424971 375 12 846 846 CD 10_1101-2020_12_31_424971 375 13 preparation preparation NN 10_1101-2020_12_31_424971 375 14 . . . 10_1101-2020_12_31_424971 376 1 847 847 CD 10_1101-2020_12_31_424971 376 2 848 848 CD 10_1101-2020_12_31_424971 376 3 smFRET smFRET NNP 10_1101-2020_12_31_424971 376 4 imaging imaging NN 10_1101-2020_12_31_424971 376 5 using use VBG 10_1101-2020_12_31_424971 376 6 total total JJ 10_1101-2020_12_31_424971 376 7 internal internal JJ 10_1101-2020_12_31_424971 376 8 reflection reflection NN 10_1101-2020_12_31_424971 376 9 fluorescence fluorescence NN 10_1101-2020_12_31_424971 376 10 ( ( -LRB- 10_1101-2020_12_31_424971 376 11 TIRF TIRF NNP 10_1101-2020_12_31_424971 376 12 ) ) -RRB- 10_1101-2020_12_31_424971 376 13 microscopy microscopy JJ 10_1101-2020_12_31_424971 376 14 . . . 10_1101-2020_12_31_424971 377 1 70S 70s CD 10_1101-2020_12_31_424971 377 2 ICs ic NNS 10_1101-2020_12_31_424971 377 3 or or CC 10_1101-2020_12_31_424971 377 4 849 849 CD 10_1101-2020_12_31_424971 377 5 PRE PRE NNP 10_1101-2020_12_31_424971 377 6 – – : 10_1101-2020_12_31_424971 377 7 A a DT 10_1101-2020_12_31_424971 377 8 complexes complex NNS 10_1101-2020_12_31_424971 377 9 were be VBD 10_1101-2020_12_31_424971 377 10 tethered tether VBN 10_1101-2020_12_31_424971 377 11 to to IN 10_1101-2020_12_31_424971 377 12 the the DT 10_1101-2020_12_31_424971 377 13 PEG peg NN 10_1101-2020_12_31_424971 377 14 / / SYM 10_1101-2020_12_31_424971 377 15 biotin biotin NN 10_1101-2020_12_31_424971 377 16 - - HYPH 10_1101-2020_12_31_424971 377 17 PEG peg NN 10_1101-2020_12_31_424971 377 18 - - HYPH 10_1101-2020_12_31_424971 377 19 passivated passivate VBN 10_1101-2020_12_31_424971 377 20 and and CC 10_1101-2020_12_31_424971 377 21 streptavidin streptavidin NNP 10_1101-2020_12_31_424971 377 22 - - HYPH 10_1101-2020_12_31_424971 377 23 derivatized derivatize VBN 10_1101-2020_12_31_424971 377 24 850 850 CD 10_1101-2020_12_31_424971 377 25 surface surface NN 10_1101-2020_12_31_424971 377 26 of of IN 10_1101-2020_12_31_424971 377 27 a a DT 10_1101-2020_12_31_424971 377 28 quartz quartz NN 10_1101-2020_12_31_424971 377 29 microfluidic microfluidic NN 10_1101-2020_12_31_424971 377 30 flowcell flowcell NN 10_1101-2020_12_31_424971 377 31 via via IN 10_1101-2020_12_31_424971 377 32 a a DT 10_1101-2020_12_31_424971 377 33 biotin biotin NN 10_1101-2020_12_31_424971 377 34 - - HYPH 10_1101-2020_12_31_424971 377 35 streptavidin streptavidin NN 10_1101-2020_12_31_424971 377 36 - - HYPH 10_1101-2020_12_31_424971 377 37 biotin biotin JJ 10_1101-2020_12_31_424971 377 38 bridge bridge NN 10_1101-2020_12_31_424971 377 39 between between IN 10_1101-2020_12_31_424971 377 40 the the DT 10_1101-2020_12_31_424971 377 41 biotin-851 biotin-851 NN 10_1101-2020_12_31_424971 377 42 mRNA mRNA NNS 10_1101-2020_12_31_424971 377 43 and and CC 10_1101-2020_12_31_424971 377 44 the the DT 10_1101-2020_12_31_424971 377 45 biotin biotin NNP 10_1101-2020_12_31_424971 377 46 - - HYPH 10_1101-2020_12_31_424971 377 47 PEG37,43 PEG37,43 NNS 10_1101-2020_12_31_424971 377 48 . . . 10_1101-2020_12_31_424971 378 1 Untethered Untethered NNP 10_1101-2020_12_31_424971 378 2 70S 70S NNP 10_1101-2020_12_31_424971 378 3 ICs ic NNS 10_1101-2020_12_31_424971 378 4 or or CC 10_1101-2020_12_31_424971 378 5 PRE PRE NNP 10_1101-2020_12_31_424971 378 6 – – : 10_1101-2020_12_31_424971 378 7 A a DT 10_1101-2020_12_31_424971 378 8 complexes complex NNS 10_1101-2020_12_31_424971 378 9 were be VBD 10_1101-2020_12_31_424971 378 10 removed remove VBN 10_1101-2020_12_31_424971 378 11 from from IN 10_1101-2020_12_31_424971 378 12 the the DT 10_1101-2020_12_31_424971 378 13 852 852 CD 10_1101-2020_12_31_424971 378 14 flowcell flowcell NN 10_1101-2020_12_31_424971 378 15 , , , 10_1101-2020_12_31_424971 378 16 and and CC 10_1101-2020_12_31_424971 378 17 the the DT 10_1101-2020_12_31_424971 378 18 flowcell flowcell NN 10_1101-2020_12_31_424971 378 19 was be VBD 10_1101-2020_12_31_424971 378 20 prepared prepare VBN 10_1101-2020_12_31_424971 378 21 for for IN 10_1101-2020_12_31_424971 378 22 smFRET smfret NN 10_1101-2020_12_31_424971 378 23 imaging image VBG 10_1101-2020_12_31_424971 378 24 experiments experiment NNS 10_1101-2020_12_31_424971 378 25 , , , 10_1101-2020_12_31_424971 378 26 by by IN 10_1101-2020_12_31_424971 378 27 flushing flush VBG 10_1101-2020_12_31_424971 378 28 it -PRON- PRP 10_1101-2020_12_31_424971 378 29 with with IN 10_1101-2020_12_31_424971 378 30 Tris-853 tris-853 CD 10_1101-2020_12_31_424971 378 31 Polymix Polymix NNP 10_1101-2020_12_31_424971 378 32 Buffer Buffer NNP 10_1101-2020_12_31_424971 378 33 at at IN 10_1101-2020_12_31_424971 378 34 15 15 CD 10_1101-2020_12_31_424971 378 35 mM mm NN 10_1101-2020_12_31_424971 378 36 Mg(OAc)2 mg(oac)2 NN 10_1101-2020_12_31_424971 378 37 supplemented supplement VBN 10_1101-2020_12_31_424971 378 38 with with IN 10_1101-2020_12_31_424971 378 39 an an DT 10_1101-2020_12_31_424971 378 40 Oxygen oxygen NN 10_1101-2020_12_31_424971 378 41 - - HYPH 10_1101-2020_12_31_424971 378 42 Scavenging scavenge VBG 10_1101-2020_12_31_424971 378 43 System system NN 10_1101-2020_12_31_424971 378 44 ( ( -LRB- 10_1101-2020_12_31_424971 378 45 2.5 2.5 CD 10_1101-2020_12_31_424971 378 46 mM mM NNP 10_1101-2020_12_31_424971 378 47 854 854 CD 10_1101-2020_12_31_424971 378 48 protocatechuic protocatechuic NNS 10_1101-2020_12_31_424971 378 49 acid acid NN 10_1101-2020_12_31_424971 378 50 ( ( -LRB- 10_1101-2020_12_31_424971 378 51 pH pH NNP 10_1101-2020_12_31_424971 378 52 = = SYM 10_1101-2020_12_31_424971 378 53 9 9 CD 10_1101-2020_12_31_424971 378 54 ) ) -RRB- 10_1101-2020_12_31_424971 378 55 ( ( -LRB- 10_1101-2020_12_31_424971 378 56 Sigma Sigma NNP 10_1101-2020_12_31_424971 378 57 Aldrich Aldrich NNP 10_1101-2020_12_31_424971 378 58 ) ) -RRB- 10_1101-2020_12_31_424971 378 59 and and CC 10_1101-2020_12_31_424971 378 60 250 250 CD 10_1101-2020_12_31_424971 378 61 nM nM NNP 10_1101-2020_12_31_424971 378 62 protocatechuate-3,4-dioxygenase protocatechuate-3,4-dioxygenase NNP 10_1101-2020_12_31_424971 378 63 ( ( -LRB- 10_1101-2020_12_31_424971 378 64 pH pH NNP 10_1101-2020_12_31_424971 378 65 855 855 CD 10_1101-2020_12_31_424971 378 66 .CC .cc SYM 10_1101-2020_12_31_424971 378 67 - - HYPH 10_1101-2020_12_31_424971 378 68 BY by IN 10_1101-2020_12_31_424971 378 69 - - HYPH 10_1101-2020_12_31_424971 378 70 NC NC NNP 10_1101-2020_12_31_424971 378 71 - - HYPH 10_1101-2020_12_31_424971 378 72 ND ND NNP 10_1101-2020_12_31_424971 378 73 4.0 4.0 CD 10_1101-2020_12_31_424971 378 74 International International NNP 10_1101-2020_12_31_424971 378 75 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 378 76 under under IN 10_1101-2020_12_31_424971 378 77 a a DT 10_1101-2020_12_31_424971 378 78 ( ( -LRB- 10_1101-2020_12_31_424971 378 79 which which WDT 10_1101-2020_12_31_424971 378 80 was be VBD 10_1101-2020_12_31_424971 378 81 not not RB 10_1101-2020_12_31_424971 378 82 certified certify VBN 10_1101-2020_12_31_424971 378 83 by by IN 10_1101-2020_12_31_424971 378 84 peer peer NN 10_1101-2020_12_31_424971 378 85 review review NN 10_1101-2020_12_31_424971 378 86 ) ) -RRB- 10_1101-2020_12_31_424971 378 87 is be VBZ 10_1101-2020_12_31_424971 378 88 the the DT 10_1101-2020_12_31_424971 378 89 author author NN 10_1101-2020_12_31_424971 378 90 / / SYM 10_1101-2020_12_31_424971 378 91 funder funder NN 10_1101-2020_12_31_424971 378 92 , , , 10_1101-2020_12_31_424971 378 93 who who WP 10_1101-2020_12_31_424971 378 94 has have VBZ 10_1101-2020_12_31_424971 378 95 granted grant VBN 10_1101-2020_12_31_424971 378 96 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 378 97 a a DT 10_1101-2020_12_31_424971 378 98 license license NN 10_1101-2020_12_31_424971 378 99 to to TO 10_1101-2020_12_31_424971 378 100 display display VB 10_1101-2020_12_31_424971 378 101 the the DT 10_1101-2020_12_31_424971 378 102 preprint preprint NN 10_1101-2020_12_31_424971 378 103 in in IN 10_1101-2020_12_31_424971 378 104 perpetuity perpetuity NN 10_1101-2020_12_31_424971 378 105 . . . 10_1101-2020_12_31_424971 379 1 It -PRON- PRP 10_1101-2020_12_31_424971 379 2 is be VBZ 10_1101-2020_12_31_424971 379 3 made make VBN 10_1101-2020_12_31_424971 379 4 The the DT 10_1101-2020_12_31_424971 379 5 copyright copyright NN 10_1101-2020_12_31_424971 379 6 holder holder NN 10_1101-2020_12_31_424971 379 7 for for IN 10_1101-2020_12_31_424971 379 8 this this DT 10_1101-2020_12_31_424971 379 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 379 10 version version NN 10_1101-2020_12_31_424971 379 11 posted post VBD 10_1101-2020_12_31_424971 379 12 January January NNP 10_1101-2020_12_31_424971 379 13 2 2 CD 10_1101-2020_12_31_424971 379 14 , , , 10_1101-2020_12_31_424971 379 15 2021 2021 CD 10_1101-2020_12_31_424971 379 16 . . . 10_1101-2020_12_31_424971 379 17 ; ; : 10_1101-2020_12_31_424971 379 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 379 19 : : : 10_1101-2020_12_31_424971 379 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 379 21 preprint preprint NN 10_1101-2020_12_31_424971 379 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 379 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 379 24 35 35 CD 10_1101-2020_12_31_424971 379 25 = = SYM 10_1101-2020_12_31_424971 379 26 7.8 7.8 CD 10_1101-2020_12_31_424971 379 27 ) ) -RRB- 10_1101-2020_12_31_424971 379 28 ( ( -LRB- 10_1101-2020_12_31_424971 379 29 Sigma Sigma NNP 10_1101-2020_12_31_424971 379 30 Aldrich))64 Aldrich))64 NNP 10_1101-2020_12_31_424971 379 31 and and CC 10_1101-2020_12_31_424971 379 32 a a DT 10_1101-2020_12_31_424971 379 33 Triplet Triplet NNP 10_1101-2020_12_31_424971 379 34 - - HYPH 10_1101-2020_12_31_424971 379 35 State State NNP 10_1101-2020_12_31_424971 379 36 - - HYPH 10_1101-2020_12_31_424971 379 37 Quencher Quencher NNP 10_1101-2020_12_31_424971 379 38 Cocktail Cocktail NNP 10_1101-2020_12_31_424971 379 39 ( ( -LRB- 10_1101-2020_12_31_424971 379 40 1 1 CD 10_1101-2020_12_31_424971 379 41 mM mM NNP 10_1101-2020_12_31_424971 379 42 1,3,5,7-cyclooctatetraene 1,3,5,7-cyclooctatetraene CD 10_1101-2020_12_31_424971 379 43 856 856 CD 10_1101-2020_12_31_424971 379 44 ( ( -LRB- 10_1101-2020_12_31_424971 379 45 Aldrich Aldrich NNP 10_1101-2020_12_31_424971 379 46 ) ) -RRB- 10_1101-2020_12_31_424971 379 47 and and CC 10_1101-2020_12_31_424971 379 48 1 1 CD 10_1101-2020_12_31_424971 379 49 mM mm NN 10_1101-2020_12_31_424971 379 50 3-nitrobenzyl 3-nitrobenzyl CD 10_1101-2020_12_31_424971 379 51 alcohol alcohol NN 10_1101-2020_12_31_424971 379 52 ( ( -LRB- 10_1101-2020_12_31_424971 379 53 Fluka))65 Fluka))65 NNP 10_1101-2020_12_31_424971 379 54 . . . 10_1101-2020_12_31_424971 380 1 857 857 CD 10_1101-2020_12_31_424971 380 2 Tethered Tethered NNP 10_1101-2020_12_31_424971 380 3 70S 70S NNP 10_1101-2020_12_31_424971 380 4 ICs ic NNS 10_1101-2020_12_31_424971 380 5 or or CC 10_1101-2020_12_31_424971 380 6 PRE PRE NNP 10_1101-2020_12_31_424971 380 7 – – : 10_1101-2020_12_31_424971 380 8 A a DT 10_1101-2020_12_31_424971 380 9 complexes complex NNS 10_1101-2020_12_31_424971 380 10 were be VBD 10_1101-2020_12_31_424971 380 11 imaged image VBN 10_1101-2020_12_31_424971 380 12 at at IN 10_1101-2020_12_31_424971 380 13 single single JJ 10_1101-2020_12_31_424971 380 14 - - HYPH 10_1101-2020_12_31_424971 380 15 molecule molecule NN 10_1101-2020_12_31_424971 380 16 resolution resolution NN 10_1101-2020_12_31_424971 380 17 using use VBG 10_1101-2020_12_31_424971 380 18 a a DT 10_1101-2020_12_31_424971 380 19 858 858 CD 10_1101-2020_12_31_424971 380 20 laboratory laboratory NN 10_1101-2020_12_31_424971 380 21 - - HYPH 10_1101-2020_12_31_424971 380 22 built build VBN 10_1101-2020_12_31_424971 380 23 , , , 10_1101-2020_12_31_424971 380 24 wide wide JJ 10_1101-2020_12_31_424971 380 25 - - HYPH 10_1101-2020_12_31_424971 380 26 field field NN 10_1101-2020_12_31_424971 380 27 , , , 10_1101-2020_12_31_424971 380 28 prism prism NN 10_1101-2020_12_31_424971 380 29 - - HYPH 10_1101-2020_12_31_424971 380 30 based base VBN 10_1101-2020_12_31_424971 380 31 total total JJ 10_1101-2020_12_31_424971 380 32 internal internal JJ 10_1101-2020_12_31_424971 380 33 reflection reflection NN 10_1101-2020_12_31_424971 380 34 fluorescence fluorescence NN 10_1101-2020_12_31_424971 380 35 ( ( -LRB- 10_1101-2020_12_31_424971 380 36 TIRF TIRF NNP 10_1101-2020_12_31_424971 380 37 ) ) -RRB- 10_1101-2020_12_31_424971 380 38 microscope microscope NN 10_1101-2020_12_31_424971 380 39 859 859 CD 10_1101-2020_12_31_424971 380 40 with with IN 10_1101-2020_12_31_424971 380 41 a a DT 10_1101-2020_12_31_424971 380 42 532-nm 532-nm CD 10_1101-2020_12_31_424971 380 43 , , , 10_1101-2020_12_31_424971 380 44 diode diode NN 10_1101-2020_12_31_424971 380 45 - - HYPH 10_1101-2020_12_31_424971 380 46 pumped pump VBN 10_1101-2020_12_31_424971 380 47 , , , 10_1101-2020_12_31_424971 380 48 solid solid JJ 10_1101-2020_12_31_424971 380 49 - - HYPH 10_1101-2020_12_31_424971 380 50 state state NN 10_1101-2020_12_31_424971 380 51 laser laser NN 10_1101-2020_12_31_424971 380 52 ( ( -LRB- 10_1101-2020_12_31_424971 380 53 Laser Laser NNP 10_1101-2020_12_31_424971 380 54 Quantum Quantum NNP 10_1101-2020_12_31_424971 380 55 ) ) -RRB- 10_1101-2020_12_31_424971 380 56 excitation excitation NN 10_1101-2020_12_31_424971 380 57 source source NN 10_1101-2020_12_31_424971 380 58 delivering deliver VBG 10_1101-2020_12_31_424971 380 59 a a DT 10_1101-2020_12_31_424971 380 60 860 860 CD 10_1101-2020_12_31_424971 380 61 power power NN 10_1101-2020_12_31_424971 380 62 of of IN 10_1101-2020_12_31_424971 380 63 16 16 CD 10_1101-2020_12_31_424971 380 64 - - SYM 10_1101-2020_12_31_424971 380 65 25 25 CD 10_1101-2020_12_31_424971 380 66 mW mW NNP 10_1101-2020_12_31_424971 380 67 as as IN 10_1101-2020_12_31_424971 380 68 measured measure VBN 10_1101-2020_12_31_424971 380 69 at at IN 10_1101-2020_12_31_424971 380 70 the the DT 10_1101-2020_12_31_424971 380 71 prism prism NN 10_1101-2020_12_31_424971 380 72 to to TO 10_1101-2020_12_31_424971 380 73 ensure ensure VB 10_1101-2020_12_31_424971 380 74 the the DT 10_1101-2020_12_31_424971 380 75 same same JJ 10_1101-2020_12_31_424971 380 76 power power NN 10_1101-2020_12_31_424971 380 77 density density NN 10_1101-2020_12_31_424971 380 78 on on IN 10_1101-2020_12_31_424971 380 79 the the DT 10_1101-2020_12_31_424971 380 80 imaging imaging JJ 10_1101-2020_12_31_424971 380 81 861 861 CD 10_1101-2020_12_31_424971 380 82 plane plane NN 10_1101-2020_12_31_424971 380 83 . . . 10_1101-2020_12_31_424971 381 1 The the DT 10_1101-2020_12_31_424971 381 2 Cy3 cy3 NN 10_1101-2020_12_31_424971 381 3 and and CC 10_1101-2020_12_31_424971 381 4 Cy5 Cy5 NNP 10_1101-2020_12_31_424971 381 5 fluorescence fluorescence NN 10_1101-2020_12_31_424971 381 6 emissions emission NNS 10_1101-2020_12_31_424971 381 7 were be VBD 10_1101-2020_12_31_424971 381 8 simultaneously simultaneously RB 10_1101-2020_12_31_424971 381 9 collected collect VBN 10_1101-2020_12_31_424971 381 10 by by IN 10_1101-2020_12_31_424971 381 11 a a DT 10_1101-2020_12_31_424971 381 12 1.2 1.2 CD 10_1101-2020_12_31_424971 381 13 862 862 CD 10_1101-2020_12_31_424971 381 14 numerical numerical JJ 10_1101-2020_12_31_424971 381 15 aperture aperture NN 10_1101-2020_12_31_424971 381 16 , , , 10_1101-2020_12_31_424971 381 17 60 60 CD 10_1101-2020_12_31_424971 381 18 ´ ´ CD 10_1101-2020_12_31_424971 381 19 , , , 10_1101-2020_12_31_424971 381 20 water water NN 10_1101-2020_12_31_424971 381 21 - - HYPH 10_1101-2020_12_31_424971 381 22 immersion immersion NN 10_1101-2020_12_31_424971 381 23 objective objective NN 10_1101-2020_12_31_424971 381 24 ( ( -LRB- 10_1101-2020_12_31_424971 381 25 Nikon Nikon NNP 10_1101-2020_12_31_424971 381 26 ) ) -RRB- 10_1101-2020_12_31_424971 381 27 and and CC 10_1101-2020_12_31_424971 381 28 separated separate VBD 10_1101-2020_12_31_424971 381 29 based base VBN 10_1101-2020_12_31_424971 381 30 on on IN 10_1101-2020_12_31_424971 381 31 wavelength wavelength NN 10_1101-2020_12_31_424971 381 32 863 863 CD 10_1101-2020_12_31_424971 381 33 using use VBG 10_1101-2020_12_31_424971 381 34 a a DT 10_1101-2020_12_31_424971 381 35 two two CD 10_1101-2020_12_31_424971 381 36 - - HYPH 10_1101-2020_12_31_424971 381 37 channel channel NN 10_1101-2020_12_31_424971 381 38 , , , 10_1101-2020_12_31_424971 381 39 simultaneous simultaneous JJ 10_1101-2020_12_31_424971 381 40 - - HYPH 10_1101-2020_12_31_424971 381 41 imaging image VBG 10_1101-2020_12_31_424971 381 42 system system NN 10_1101-2020_12_31_424971 381 43 ( ( -LRB- 10_1101-2020_12_31_424971 381 44 Dual Dual NNP 10_1101-2020_12_31_424971 381 45 ViewTM ViewTM NNP 10_1101-2020_12_31_424971 381 46 , , , 10_1101-2020_12_31_424971 381 47 Optical Optical NNP 10_1101-2020_12_31_424971 381 48 Insights Insights NNPS 10_1101-2020_12_31_424971 381 49 LLC LLC NNP 10_1101-2020_12_31_424971 381 50 ) ) -RRB- 10_1101-2020_12_31_424971 381 51 . . . 10_1101-2020_12_31_424971 382 1 The the DT 10_1101-2020_12_31_424971 382 2 Cy3 cy3 CD 10_1101-2020_12_31_424971 382 3 864 864 CD 10_1101-2020_12_31_424971 382 4 and and CC 10_1101-2020_12_31_424971 382 5 Cy5 Cy5 NNP 10_1101-2020_12_31_424971 382 6 fluorescence fluorescence NN 10_1101-2020_12_31_424971 382 7 intensities intensity NNS 10_1101-2020_12_31_424971 382 8 were be VBD 10_1101-2020_12_31_424971 382 9 recorded record VBN 10_1101-2020_12_31_424971 382 10 using use VBG 10_1101-2020_12_31_424971 382 11 a a DT 10_1101-2020_12_31_424971 382 12 1024 1024 CD 10_1101-2020_12_31_424971 382 13 ´ ´ CD 10_1101-2020_12_31_424971 382 14 1024 1024 CD 10_1101-2020_12_31_424971 382 15 pixel pixel NN 10_1101-2020_12_31_424971 382 16 , , , 10_1101-2020_12_31_424971 382 17 back back RB 10_1101-2020_12_31_424971 382 18 - - HYPH 10_1101-2020_12_31_424971 382 19 illuminated illuminate VBN 10_1101-2020_12_31_424971 382 20 865 865 CD 10_1101-2020_12_31_424971 382 21 electron electron NN 10_1101-2020_12_31_424971 382 22 - - HYPH 10_1101-2020_12_31_424971 382 23 multiplying multiply VBG 10_1101-2020_12_31_424971 382 24 charge charge NN 10_1101-2020_12_31_424971 382 25 - - HYPH 10_1101-2020_12_31_424971 382 26 coupled couple VBN 10_1101-2020_12_31_424971 382 27 - - HYPH 10_1101-2020_12_31_424971 382 28 device device NN 10_1101-2020_12_31_424971 382 29 ( ( -LRB- 10_1101-2020_12_31_424971 382 30 EMCCD EMCCD NNP 10_1101-2020_12_31_424971 382 31 ) ) -RRB- 10_1101-2020_12_31_424971 382 32 camera camera NN 10_1101-2020_12_31_424971 382 33 ( ( -LRB- 10_1101-2020_12_31_424971 382 34 Andor Andor NNP 10_1101-2020_12_31_424971 382 35 iXon iXon NNP 10_1101-2020_12_31_424971 382 36 Ultra Ultra NNP 10_1101-2020_12_31_424971 382 37 888 888 CD 10_1101-2020_12_31_424971 382 38 ) ) -RRB- 10_1101-2020_12_31_424971 382 39 operating operate VBG 10_1101-2020_12_31_424971 382 40 866 866 CD 10_1101-2020_12_31_424971 382 41 with with IN 10_1101-2020_12_31_424971 382 42 2 2 CD 10_1101-2020_12_31_424971 382 43 ´ ´ CD 10_1101-2020_12_31_424971 382 44 2 2 CD 10_1101-2020_12_31_424971 382 45 pixel pixel NN 10_1101-2020_12_31_424971 382 46 binning bin VBG 10_1101-2020_12_31_424971 382 47 at at IN 10_1101-2020_12_31_424971 382 48 an an DT 10_1101-2020_12_31_424971 382 49 acquisition acquisition NN 10_1101-2020_12_31_424971 382 50 time time NN 10_1101-2020_12_31_424971 382 51 of of IN 10_1101-2020_12_31_424971 382 52 0.1 0.1 CD 10_1101-2020_12_31_424971 382 53 seconds second NNS 10_1101-2020_12_31_424971 382 54 per per IN 10_1101-2020_12_31_424971 382 55 frame frame NN 10_1101-2020_12_31_424971 382 56 controlled control VBN 10_1101-2020_12_31_424971 382 57 by by IN 10_1101-2020_12_31_424971 382 58 software software NN 10_1101-2020_12_31_424971 382 59 867 867 CD 10_1101-2020_12_31_424971 382 60 μManager μManager NNP 10_1101-2020_12_31_424971 382 61 1.4 1.4 CD 10_1101-2020_12_31_424971 382 62 . . . 10_1101-2020_12_31_424971 383 1 This this DT 10_1101-2020_12_31_424971 383 2 microscope microscope NN 10_1101-2020_12_31_424971 383 3 allows allow VBZ 10_1101-2020_12_31_424971 383 4 direct direct JJ 10_1101-2020_12_31_424971 383 5 visualization visualization NN 10_1101-2020_12_31_424971 383 6 of of IN 10_1101-2020_12_31_424971 383 7 thousands thousand NNS 10_1101-2020_12_31_424971 383 8 of of IN 10_1101-2020_12_31_424971 383 9 individual individual JJ 10_1101-2020_12_31_424971 383 10 70S 70S NNP 10_1101-2020_12_31_424971 383 11 ICs ic NNS 10_1101-2020_12_31_424971 383 12 or or CC 10_1101-2020_12_31_424971 383 13 868 868 CD 10_1101-2020_12_31_424971 383 14 PRE PRE NNP 10_1101-2020_12_31_424971 383 15 - - HYPH 10_1101-2020_12_31_424971 383 16 A a DT 10_1101-2020_12_31_424971 383 17 complexes complex NNS 10_1101-2020_12_31_424971 383 18 in in IN 10_1101-2020_12_31_424971 383 19 a a DT 10_1101-2020_12_31_424971 383 20 field field NN 10_1101-2020_12_31_424971 383 21 - - HYPH 10_1101-2020_12_31_424971 383 22 of of IN 10_1101-2020_12_31_424971 383 23 - - HYPH 10_1101-2020_12_31_424971 383 24 view view NN 10_1101-2020_12_31_424971 383 25 of of IN 10_1101-2020_12_31_424971 383 26 115 115 CD 10_1101-2020_12_31_424971 383 27 × × CD 10_1101-2020_12_31_424971 383 28 230 230 CD 10_1101-2020_12_31_424971 383 29 µm2 µm2 RB 10_1101-2020_12_31_424971 383 30 . . . 10_1101-2020_12_31_424971 384 1 Each each DT 10_1101-2020_12_31_424971 384 2 movie movie NN 10_1101-2020_12_31_424971 384 3 was be VBD 10_1101-2020_12_31_424971 384 4 composed compose VBN 10_1101-2020_12_31_424971 384 5 of of IN 10_1101-2020_12_31_424971 384 6 600 600 CD 10_1101-2020_12_31_424971 384 7 frames frame NNS 10_1101-2020_12_31_424971 384 8 869 869 CD 10_1101-2020_12_31_424971 384 9 in in IN 10_1101-2020_12_31_424971 384 10 order order NN 10_1101-2020_12_31_424971 384 11 to to TO 10_1101-2020_12_31_424971 384 12 ensure ensure VB 10_1101-2020_12_31_424971 384 13 that that IN 10_1101-2020_12_31_424971 384 14 the the DT 10_1101-2020_12_31_424971 384 15 majority majority NN 10_1101-2020_12_31_424971 384 16 of of IN 10_1101-2020_12_31_424971 384 17 the the DT 10_1101-2020_12_31_424971 384 18 fluorophores fluorophore NNS 10_1101-2020_12_31_424971 384 19 in in IN 10_1101-2020_12_31_424971 384 20 the the DT 10_1101-2020_12_31_424971 384 21 field field NN 10_1101-2020_12_31_424971 384 22 - - HYPH 10_1101-2020_12_31_424971 384 23 of of IN 10_1101-2020_12_31_424971 384 24 - - HYPH 10_1101-2020_12_31_424971 384 25 view view NN 10_1101-2020_12_31_424971 384 26 were be VBD 10_1101-2020_12_31_424971 384 27 photobleached photobleache VBN 10_1101-2020_12_31_424971 384 28 870 870 CD 10_1101-2020_12_31_424971 384 29 within within IN 10_1101-2020_12_31_424971 384 30 the the DT 10_1101-2020_12_31_424971 384 31 observation observation NN 10_1101-2020_12_31_424971 384 32 period period NN 10_1101-2020_12_31_424971 384 33 . . . 10_1101-2020_12_31_424971 385 1 For for IN 10_1101-2020_12_31_424971 385 2 stopped stop VBN 10_1101-2020_12_31_424971 385 3 - - HYPH 10_1101-2020_12_31_424971 385 4 flow flow NN 10_1101-2020_12_31_424971 385 5 experiments experiment NNS 10_1101-2020_12_31_424971 385 6 using use VBG 10_1101-2020_12_31_424971 385 7 tethered tethered JJ 10_1101-2020_12_31_424971 385 8 70S 70S NNP 10_1101-2020_12_31_424971 385 9 ICs ic NNS 10_1101-2020_12_31_424971 385 10 , , , 10_1101-2020_12_31_424971 385 11 we -PRON- PRP 10_1101-2020_12_31_424971 385 12 delivered deliver VBD 10_1101-2020_12_31_424971 385 13 871 871 CD 10_1101-2020_12_31_424971 385 14 0.25 0.25 CD 10_1101-2020_12_31_424971 385 15 µM µM NNP 10_1101-2020_12_31_424971 385 16 of of IN 10_1101-2020_12_31_424971 385 17 G37-state G37-state NNP 10_1101-2020_12_31_424971 385 18 SufB2- sufb2- NN 10_1101-2020_12_31_424971 385 19 or or CC 10_1101-2020_12_31_424971 385 20 ProL ProL NNP 10_1101-2020_12_31_424971 385 21 - - HYPH 10_1101-2020_12_31_424971 385 22 TC TC NNP 10_1101-2020_12_31_424971 385 23 in in IN 10_1101-2020_12_31_424971 385 24 the the DT 10_1101-2020_12_31_424971 385 25 absence absence NN 10_1101-2020_12_31_424971 385 26 of of IN 10_1101-2020_12_31_424971 385 27 EF EF NNP 10_1101-2020_12_31_424971 385 28 - - HYPH 10_1101-2020_12_31_424971 385 29 G G NNP 10_1101-2020_12_31_424971 385 30 or or CC 10_1101-2020_12_31_424971 385 31 , , , 10_1101-2020_12_31_424971 385 32 when when WRB 10_1101-2020_12_31_424971 385 33 specified specify VBN 10_1101-2020_12_31_424971 385 34 , , , 10_1101-2020_12_31_424971 385 35 in in IN 10_1101-2020_12_31_424971 385 36 the the DT 10_1101-2020_12_31_424971 385 37 872 872 CD 10_1101-2020_12_31_424971 385 38 presence presence NN 10_1101-2020_12_31_424971 385 39 of of IN 10_1101-2020_12_31_424971 385 40 a a DT 10_1101-2020_12_31_424971 385 41 2 2 CD 10_1101-2020_12_31_424971 385 42 µM µM NNP 10_1101-2020_12_31_424971 385 43 saturating saturating NN 10_1101-2020_12_31_424971 385 44 concentration concentration NN 10_1101-2020_12_31_424971 385 45 of of IN 10_1101-2020_12_31_424971 385 46 EF EF NNP 10_1101-2020_12_31_424971 385 47 - - HYPH 10_1101-2020_12_31_424971 385 48 G. G. NNP 10_1101-2020_12_31_424971 385 49 Stopped Stopped NNP 10_1101-2020_12_31_424971 385 50 - - HYPH 10_1101-2020_12_31_424971 385 51 flow flow NN 10_1101-2020_12_31_424971 385 52 experiments experiment NNS 10_1101-2020_12_31_424971 385 53 proceeded proceed VBN 10_1101-2020_12_31_424971 385 54 by by IN 10_1101-2020_12_31_424971 385 55 873 873 CD 10_1101-2020_12_31_424971 385 56 recording record VBG 10_1101-2020_12_31_424971 385 57 an an DT 10_1101-2020_12_31_424971 385 58 initial initial JJ 10_1101-2020_12_31_424971 385 59 pre pre JJ 10_1101-2020_12_31_424971 385 60 - - JJ 10_1101-2020_12_31_424971 385 61 steady steady JJ 10_1101-2020_12_31_424971 385 62 - - HYPH 10_1101-2020_12_31_424971 385 63 state state NN 10_1101-2020_12_31_424971 385 64 movie movie NN 10_1101-2020_12_31_424971 385 65 of of IN 10_1101-2020_12_31_424971 385 66 a a DT 10_1101-2020_12_31_424971 385 67 field field NN 10_1101-2020_12_31_424971 385 68 - - HYPH 10_1101-2020_12_31_424971 385 69 of of IN 10_1101-2020_12_31_424971 385 70 - - HYPH 10_1101-2020_12_31_424971 385 71 view view NN 10_1101-2020_12_31_424971 385 72 that that IN 10_1101-2020_12_31_424971 385 73 captured capture VBD 10_1101-2020_12_31_424971 385 74 conformational conformational JJ 10_1101-2020_12_31_424971 385 75 changes change NNS 10_1101-2020_12_31_424971 385 76 874 874 CD 10_1101-2020_12_31_424971 385 77 taking take VBG 10_1101-2020_12_31_424971 385 78 place place NN 10_1101-2020_12_31_424971 385 79 during during IN 10_1101-2020_12_31_424971 385 80 delivery delivery NN 10_1101-2020_12_31_424971 385 81 followed follow VBN 10_1101-2020_12_31_424971 385 82 by by IN 10_1101-2020_12_31_424971 385 83 recording record VBG 10_1101-2020_12_31_424971 385 84 of of IN 10_1101-2020_12_31_424971 385 85 one one CD 10_1101-2020_12_31_424971 385 86 or or CC 10_1101-2020_12_31_424971 385 87 more more RBR 10_1101-2020_12_31_424971 385 88 steady steady JJ 10_1101-2020_12_31_424971 385 89 - - HYPH 10_1101-2020_12_31_424971 385 90 state state NN 10_1101-2020_12_31_424971 385 91 movies movie NNS 10_1101-2020_12_31_424971 385 92 of of IN 10_1101-2020_12_31_424971 385 93 different different JJ 10_1101-2020_12_31_424971 385 94 875 875 CD 10_1101-2020_12_31_424971 385 95 fields field NNS 10_1101-2020_12_31_424971 385 96 - - HYPH 10_1101-2020_12_31_424971 385 97 of of IN 10_1101-2020_12_31_424971 385 98 - - HYPH 10_1101-2020_12_31_424971 385 99 view view NN 10_1101-2020_12_31_424971 385 100 that that IN 10_1101-2020_12_31_424971 385 101 captured capture VBD 10_1101-2020_12_31_424971 385 102 conformational conformational JJ 10_1101-2020_12_31_424971 385 103 changes change NNS 10_1101-2020_12_31_424971 385 104 taking take VBG 10_1101-2020_12_31_424971 385 105 place place NN 10_1101-2020_12_31_424971 385 106 the the DT 10_1101-2020_12_31_424971 385 107 specified specified JJ 10_1101-2020_12_31_424971 385 108 number number NN 10_1101-2020_12_31_424971 385 109 of of IN 10_1101-2020_12_31_424971 385 110 minutes minute NNS 10_1101-2020_12_31_424971 385 111 876 876 CD 10_1101-2020_12_31_424971 385 112 post post NN 10_1101-2020_12_31_424971 385 113 - - NN 10_1101-2020_12_31_424971 385 114 delivery delivery NN 10_1101-2020_12_31_424971 385 115 . . . 10_1101-2020_12_31_424971 386 1 877 877 CD 10_1101-2020_12_31_424971 386 2 878 878 CD 10_1101-2020_12_31_424971 386 3 Analysis analysis NN 10_1101-2020_12_31_424971 386 4 of of IN 10_1101-2020_12_31_424971 386 5 smFRET smFRET NNP 10_1101-2020_12_31_424971 386 6 experiments experiment NNS 10_1101-2020_12_31_424971 386 7 . . . 10_1101-2020_12_31_424971 387 1 For for IN 10_1101-2020_12_31_424971 387 2 each each DT 10_1101-2020_12_31_424971 387 3 TIRF TIRF NNP 10_1101-2020_12_31_424971 387 4 microscopy microscopy JJ 10_1101-2020_12_31_424971 387 5 movie movie NN 10_1101-2020_12_31_424971 387 6 , , , 10_1101-2020_12_31_424971 387 7 we -PRON- PRP 10_1101-2020_12_31_424971 387 8 identified identify VBD 10_1101-2020_12_31_424971 387 9 879 879 CD 10_1101-2020_12_31_424971 387 10 fluorophores fluorophore NNS 10_1101-2020_12_31_424971 387 11 , , , 10_1101-2020_12_31_424971 387 12 aligned align VBN 10_1101-2020_12_31_424971 387 13 Cy3 cy3 NN 10_1101-2020_12_31_424971 387 14 and and CC 10_1101-2020_12_31_424971 387 15 Cy5 cy5 NN 10_1101-2020_12_31_424971 387 16 imaging imaging NN 10_1101-2020_12_31_424971 387 17 channels channel NNS 10_1101-2020_12_31_424971 387 18 , , , 10_1101-2020_12_31_424971 387 19 and and CC 10_1101-2020_12_31_424971 387 20 generated generate VBN 10_1101-2020_12_31_424971 387 21 fluorescence fluorescence NN 10_1101-2020_12_31_424971 387 22 intensity intensity NN 10_1101-2020_12_31_424971 387 23 vs. vs. IN 10_1101-2020_12_31_424971 387 24 880 880 CD 10_1101-2020_12_31_424971 387 25 .CC .CC NFP 10_1101-2020_12_31_424971 387 26 - - HYPH 10_1101-2020_12_31_424971 387 27 BY by IN 10_1101-2020_12_31_424971 387 28 - - HYPH 10_1101-2020_12_31_424971 387 29 NC NC NNP 10_1101-2020_12_31_424971 387 30 - - HYPH 10_1101-2020_12_31_424971 387 31 ND ND NNP 10_1101-2020_12_31_424971 387 32 4.0 4.0 CD 10_1101-2020_12_31_424971 387 33 International International NNP 10_1101-2020_12_31_424971 387 34 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 387 35 under under IN 10_1101-2020_12_31_424971 387 36 a a DT 10_1101-2020_12_31_424971 387 37 ( ( -LRB- 10_1101-2020_12_31_424971 387 38 which which WDT 10_1101-2020_12_31_424971 387 39 was be VBD 10_1101-2020_12_31_424971 387 40 not not RB 10_1101-2020_12_31_424971 387 41 certified certify VBN 10_1101-2020_12_31_424971 387 42 by by IN 10_1101-2020_12_31_424971 387 43 peer peer NN 10_1101-2020_12_31_424971 387 44 review review NN 10_1101-2020_12_31_424971 387 45 ) ) -RRB- 10_1101-2020_12_31_424971 387 46 is be VBZ 10_1101-2020_12_31_424971 387 47 the the DT 10_1101-2020_12_31_424971 387 48 author author NN 10_1101-2020_12_31_424971 387 49 / / SYM 10_1101-2020_12_31_424971 387 50 funder funder NN 10_1101-2020_12_31_424971 387 51 , , , 10_1101-2020_12_31_424971 387 52 who who WP 10_1101-2020_12_31_424971 387 53 has have VBZ 10_1101-2020_12_31_424971 387 54 granted grant VBN 10_1101-2020_12_31_424971 387 55 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 387 56 a a DT 10_1101-2020_12_31_424971 387 57 license license NN 10_1101-2020_12_31_424971 387 58 to to TO 10_1101-2020_12_31_424971 387 59 display display VB 10_1101-2020_12_31_424971 387 60 the the DT 10_1101-2020_12_31_424971 387 61 preprint preprint NN 10_1101-2020_12_31_424971 387 62 in in IN 10_1101-2020_12_31_424971 387 63 perpetuity perpetuity NN 10_1101-2020_12_31_424971 387 64 . . . 10_1101-2020_12_31_424971 388 1 It -PRON- PRP 10_1101-2020_12_31_424971 388 2 is be VBZ 10_1101-2020_12_31_424971 388 3 made make VBN 10_1101-2020_12_31_424971 388 4 The the DT 10_1101-2020_12_31_424971 388 5 copyright copyright NN 10_1101-2020_12_31_424971 388 6 holder holder NN 10_1101-2020_12_31_424971 388 7 for for IN 10_1101-2020_12_31_424971 388 8 this this DT 10_1101-2020_12_31_424971 388 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 388 10 version version NN 10_1101-2020_12_31_424971 388 11 posted post VBD 10_1101-2020_12_31_424971 388 12 January January NNP 10_1101-2020_12_31_424971 388 13 2 2 CD 10_1101-2020_12_31_424971 388 14 , , , 10_1101-2020_12_31_424971 388 15 2021 2021 CD 10_1101-2020_12_31_424971 388 16 . . . 10_1101-2020_12_31_424971 388 17 ; ; : 10_1101-2020_12_31_424971 388 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 388 19 : : : 10_1101-2020_12_31_424971 388 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 388 21 preprint preprint NN 10_1101-2020_12_31_424971 388 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 388 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 388 24 36 36 CD 10_1101-2020_12_31_424971 388 25 time time NN 10_1101-2020_12_31_424971 388 26 trajectories trajectory NNS 10_1101-2020_12_31_424971 388 27 for for IN 10_1101-2020_12_31_424971 388 28 each each DT 10_1101-2020_12_31_424971 388 29 pair pair NN 10_1101-2020_12_31_424971 388 30 of of IN 10_1101-2020_12_31_424971 388 31 Cy3 cy3 NN 10_1101-2020_12_31_424971 388 32 and and CC 10_1101-2020_12_31_424971 388 33 Cy5 cy5 NN 10_1101-2020_12_31_424971 388 34 fluorophores fluorophore NNS 10_1101-2020_12_31_424971 388 35 using use VBG 10_1101-2020_12_31_424971 388 36 custom custom RB 10_1101-2020_12_31_424971 388 37 - - HYPH 10_1101-2020_12_31_424971 388 38 written write VBN 10_1101-2020_12_31_424971 388 39 software software NN 10_1101-2020_12_31_424971 388 40 881 881 CD 10_1101-2020_12_31_424971 388 41 ( ( -LRB- 10_1101-2020_12_31_424971 388 42 manuscript manuscript NN 10_1101-2020_12_31_424971 388 43 in in IN 10_1101-2020_12_31_424971 388 44 preparation preparation NN 10_1101-2020_12_31_424971 388 45 ; ; : 10_1101-2020_12_31_424971 388 46 Jason Jason NNP 10_1101-2020_12_31_424971 388 47 Hon Hon NNP 10_1101-2020_12_31_424971 388 48 , , , 10_1101-2020_12_31_424971 388 49 Colin Colin NNP 10_1101-2020_12_31_424971 388 50 Kinz Kinz NNP 10_1101-2020_12_31_424971 388 51 - - HYPH 10_1101-2020_12_31_424971 388 52 Thompson Thompson NNP 10_1101-2020_12_31_424971 388 53 , , , 10_1101-2020_12_31_424971 388 54 Ruben Ruben NNP 10_1101-2020_12_31_424971 388 55 L. L. NNP 10_1101-2020_12_31_424971 388 56 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 388 57 ) ) -RRB- 10_1101-2020_12_31_424971 388 58 as as IN 10_1101-2020_12_31_424971 388 59 described describe VBD 10_1101-2020_12_31_424971 388 60 882 882 CD 10_1101-2020_12_31_424971 388 61 previously66 previously66 NN 10_1101-2020_12_31_424971 388 62 . . . 10_1101-2020_12_31_424971 389 1 For for IN 10_1101-2020_12_31_424971 389 2 each each DT 10_1101-2020_12_31_424971 389 3 time time NN 10_1101-2020_12_31_424971 389 4 point point NN 10_1101-2020_12_31_424971 389 5 , , , 10_1101-2020_12_31_424971 389 6 Cy5 cy5 NN 10_1101-2020_12_31_424971 389 7 fluorescence fluorescence NN 10_1101-2020_12_31_424971 389 8 intensity intensity NN 10_1101-2020_12_31_424971 389 9 values value NNS 10_1101-2020_12_31_424971 389 10 were be VBD 10_1101-2020_12_31_424971 389 11 corrected correct VBN 10_1101-2020_12_31_424971 389 12 for for IN 10_1101-2020_12_31_424971 389 13 Cy3 cy3 CD 10_1101-2020_12_31_424971 389 14 883 883 CD 10_1101-2020_12_31_424971 389 15 bleedthrough bleedthrough NN 10_1101-2020_12_31_424971 389 16 by by IN 10_1101-2020_12_31_424971 389 17 subtracting subtract VBG 10_1101-2020_12_31_424971 389 18 5 5 CD 10_1101-2020_12_31_424971 389 19 % % NN 10_1101-2020_12_31_424971 389 20 of of IN 10_1101-2020_12_31_424971 389 21 the the DT 10_1101-2020_12_31_424971 389 22 Cy3 Cy3 NNP 10_1101-2020_12_31_424971 389 23 fluorescence fluorescence NN 10_1101-2020_12_31_424971 389 24 intensity intensity NN 10_1101-2020_12_31_424971 389 25 value value NN 10_1101-2020_12_31_424971 389 26 in in IN 10_1101-2020_12_31_424971 389 27 the the DT 10_1101-2020_12_31_424971 389 28 corresponding corresponding JJ 10_1101-2020_12_31_424971 389 29 Cy3 cy3 CD 10_1101-2020_12_31_424971 389 30 884 884 CD 10_1101-2020_12_31_424971 389 31 fluorescence fluorescence NN 10_1101-2020_12_31_424971 389 32 intensity intensity NN 10_1101-2020_12_31_424971 389 33 vs. vs. IN 10_1101-2020_12_31_424971 389 34 time time NN 10_1101-2020_12_31_424971 389 35 trajectory trajectory CD 10_1101-2020_12_31_424971 389 36 . . . 10_1101-2020_12_31_424971 390 1 EFRET EFRET NNP 10_1101-2020_12_31_424971 390 2 vs. vs. IN 10_1101-2020_12_31_424971 390 3 time time NN 10_1101-2020_12_31_424971 390 4 trajectories trajectory NNS 10_1101-2020_12_31_424971 390 5 were be VBD 10_1101-2020_12_31_424971 390 6 generated generate VBN 10_1101-2020_12_31_424971 390 7 by by IN 10_1101-2020_12_31_424971 390 8 using use VBG 10_1101-2020_12_31_424971 390 9 the the DT 10_1101-2020_12_31_424971 390 10 885 885 CD 10_1101-2020_12_31_424971 390 11 Cy3 cy3 CD 10_1101-2020_12_31_424971 390 12 fluorescence fluorescence NN 10_1101-2020_12_31_424971 390 13 intensity intensity NN 10_1101-2020_12_31_424971 390 14 ( ( -LRB- 10_1101-2020_12_31_424971 390 15 ICy3 ICy3 NNP 10_1101-2020_12_31_424971 390 16 ) ) -RRB- 10_1101-2020_12_31_424971 390 17 and and CC 10_1101-2020_12_31_424971 390 18 the the DT 10_1101-2020_12_31_424971 390 19 bleedthrough bleedthrough RB 10_1101-2020_12_31_424971 390 20 - - HYPH 10_1101-2020_12_31_424971 390 21 corrected correct VBN 10_1101-2020_12_31_424971 390 22 Cy5 cy5 NN 10_1101-2020_12_31_424971 390 23 fluorescence fluorescence NN 10_1101-2020_12_31_424971 390 24 intensity intensity NN 10_1101-2020_12_31_424971 390 25 ( ( -LRB- 10_1101-2020_12_31_424971 390 26 ICy5 ICy5 NNP 10_1101-2020_12_31_424971 390 27 ) ) -RRB- 10_1101-2020_12_31_424971 390 28 886 886 CD 10_1101-2020_12_31_424971 390 29 from from IN 10_1101-2020_12_31_424971 390 30 each each DT 10_1101-2020_12_31_424971 390 31 aligned align VBN 10_1101-2020_12_31_424971 390 32 pair pair NN 10_1101-2020_12_31_424971 390 33 of of IN 10_1101-2020_12_31_424971 390 34 Cy3 cy3 NN 10_1101-2020_12_31_424971 390 35 and and CC 10_1101-2020_12_31_424971 390 36 Cy5 cy5 NN 10_1101-2020_12_31_424971 390 37 fluorophores fluorophore NNS 10_1101-2020_12_31_424971 390 38 to to TO 10_1101-2020_12_31_424971 390 39 calculate calculate VB 10_1101-2020_12_31_424971 390 40 the the DT 10_1101-2020_12_31_424971 390 41 EFRET EFRET NNP 10_1101-2020_12_31_424971 390 42 value value NN 10_1101-2020_12_31_424971 390 43 at at IN 10_1101-2020_12_31_424971 390 44 each each DT 10_1101-2020_12_31_424971 390 45 time time NN 10_1101-2020_12_31_424971 390 46 point point NN 10_1101-2020_12_31_424971 390 47 887 887 CD 10_1101-2020_12_31_424971 390 48 using use VBG 10_1101-2020_12_31_424971 390 49 EFRET EFRET NNP 10_1101-2020_12_31_424971 390 50 = = NFP 10_1101-2020_12_31_424971 390 51 ( ( -LRB- 10_1101-2020_12_31_424971 390 52 ICy5 ICy5 NNP 10_1101-2020_12_31_424971 390 53 / / , 10_1101-2020_12_31_424971 390 54 ( ( -LRB- 10_1101-2020_12_31_424971 390 55 ICy5 ICy5 NNP 10_1101-2020_12_31_424971 390 56 + + SYM 10_1101-2020_12_31_424971 390 57 ICy3 ICy3 NNP 10_1101-2020_12_31_424971 390 58 ) ) -RRB- 10_1101-2020_12_31_424971 390 59 ) ) -RRB- 10_1101-2020_12_31_424971 390 60 . . . 10_1101-2020_12_31_424971 391 1 888 888 CD 10_1101-2020_12_31_424971 391 2 For for IN 10_1101-2020_12_31_424971 391 3 both both DT 10_1101-2020_12_31_424971 391 4 pre pre JJ 10_1101-2020_12_31_424971 391 5 - - JJ 10_1101-2020_12_31_424971 391 6 steady steady JJ 10_1101-2020_12_31_424971 391 7 - - HYPH 10_1101-2020_12_31_424971 391 8 state state NN 10_1101-2020_12_31_424971 391 9 and and CC 10_1101-2020_12_31_424971 391 10 steady steady JJ 10_1101-2020_12_31_424971 391 11 - - HYPH 10_1101-2020_12_31_424971 391 12 state state NN 10_1101-2020_12_31_424971 391 13 movies movie NNS 10_1101-2020_12_31_424971 391 14 ( ( -LRB- 10_1101-2020_12_31_424971 391 15 Figures figure NNS 10_1101-2020_12_31_424971 391 16 6d-6h 6d-6h CD 10_1101-2020_12_31_424971 391 17 and and CC 10_1101-2020_12_31_424971 391 18 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 391 19 889 889 CD 10_1101-2020_12_31_424971 391 20 Figures figure NNS 10_1101-2020_12_31_424971 391 21 3 3 CD 10_1101-2020_12_31_424971 391 22 , , , 10_1101-2020_12_31_424971 391 23 5 5 CD 10_1101-2020_12_31_424971 391 24 , , , 10_1101-2020_12_31_424971 391 25 and and CC 10_1101-2020_12_31_424971 391 26 6 6 CD 10_1101-2020_12_31_424971 391 27 , , , 10_1101-2020_12_31_424971 391 28 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 391 29 Tables Tables NNP 10_1101-2020_12_31_424971 391 30 4 4 CD 10_1101-2020_12_31_424971 391 31 - - SYM 10_1101-2020_12_31_424971 391 32 7 7 CD 10_1101-2020_12_31_424971 391 33 ) ) -RRB- 10_1101-2020_12_31_424971 391 34 , , , 10_1101-2020_12_31_424971 391 35 an an DT 10_1101-2020_12_31_424971 391 36 EFRET EFRET NNP 10_1101-2020_12_31_424971 391 37 vs. vs. NN 10_1101-2020_12_31_424971 391 38 time time NN 10_1101-2020_12_31_424971 391 39 trajectory trajectory CD 10_1101-2020_12_31_424971 391 40 was be VBD 10_1101-2020_12_31_424971 391 41 selected select VBN 10_1101-2020_12_31_424971 391 42 for for IN 10_1101-2020_12_31_424971 391 43 890 890 CD 10_1101-2020_12_31_424971 391 44 further further JJ 10_1101-2020_12_31_424971 391 45 analysis analysis NN 10_1101-2020_12_31_424971 391 46 if if IN 10_1101-2020_12_31_424971 391 47 all all DT 10_1101-2020_12_31_424971 391 48 of of IN 10_1101-2020_12_31_424971 391 49 the the DT 10_1101-2020_12_31_424971 391 50 transitions transition NNS 10_1101-2020_12_31_424971 391 51 in in IN 10_1101-2020_12_31_424971 391 52 the the DT 10_1101-2020_12_31_424971 391 53 fluorescence fluorescence NN 10_1101-2020_12_31_424971 391 54 intensity intensity NN 10_1101-2020_12_31_424971 391 55 vs. vs. IN 10_1101-2020_12_31_424971 391 56 time time NN 10_1101-2020_12_31_424971 391 57 trajectory trajectory CD 10_1101-2020_12_31_424971 391 58 were be VBD 10_1101-2020_12_31_424971 391 59 anti-891 anti-891 NNP 10_1101-2020_12_31_424971 391 60 correlated correlate VBN 10_1101-2020_12_31_424971 391 61 for for IN 10_1101-2020_12_31_424971 391 62 the the DT 10_1101-2020_12_31_424971 391 63 corresponding corresponding JJ 10_1101-2020_12_31_424971 391 64 , , , 10_1101-2020_12_31_424971 391 65 aligned align VBN 10_1101-2020_12_31_424971 391 66 pair pair NN 10_1101-2020_12_31_424971 391 67 of of IN 10_1101-2020_12_31_424971 391 68 Cy3 cy3 NN 10_1101-2020_12_31_424971 391 69 and and CC 10_1101-2020_12_31_424971 391 70 Cy5 cy5 NN 10_1101-2020_12_31_424971 391 71 fluorophores fluorophore NNS 10_1101-2020_12_31_424971 391 72 , , , 10_1101-2020_12_31_424971 391 73 and and CC 10_1101-2020_12_31_424971 391 74 the the DT 10_1101-2020_12_31_424971 391 75 Cy3 cy3 CD 10_1101-2020_12_31_424971 391 76 892 892 CD 10_1101-2020_12_31_424971 391 77 fluorescence fluorescence NN 10_1101-2020_12_31_424971 391 78 intensity intensity NN 10_1101-2020_12_31_424971 391 79 vs. vs. NN 10_1101-2020_12_31_424971 391 80 time time NN 10_1101-2020_12_31_424971 391 81 trajectory trajectory CD 10_1101-2020_12_31_424971 391 82 underwent undergo VBD 10_1101-2020_12_31_424971 391 83 single single JJ 10_1101-2020_12_31_424971 391 84 - - HYPH 10_1101-2020_12_31_424971 391 85 step step NN 10_1101-2020_12_31_424971 391 86 Cy3 cy3 NN 10_1101-2020_12_31_424971 391 87 photobleaching photobleaching NN 10_1101-2020_12_31_424971 391 88 , , , 10_1101-2020_12_31_424971 391 89 893 893 CD 10_1101-2020_12_31_424971 391 90 demonstrating demonstrate VBG 10_1101-2020_12_31_424971 391 91 it -PRON- PRP 10_1101-2020_12_31_424971 391 92 arose arise VBD 10_1101-2020_12_31_424971 391 93 from from IN 10_1101-2020_12_31_424971 391 94 a a DT 10_1101-2020_12_31_424971 391 95 single single JJ 10_1101-2020_12_31_424971 391 96 ribosomal ribosomal NN 10_1101-2020_12_31_424971 391 97 complex complex NN 10_1101-2020_12_31_424971 391 98 . . . 10_1101-2020_12_31_424971 392 1 In in IN 10_1101-2020_12_31_424971 392 2 the the DT 10_1101-2020_12_31_424971 392 3 case case NN 10_1101-2020_12_31_424971 392 4 of of IN 10_1101-2020_12_31_424971 392 5 pre pre JJ 10_1101-2020_12_31_424971 392 6 - - JJ 10_1101-2020_12_31_424971 392 7 steady steady JJ 10_1101-2020_12_31_424971 392 8 - - HYPH 10_1101-2020_12_31_424971 392 9 state state NN 10_1101-2020_12_31_424971 392 10 movies movie NNS 10_1101-2020_12_31_424971 392 11 894 894 CD 10_1101-2020_12_31_424971 392 12 ( ( -LRB- 10_1101-2020_12_31_424971 392 13 Figures figure NNS 10_1101-2020_12_31_424971 392 14 6d-6 6d-6 CD 10_1101-2020_12_31_424971 392 15 g g NN 10_1101-2020_12_31_424971 392 16 , , , 10_1101-2020_12_31_424971 392 17 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 392 18 Figures Figures NNPS 10_1101-2020_12_31_424971 392 19 3 3 CD 10_1101-2020_12_31_424971 392 20 and and CC 10_1101-2020_12_31_424971 392 21 5 5 CD 10_1101-2020_12_31_424971 392 22 and and CC 10_1101-2020_12_31_424971 392 23 Tables Tables NNP 10_1101-2020_12_31_424971 392 24 4 4 CD 10_1101-2020_12_31_424971 392 25 - - SYM 10_1101-2020_12_31_424971 392 26 6 6 CD 10_1101-2020_12_31_424971 392 27 ) ) -RRB- 10_1101-2020_12_31_424971 392 28 , , , 10_1101-2020_12_31_424971 392 29 EFRET EFRET NNP 10_1101-2020_12_31_424971 392 30 vs. vs. IN 10_1101-2020_12_31_424971 392 31 time time NN 10_1101-2020_12_31_424971 392 32 trajectories trajectory NNS 10_1101-2020_12_31_424971 392 33 had have VBD 10_1101-2020_12_31_424971 392 34 895 895 CD 10_1101-2020_12_31_424971 392 35 to to TO 10_1101-2020_12_31_424971 392 36 meet meet VB 10_1101-2020_12_31_424971 392 37 two two CD 10_1101-2020_12_31_424971 392 38 additional additional JJ 10_1101-2020_12_31_424971 392 39 criteria criterion NNS 10_1101-2020_12_31_424971 392 40 in in IN 10_1101-2020_12_31_424971 392 41 order order NN 10_1101-2020_12_31_424971 392 42 to to TO 10_1101-2020_12_31_424971 392 43 be be VB 10_1101-2020_12_31_424971 392 44 selected select VBN 10_1101-2020_12_31_424971 392 45 for for IN 10_1101-2020_12_31_424971 392 46 further further JJ 10_1101-2020_12_31_424971 392 47 analysis analysis NN 10_1101-2020_12_31_424971 392 48 : : : 10_1101-2020_12_31_424971 392 49 ( ( -LRB- 10_1101-2020_12_31_424971 392 50 i i NN 10_1101-2020_12_31_424971 392 51 ) ) -RRB- 10_1101-2020_12_31_424971 392 52 EFRET efret FW 10_1101-2020_12_31_424971 392 53 vs. vs. NN 10_1101-2020_12_31_424971 392 54 time time NN 10_1101-2020_12_31_424971 392 55 896 896 CD 10_1101-2020_12_31_424971 392 56 trajectories trajectory NNS 10_1101-2020_12_31_424971 392 57 had have VBD 10_1101-2020_12_31_424971 392 58 to to TO 10_1101-2020_12_31_424971 392 59 be be VB 10_1101-2020_12_31_424971 392 60 stably stably RB 10_1101-2020_12_31_424971 392 61 sampling sample VBG 10_1101-2020_12_31_424971 392 62 EFRET EFRET NNP 10_1101-2020_12_31_424971 392 63 = = NFP 10_1101-2020_12_31_424971 392 64 0.55 0.55 CD 10_1101-2020_12_31_424971 392 65 prior prior RB 10_1101-2020_12_31_424971 392 66 to to IN 10_1101-2020_12_31_424971 392 67 TC TC NNP 10_1101-2020_12_31_424971 392 68 delivery delivery NN 10_1101-2020_12_31_424971 392 69 , , , 10_1101-2020_12_31_424971 392 70 thereby thereby RB 10_1101-2020_12_31_424971 392 71 confirming confirm VBG 10_1101-2020_12_31_424971 392 72 that that IN 10_1101-2020_12_31_424971 392 73 897 897 CD 10_1101-2020_12_31_424971 392 74 the the DT 10_1101-2020_12_31_424971 392 75 corresponding correspond VBG 10_1101-2020_12_31_424971 392 76 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 392 77 complex complex NN 10_1101-2020_12_31_424971 392 78 was be VBD 10_1101-2020_12_31_424971 392 79 a a DT 10_1101-2020_12_31_424971 392 80 70S 70S NNP 10_1101-2020_12_31_424971 392 81 IC IC NNP 10_1101-2020_12_31_424971 392 82 carrying carry VBG 10_1101-2020_12_31_424971 392 83 an an DT 10_1101-2020_12_31_424971 392 84 fMet fMet NNP 10_1101-2020_12_31_424971 392 85 - - HYPH 10_1101-2020_12_31_424971 392 86 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 392 87 at at IN 10_1101-2020_12_31_424971 392 88 the the DT 10_1101-2020_12_31_424971 392 89 P p NN 10_1101-2020_12_31_424971 392 90 site site NN 10_1101-2020_12_31_424971 392 91 and and CC 10_1101-2020_12_31_424971 392 92 898 898 CD 10_1101-2020_12_31_424971 392 93 ( ( -LRB- 10_1101-2020_12_31_424971 392 94 ii ii NN 10_1101-2020_12_31_424971 392 95 ) ) -RRB- 10_1101-2020_12_31_424971 392 96 EFRET efret FW 10_1101-2020_12_31_424971 392 97 vs. vs. NN 10_1101-2020_12_31_424971 392 98 time time NN 10_1101-2020_12_31_424971 392 99 trajectories trajectory NNS 10_1101-2020_12_31_424971 392 100 had have VBD 10_1101-2020_12_31_424971 392 101 to to TO 10_1101-2020_12_31_424971 392 102 exhibit exhibit VB 10_1101-2020_12_31_424971 392 103 at at RB 10_1101-2020_12_31_424971 392 104 least least RBS 10_1101-2020_12_31_424971 392 105 one one CD 10_1101-2020_12_31_424971 392 106 0.55→0.31 0.55→0.31 CD 10_1101-2020_12_31_424971 392 107 transition transition NN 10_1101-2020_12_31_424971 392 108 after after IN 10_1101-2020_12_31_424971 392 109 delivery delivery NN 10_1101-2020_12_31_424971 392 110 of of IN 10_1101-2020_12_31_424971 392 111 TCs tc NNS 10_1101-2020_12_31_424971 392 112 , , , 10_1101-2020_12_31_424971 392 113 899 899 CD 10_1101-2020_12_31_424971 392 114 thereby thereby RB 10_1101-2020_12_31_424971 392 115 confirming confirm VBG 10_1101-2020_12_31_424971 392 116 that that IN 10_1101-2020_12_31_424971 392 117 the the DT 10_1101-2020_12_31_424971 392 118 corresponding correspond VBG 10_1101-2020_12_31_424971 392 119 70S 70S NNP 10_1101-2020_12_31_424971 392 120 IC IC NNP 10_1101-2020_12_31_424971 392 121 had have VBD 10_1101-2020_12_31_424971 392 122 accommodated accommodate VBN 10_1101-2020_12_31_424971 392 123 a a DT 10_1101-2020_12_31_424971 392 124 Pro Pro NNP 10_1101-2020_12_31_424971 392 125 - - JJ 10_1101-2020_12_31_424971 392 126 SufB2 sufb2 JJ 10_1101-2020_12_31_424971 392 127 or or CC 10_1101-2020_12_31_424971 392 128 Pro pro JJ 10_1101-2020_12_31_424971 392 129 - - JJ 10_1101-2020_12_31_424971 392 130 ProL ProL NNP 10_1101-2020_12_31_424971 392 131 900 900 CD 10_1101-2020_12_31_424971 392 132 into into IN 10_1101-2020_12_31_424971 392 133 the the DT 10_1101-2020_12_31_424971 392 134 A A NNP 10_1101-2020_12_31_424971 392 135 site site NN 10_1101-2020_12_31_424971 392 136 , , , 10_1101-2020_12_31_424971 392 137 that that IN 10_1101-2020_12_31_424971 392 138 the the DT 10_1101-2020_12_31_424971 392 139 A a DT 10_1101-2020_12_31_424971 392 140 site site NN 10_1101-2020_12_31_424971 392 141 - - HYPH 10_1101-2020_12_31_424971 392 142 bound bind VBN 10_1101-2020_12_31_424971 392 143 Pro pro JJ 10_1101-2020_12_31_424971 392 144 - - JJ 10_1101-2020_12_31_424971 392 145 SufB2 sufb2 JJ 10_1101-2020_12_31_424971 392 146 or or CC 10_1101-2020_12_31_424971 392 147 Pro pro JJ 10_1101-2020_12_31_424971 392 148 - - JJ 10_1101-2020_12_31_424971 392 149 ProL ProL NNP 10_1101-2020_12_31_424971 392 150 had have VBD 10_1101-2020_12_31_424971 392 151 participated participate VBN 10_1101-2020_12_31_424971 392 152 as as IN 10_1101-2020_12_31_424971 392 153 the the DT 10_1101-2020_12_31_424971 392 154 acceptor acceptor NN 10_1101-2020_12_31_424971 392 155 in in IN 10_1101-2020_12_31_424971 392 156 901 901 CD 10_1101-2020_12_31_424971 392 157 peptide peptide NN 10_1101-2020_12_31_424971 392 158 - - HYPH 10_1101-2020_12_31_424971 392 159 bond bond NN 10_1101-2020_12_31_424971 392 160 formation formation NN 10_1101-2020_12_31_424971 392 161 , , , 10_1101-2020_12_31_424971 392 162 and and CC 10_1101-2020_12_31_424971 392 163 that that IN 10_1101-2020_12_31_424971 392 164 the the DT 10_1101-2020_12_31_424971 392 165 resulting result VBG 10_1101-2020_12_31_424971 392 166 PRE PRE NNP 10_1101-2020_12_31_424971 392 167 complex complex NN 10_1101-2020_12_31_424971 392 168 was be VBD 10_1101-2020_12_31_424971 392 169 capable capable JJ 10_1101-2020_12_31_424971 392 170 of of IN 10_1101-2020_12_31_424971 392 171 undergoing undergo VBG 10_1101-2020_12_31_424971 392 172 902 902 CD 10_1101-2020_12_31_424971 392 173 GS1→GS2 GS1→GS2 NNP 10_1101-2020_12_31_424971 392 174 transitions transition NNS 10_1101-2020_12_31_424971 392 175 . . . 10_1101-2020_12_31_424971 393 1 We -PRON- PRP 10_1101-2020_12_31_424971 393 2 note note VBP 10_1101-2020_12_31_424971 393 3 that that IN 10_1101-2020_12_31_424971 393 4 the the DT 10_1101-2020_12_31_424971 393 5 second second JJ 10_1101-2020_12_31_424971 393 6 criterion criterion NN 10_1101-2020_12_31_424971 393 7 might may MD 10_1101-2020_12_31_424971 393 8 result result VB 10_1101-2020_12_31_424971 393 9 in in IN 10_1101-2020_12_31_424971 393 10 the the DT 10_1101-2020_12_31_424971 393 11 exclusion exclusion NN 10_1101-2020_12_31_424971 393 12 of of IN 10_1101-2020_12_31_424971 393 13 EFRET EFRET NNP 10_1101-2020_12_31_424971 393 14 vs. vs. NN 10_1101-2020_12_31_424971 393 15 903 903 CD 10_1101-2020_12_31_424971 393 16 time time NN 10_1101-2020_12_31_424971 393 17 trajectories trajectory NNS 10_1101-2020_12_31_424971 393 18 in in IN 10_1101-2020_12_31_424971 393 19 which which WDT 10_1101-2020_12_31_424971 393 20 Cy3 cy3 NN 10_1101-2020_12_31_424971 393 21 or or CC 10_1101-2020_12_31_424971 393 22 Cy5 cy5 NN 10_1101-2020_12_31_424971 393 23 simply simply RB 10_1101-2020_12_31_424971 393 24 photobleached photobleache VBN 10_1101-2020_12_31_424971 393 25 prior prior RB 10_1101-2020_12_31_424971 393 26 to to IN 10_1101-2020_12_31_424971 393 27 undergoing undergo VBG 10_1101-2020_12_31_424971 393 28 a a DT 10_1101-2020_12_31_424971 393 29 0.55→0.31 0.55→0.31 NNP 10_1101-2020_12_31_424971 393 30 904 904 CD 10_1101-2020_12_31_424971 393 31 transition transition NN 10_1101-2020_12_31_424971 393 32 , , , 10_1101-2020_12_31_424971 393 33 and and CC 10_1101-2020_12_31_424971 393 34 could could MD 10_1101-2020_12_31_424971 393 35 therefore therefore RB 10_1101-2020_12_31_424971 393 36 result result VB 10_1101-2020_12_31_424971 393 37 in in IN 10_1101-2020_12_31_424971 393 38 a a DT 10_1101-2020_12_31_424971 393 39 slight slight JJ 10_1101-2020_12_31_424971 393 40 overestimation overestimation NN 10_1101-2020_12_31_424971 393 41 of of IN 10_1101-2020_12_31_424971 393 42 k70S k70s ADD 10_1101-2020_12_31_424971 393 43 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 393 44 and/or and/or CC 10_1101-2020_12_31_424971 393 45 kGS1→GS2 kGS1→GS2 NNP 10_1101-2020_12_31_424971 393 46 ( ( -LRB- 10_1101-2020_12_31_424971 393 47 see see VB 10_1101-2020_12_31_424971 393 48 905 905 CD 10_1101-2020_12_31_424971 393 49 below below IN 10_1101-2020_12_31_424971 393 50 for for IN 10_1101-2020_12_31_424971 393 51 a a DT 10_1101-2020_12_31_424971 393 52 detailed detailed JJ 10_1101-2020_12_31_424971 393 53 description description NN 10_1101-2020_12_31_424971 393 54 of of IN 10_1101-2020_12_31_424971 393 55 how how WRB 10_1101-2020_12_31_424971 393 56 k70S k70S NNP 10_1101-2020_12_31_424971 393 57 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 393 58 , , , 10_1101-2020_12_31_424971 393 59 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 393 60 , , , 10_1101-2020_12_31_424971 393 61 and and CC 10_1101-2020_12_31_424971 393 62 other other JJ 10_1101-2020_12_31_424971 393 63 kinetic kinetic NN 10_1101-2020_12_31_424971 393 64 and and CC 10_1101-2020_12_31_424971 393 65 thermodynamic thermodynamic JJ 10_1101-2020_12_31_424971 393 66 906 906 CD 10_1101-2020_12_31_424971 393 67 .CC .CC , 10_1101-2020_12_31_424971 393 68 - - HYPH 10_1101-2020_12_31_424971 393 69 BY by IN 10_1101-2020_12_31_424971 393 70 - - HYPH 10_1101-2020_12_31_424971 393 71 NC NC NNP 10_1101-2020_12_31_424971 393 72 - - HYPH 10_1101-2020_12_31_424971 393 73 ND ND NNP 10_1101-2020_12_31_424971 393 74 4.0 4.0 CD 10_1101-2020_12_31_424971 393 75 International International NNP 10_1101-2020_12_31_424971 393 76 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 393 77 under under IN 10_1101-2020_12_31_424971 393 78 a a DT 10_1101-2020_12_31_424971 393 79 ( ( -LRB- 10_1101-2020_12_31_424971 393 80 which which WDT 10_1101-2020_12_31_424971 393 81 was be VBD 10_1101-2020_12_31_424971 393 82 not not RB 10_1101-2020_12_31_424971 393 83 certified certify VBN 10_1101-2020_12_31_424971 393 84 by by IN 10_1101-2020_12_31_424971 393 85 peer peer NN 10_1101-2020_12_31_424971 393 86 review review NN 10_1101-2020_12_31_424971 393 87 ) ) -RRB- 10_1101-2020_12_31_424971 393 88 is be VBZ 10_1101-2020_12_31_424971 393 89 the the DT 10_1101-2020_12_31_424971 393 90 author author NN 10_1101-2020_12_31_424971 393 91 / / SYM 10_1101-2020_12_31_424971 393 92 funder funder NN 10_1101-2020_12_31_424971 393 93 , , , 10_1101-2020_12_31_424971 393 94 who who WP 10_1101-2020_12_31_424971 393 95 has have VBZ 10_1101-2020_12_31_424971 393 96 granted grant VBN 10_1101-2020_12_31_424971 393 97 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 393 98 a a DT 10_1101-2020_12_31_424971 393 99 license license NN 10_1101-2020_12_31_424971 393 100 to to TO 10_1101-2020_12_31_424971 393 101 display display VB 10_1101-2020_12_31_424971 393 102 the the DT 10_1101-2020_12_31_424971 393 103 preprint preprint NN 10_1101-2020_12_31_424971 393 104 in in IN 10_1101-2020_12_31_424971 393 105 perpetuity perpetuity NN 10_1101-2020_12_31_424971 393 106 . . . 10_1101-2020_12_31_424971 394 1 It -PRON- PRP 10_1101-2020_12_31_424971 394 2 is be VBZ 10_1101-2020_12_31_424971 394 3 made make VBN 10_1101-2020_12_31_424971 394 4 The the DT 10_1101-2020_12_31_424971 394 5 copyright copyright NN 10_1101-2020_12_31_424971 394 6 holder holder NN 10_1101-2020_12_31_424971 394 7 for for IN 10_1101-2020_12_31_424971 394 8 this this DT 10_1101-2020_12_31_424971 394 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 394 10 version version NN 10_1101-2020_12_31_424971 394 11 posted post VBD 10_1101-2020_12_31_424971 394 12 January January NNP 10_1101-2020_12_31_424971 394 13 2 2 CD 10_1101-2020_12_31_424971 394 14 , , , 10_1101-2020_12_31_424971 394 15 2021 2021 CD 10_1101-2020_12_31_424971 394 16 . . . 10_1101-2020_12_31_424971 394 17 ; ; : 10_1101-2020_12_31_424971 394 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 394 19 : : : 10_1101-2020_12_31_424971 394 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 394 21 preprint preprint NN 10_1101-2020_12_31_424971 394 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 394 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 394 24 37 37 CD 10_1101-2020_12_31_424971 394 25 parameters parameter NNS 10_1101-2020_12_31_424971 394 26 were be VBD 10_1101-2020_12_31_424971 394 27 estimated estimate VBN 10_1101-2020_12_31_424971 394 28 ) ) -RRB- 10_1101-2020_12_31_424971 394 29 . . . 10_1101-2020_12_31_424971 395 1 Nonetheless nonetheless RB 10_1101-2020_12_31_424971 395 2 , , , 10_1101-2020_12_31_424971 395 3 the the DT 10_1101-2020_12_31_424971 395 4 number number NN 10_1101-2020_12_31_424971 395 5 of of IN 10_1101-2020_12_31_424971 395 6 such such JJ 10_1101-2020_12_31_424971 395 7 EFRET EFRET NNP 10_1101-2020_12_31_424971 395 8 vs. vs. IN 10_1101-2020_12_31_424971 395 9 time time NN 10_1101-2020_12_31_424971 395 10 trajectories trajectory NNS 10_1101-2020_12_31_424971 395 11 should should MD 10_1101-2020_12_31_424971 395 12 907 907 CD 10_1101-2020_12_31_424971 395 13 be be VB 10_1101-2020_12_31_424971 395 14 exceedingly exceedingly RB 10_1101-2020_12_31_424971 395 15 small small JJ 10_1101-2020_12_31_424971 395 16 . . . 10_1101-2020_12_31_424971 396 1 This this DT 10_1101-2020_12_31_424971 396 2 is be VBZ 10_1101-2020_12_31_424971 396 3 because because IN 10_1101-2020_12_31_424971 396 4 the the DT 10_1101-2020_12_31_424971 396 5 rates rate NNS 10_1101-2020_12_31_424971 396 6 with with IN 10_1101-2020_12_31_424971 396 7 which which WDT 10_1101-2020_12_31_424971 396 8 the the DT 10_1101-2020_12_31_424971 396 9 fluorophore fluorophore NN 10_1101-2020_12_31_424971 396 10 that that WDT 10_1101-2020_12_31_424971 396 11 photobleached photobleache VBD 10_1101-2020_12_31_424971 396 12 908 908 CD 10_1101-2020_12_31_424971 396 13 the the DT 10_1101-2020_12_31_424971 396 14 fastest fast JJS 10_1101-2020_12_31_424971 396 15 , , , 10_1101-2020_12_31_424971 396 16 Cy5 cy5 NN 10_1101-2020_12_31_424971 396 17 , , , 10_1101-2020_12_31_424971 396 18 entered enter VBD 10_1101-2020_12_31_424971 396 19 into into IN 10_1101-2020_12_31_424971 396 20 the the DT 10_1101-2020_12_31_424971 396 21 photobleached photobleache VBN 10_1101-2020_12_31_424971 396 22 state state NN 10_1101-2020_12_31_424971 396 23 ( ( -LRB- 10_1101-2020_12_31_424971 396 24 Æ æ NN 10_1101-2020_12_31_424971 396 25 ) ) -RRB- 10_1101-2020_12_31_424971 396 26 from from IN 10_1101-2020_12_31_424971 396 27 the the DT 10_1101-2020_12_31_424971 396 28 GS1 GS1 NNP 10_1101-2020_12_31_424971 396 29 , , , 10_1101-2020_12_31_424971 396 30 GS2 GS2 NNP 10_1101-2020_12_31_424971 396 31 , , , 10_1101-2020_12_31_424971 396 32 EF EF NNP 10_1101-2020_12_31_424971 396 33 - - HYPH 10_1101-2020_12_31_424971 396 34 G G NNP 10_1101-2020_12_31_424971 396 35 - - HYPH 10_1101-2020_12_31_424971 396 36 bound bind VBN 10_1101-2020_12_31_424971 396 37 GS2 GS2 NNP 10_1101-2020_12_31_424971 396 38 - - , 10_1101-2020_12_31_424971 396 39 909 909 CD 10_1101-2020_12_31_424971 396 40 like like IN 10_1101-2020_12_31_424971 396 41 , , , 10_1101-2020_12_31_424971 396 42 and and CC 10_1101-2020_12_31_424971 396 43 POST POST NNP 10_1101-2020_12_31_424971 396 44 states state NNS 10_1101-2020_12_31_424971 396 45 were be VBD 10_1101-2020_12_31_424971 396 46 kGS1→Æ kGS1→Æ NNP 10_1101-2020_12_31_424971 396 47 = = SYM 10_1101-2020_12_31_424971 396 48 0.04 0.04 CD 10_1101-2020_12_31_424971 396 49 ± ± CD 10_1101-2020_12_31_424971 396 50 0.02 0.02 CD 10_1101-2020_12_31_424971 396 51 s–1 s–1 NNS 10_1101-2020_12_31_424971 396 52 , , , 10_1101-2020_12_31_424971 396 53 kGS2→Æ kGS2→Æ NNP 10_1101-2020_12_31_424971 396 54 = = SYM 10_1101-2020_12_31_424971 396 55 0.07 0.07 CD 10_1101-2020_12_31_424971 396 56 ± ± CD 10_1101-2020_12_31_424971 396 57 0.01 0.01 CD 10_1101-2020_12_31_424971 396 58 s–1 s–1 NNS 10_1101-2020_12_31_424971 396 59 , , , 10_1101-2020_12_31_424971 396 60 kGS2(G)→Æ kGS2(G)→Æ NNP 10_1101-2020_12_31_424971 396 61 = = SYM 10_1101-2020_12_31_424971 396 62 0.07 0.07 CD 10_1101-2020_12_31_424971 396 63 ± ± CD 10_1101-2020_12_31_424971 396 64 910 910 CD 10_1101-2020_12_31_424971 396 65 0.01 0.01 CD 10_1101-2020_12_31_424971 396 66 s–1 s–1 NNS 10_1101-2020_12_31_424971 396 67 ( ( -LRB- 10_1101-2020_12_31_424971 396 68 where where WRB 10_1101-2020_12_31_424971 396 69 the the DT 10_1101-2020_12_31_424971 396 70 subscript subscript NN 10_1101-2020_12_31_424971 396 71 “ " `` 10_1101-2020_12_31_424971 396 72 ( ( -LRB- 10_1101-2020_12_31_424971 396 73 G g NN 10_1101-2020_12_31_424971 396 74 ) ) -RRB- 10_1101-2020_12_31_424971 396 75 ” " '' 10_1101-2020_12_31_424971 396 76 denotes denote VBZ 10_1101-2020_12_31_424971 396 77 experiments experiment NNS 10_1101-2020_12_31_424971 396 78 performed perform VBN 10_1101-2020_12_31_424971 396 79 in in IN 10_1101-2020_12_31_424971 396 80 the the DT 10_1101-2020_12_31_424971 396 81 presence presence NN 10_1101-2020_12_31_424971 396 82 of of IN 10_1101-2020_12_31_424971 396 83 EF EF NNP 10_1101-2020_12_31_424971 396 84 - - HYPH 10_1101-2020_12_31_424971 396 85 G G NNP 10_1101-2020_12_31_424971 396 86 ) ) -RRB- 10_1101-2020_12_31_424971 396 87 , , , 10_1101-2020_12_31_424971 396 88 and and CC 10_1101-2020_12_31_424971 396 89 911 911 CD 10_1101-2020_12_31_424971 396 90 kPOST→Æ kpost→æ CD 10_1101-2020_12_31_424971 396 91 0.05 0.05 CD 10_1101-2020_12_31_424971 396 92 ± ± CD 10_1101-2020_12_31_424971 396 93 0.02 0.02 CD 10_1101-2020_12_31_424971 396 94 s–1 s–1 NNS 10_1101-2020_12_31_424971 396 95 , , , 10_1101-2020_12_31_424971 396 96 respectively respectively RB 10_1101-2020_12_31_424971 396 97 ( ( -LRB- 10_1101-2020_12_31_424971 396 98 see see VB 10_1101-2020_12_31_424971 396 99 below below RP 10_1101-2020_12_31_424971 396 100 for for IN 10_1101-2020_12_31_424971 396 101 a a DT 10_1101-2020_12_31_424971 396 102 detailed detailed JJ 10_1101-2020_12_31_424971 396 103 description description NN 10_1101-2020_12_31_424971 396 104 of of IN 10_1101-2020_12_31_424971 396 105 how how WRB 10_1101-2020_12_31_424971 396 106 kGS1→Æ kGS1→Æ NNP 10_1101-2020_12_31_424971 396 107 , , , 10_1101-2020_12_31_424971 396 108 kGS2→Æ kGS2→Æ NNP 10_1101-2020_12_31_424971 396 109 , , , 10_1101-2020_12_31_424971 396 110 912 912 CD 10_1101-2020_12_31_424971 396 111 kGS2(G)→Æ kGS2(G)→Æ NNS 10_1101-2020_12_31_424971 396 112 , , , 10_1101-2020_12_31_424971 396 113 and and CC 10_1101-2020_12_31_424971 396 114 kPOST→Æ kpost→æ NN 10_1101-2020_12_31_424971 396 115 were be VBD 10_1101-2020_12_31_424971 396 116 estimated estimate VBN 10_1101-2020_12_31_424971 396 117 ) ) -RRB- 10_1101-2020_12_31_424971 396 118 . . . 10_1101-2020_12_31_424971 397 1 These these DT 10_1101-2020_12_31_424971 397 2 rates rate NNS 10_1101-2020_12_31_424971 397 3 are be VBP 10_1101-2020_12_31_424971 397 4 , , , 10_1101-2020_12_31_424971 397 5 on on IN 10_1101-2020_12_31_424971 397 6 average average JJ 10_1101-2020_12_31_424971 397 7 , , , 10_1101-2020_12_31_424971 397 8 about about RB 10_1101-2020_12_31_424971 397 9 11-fold 11-fold CD 10_1101-2020_12_31_424971 397 10 lower low JJR 10_1101-2020_12_31_424971 397 11 than than IN 10_1101-2020_12_31_424971 397 12 913 913 CD 10_1101-2020_12_31_424971 397 13 those those DT 10_1101-2020_12_31_424971 397 14 of of IN 10_1101-2020_12_31_424971 397 15 k70S k70s NN 10_1101-2020_12_31_424971 397 16 IC IC NNP 10_1101-2020_12_31_424971 397 17 → → SYM 10_1101-2020_12_31_424971 397 18 GS2 GS2 NNP 10_1101-2020_12_31_424971 397 19 and and CC 10_1101-2020_12_31_424971 397 20 kGS1 kGS1 NNP 10_1101-2020_12_31_424971 397 21 → → SYM 10_1101-2020_12_31_424971 397 22 GS2 GS2 NNP 10_1101-2020_12_31_424971 397 23 ( ( -LRB- 10_1101-2020_12_31_424971 397 24 0.3–0.6 0.3–0.6 CD 10_1101-2020_12_31_424971 397 25 s–1 s–1 NNS 10_1101-2020_12_31_424971 397 26 and and CC 10_1101-2020_12_31_424971 397 27 0.58–0.82 0.58–0.82 NN 10_1101-2020_12_31_424971 397 28 s–1 s–1 NNP 10_1101-2020_12_31_424971 397 29 ( ( -LRB- 10_1101-2020_12_31_424971 397 30 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 397 31 Table Table NNP 10_1101-2020_12_31_424971 397 32 4 4 CD 10_1101-2020_12_31_424971 397 33 ) ) -RRB- 10_1101-2020_12_31_424971 397 34 ) ) -RRB- 10_1101-2020_12_31_424971 397 35 . . . 10_1101-2020_12_31_424971 398 1 914 914 CD 10_1101-2020_12_31_424971 398 2 Consequently consequently RB 10_1101-2020_12_31_424971 398 3 , , , 10_1101-2020_12_31_424971 398 4 we -PRON- PRP 10_1101-2020_12_31_424971 398 5 do do VBP 10_1101-2020_12_31_424971 398 6 not not RB 10_1101-2020_12_31_424971 398 7 expect expect VB 10_1101-2020_12_31_424971 398 8 the the DT 10_1101-2020_12_31_424971 398 9 measurements measurement NNS 10_1101-2020_12_31_424971 398 10 of of IN 10_1101-2020_12_31_424971 398 11 k70S k70S NNP 10_1101-2020_12_31_424971 398 12 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 398 13 and and CC 10_1101-2020_12_31_424971 398 14 kGS1→GS2 kGS1→GS2 NNP 10_1101-2020_12_31_424971 398 15 to to TO 10_1101-2020_12_31_424971 398 16 be be VB 10_1101-2020_12_31_424971 398 17 limited limit VBN 10_1101-2020_12_31_424971 398 18 by by IN 10_1101-2020_12_31_424971 398 19 915 915 CD 10_1101-2020_12_31_424971 398 20 Cy3 cy3 CD 10_1101-2020_12_31_424971 398 21 or or CC 10_1101-2020_12_31_424971 398 22 Cy5 cy5 NN 10_1101-2020_12_31_424971 398 23 photobleaching photobleaching NN 10_1101-2020_12_31_424971 398 24 . . . 10_1101-2020_12_31_424971 399 1 Additionally additionally RB 10_1101-2020_12_31_424971 399 2 , , , 10_1101-2020_12_31_424971 399 3 even even RB 10_1101-2020_12_31_424971 399 4 if if IN 10_1101-2020_12_31_424971 399 5 k70S k70S NNP 10_1101-2020_12_31_424971 399 6 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 399 7 and and CC 10_1101-2020_12_31_424971 399 8 kGS1→GS2 kGS1→GS2 NNP 10_1101-2020_12_31_424971 399 9 were be VBD 10_1101-2020_12_31_424971 399 10 slightly slightly RB 10_1101-2020_12_31_424971 399 11 916 916 CD 10_1101-2020_12_31_424971 399 12 overestimated overestimate VBN 10_1101-2020_12_31_424971 399 13 , , , 10_1101-2020_12_31_424971 399 14 they -PRON- PRP 10_1101-2020_12_31_424971 399 15 would would MD 10_1101-2020_12_31_424971 399 16 be be VB 10_1101-2020_12_31_424971 399 17 expected expect VBN 10_1101-2020_12_31_424971 399 18 to to TO 10_1101-2020_12_31_424971 399 19 be be VB 10_1101-2020_12_31_424971 399 20 equally equally RB 10_1101-2020_12_31_424971 399 21 overestimated overestimate VBN 10_1101-2020_12_31_424971 399 22 for for IN 10_1101-2020_12_31_424971 399 23 SufB2- SufB2- NNP 10_1101-2020_12_31_424971 399 24 and and CC 10_1101-2020_12_31_424971 399 25 ProL ProL NNP 10_1101-2020_12_31_424971 399 26 917 917 CD 10_1101-2020_12_31_424971 399 27 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 399 28 complexes complex NNS 10_1101-2020_12_31_424971 399 29 given give VBN 10_1101-2020_12_31_424971 399 30 that that IN 10_1101-2020_12_31_424971 399 31 the the DT 10_1101-2020_12_31_424971 399 32 rate rate NN 10_1101-2020_12_31_424971 399 33 of of IN 10_1101-2020_12_31_424971 399 34 photobleaching photobleaching NN 10_1101-2020_12_31_424971 399 35 would would MD 10_1101-2020_12_31_424971 399 36 be be VB 10_1101-2020_12_31_424971 399 37 expected expect VBN 10_1101-2020_12_31_424971 399 38 to to TO 10_1101-2020_12_31_424971 399 39 be be VB 10_1101-2020_12_31_424971 399 40 very very RB 10_1101-2020_12_31_424971 399 41 similar similar JJ 10_1101-2020_12_31_424971 399 42 918 918 CD 10_1101-2020_12_31_424971 399 43 for for IN 10_1101-2020_12_31_424971 399 44 SufB2- SufB2- NNP 10_1101-2020_12_31_424971 399 45 and and CC 10_1101-2020_12_31_424971 399 46 ProL ProL NNP 10_1101-2020_12_31_424971 399 47 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 399 48 complexes complex NNS 10_1101-2020_12_31_424971 399 49 . . . 10_1101-2020_12_31_424971 400 1 Furthermore furthermore RB 10_1101-2020_12_31_424971 400 2 , , , 10_1101-2020_12_31_424971 400 3 because because IN 10_1101-2020_12_31_424971 400 4 we -PRON- PRP 10_1101-2020_12_31_424971 400 5 are be VBP 10_1101-2020_12_31_424971 400 6 primarily primarily RB 10_1101-2020_12_31_424971 400 7 concerned concerned JJ 10_1101-2020_12_31_424971 400 8 with with IN 10_1101-2020_12_31_424971 400 9 919 919 CD 10_1101-2020_12_31_424971 400 10 the the DT 10_1101-2020_12_31_424971 400 11 relative relative JJ 10_1101-2020_12_31_424971 400 12 values value NNS 10_1101-2020_12_31_424971 400 13 of of IN 10_1101-2020_12_31_424971 400 14 k70S k70S NNP 10_1101-2020_12_31_424971 400 15 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 400 16 and and CC 10_1101-2020_12_31_424971 400 17 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 400 18 for for IN 10_1101-2020_12_31_424971 400 19 SufB2- SufB2- NNP 10_1101-2020_12_31_424971 400 20 vs. vs. IN 10_1101-2020_12_31_424971 400 21 ProL ProL NNP 10_1101-2020_12_31_424971 400 22 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 400 23 complexes complex NNS 10_1101-2020_12_31_424971 400 24 , , , 10_1101-2020_12_31_424971 400 25 rather rather RB 10_1101-2020_12_31_424971 400 26 920 920 CD 10_1101-2020_12_31_424971 400 27 than than IN 10_1101-2020_12_31_424971 400 28 with with IN 10_1101-2020_12_31_424971 400 29 the the DT 10_1101-2020_12_31_424971 400 30 absolute absolute JJ 10_1101-2020_12_31_424971 400 31 values value NNS 10_1101-2020_12_31_424971 400 32 of of IN 10_1101-2020_12_31_424971 400 33 k70S k70S NNP 10_1101-2020_12_31_424971 400 34 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 400 35 and and CC 10_1101-2020_12_31_424971 400 36 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 400 37 for for IN 10_1101-2020_12_31_424971 400 38 the the DT 10_1101-2020_12_31_424971 400 39 SufB2- SufB2- NNP 10_1101-2020_12_31_424971 400 40 and and CC 10_1101-2020_12_31_424971 400 41 ProL ProL NNP 10_1101-2020_12_31_424971 400 42 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 400 43 921 921 CD 10_1101-2020_12_31_424971 400 44 complexes complex NNS 10_1101-2020_12_31_424971 400 45 , , , 10_1101-2020_12_31_424971 400 46 such such JJ 10_1101-2020_12_31_424971 400 47 slight slight JJ 10_1101-2020_12_31_424971 400 48 overestimations overestimation NNS 10_1101-2020_12_31_424971 400 49 do do VBP 10_1101-2020_12_31_424971 400 50 not not RB 10_1101-2020_12_31_424971 400 51 affect affect VB 10_1101-2020_12_31_424971 400 52 the the DT 10_1101-2020_12_31_424971 400 53 conclusions conclusion NNS 10_1101-2020_12_31_424971 400 54 of of IN 10_1101-2020_12_31_424971 400 55 the the DT 10_1101-2020_12_31_424971 400 56 work work NN 10_1101-2020_12_31_424971 400 57 presented present VBN 10_1101-2020_12_31_424971 400 58 here here RB 10_1101-2020_12_31_424971 400 59 . . . 10_1101-2020_12_31_424971 401 1 922 922 CD 10_1101-2020_12_31_424971 401 2 To to TO 10_1101-2020_12_31_424971 401 3 calculate calculate VB 10_1101-2020_12_31_424971 401 4 k70S k70s ADD 10_1101-2020_12_31_424971 401 5 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 401 6 and and CC 10_1101-2020_12_31_424971 401 7 the the DT 10_1101-2020_12_31_424971 401 8 corresponding corresponding JJ 10_1101-2020_12_31_424971 401 9 error error NN 10_1101-2020_12_31_424971 401 10 from from IN 10_1101-2020_12_31_424971 401 11 the the DT 10_1101-2020_12_31_424971 401 12 pre pre JJ 10_1101-2020_12_31_424971 401 13 - - JJ 10_1101-2020_12_31_424971 401 14 steady steady JJ 10_1101-2020_12_31_424971 401 15 - - HYPH 10_1101-2020_12_31_424971 401 16 state state NN 10_1101-2020_12_31_424971 401 17 experiments experiment NNS 10_1101-2020_12_31_424971 401 18 , , , 10_1101-2020_12_31_424971 401 19 923 923 CD 10_1101-2020_12_31_424971 401 20 we -PRON- PRP 10_1101-2020_12_31_424971 401 21 analyzed analyze VBD 10_1101-2020_12_31_424971 401 22 the the DT 10_1101-2020_12_31_424971 401 23 70S 70S NNP 10_1101-2020_12_31_424971 401 24 IC IC NNP 10_1101-2020_12_31_424971 401 25 survival survival NN 10_1101-2020_12_31_424971 401 26 probabilities probability NNS 10_1101-2020_12_31_424971 401 27 ( ( -LRB- 10_1101-2020_12_31_424971 401 28 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 401 29 Figure Figure NNP 10_1101-2020_12_31_424971 401 30 4 4 CD 10_1101-2020_12_31_424971 401 31 , , , 10_1101-2020_12_31_424971 401 32 Tables Tables NNP 10_1101-2020_12_31_424971 401 33 4 4 CD 10_1101-2020_12_31_424971 401 34 and and CC 10_1101-2020_12_31_424971 401 35 5)37,67 5)37,67 CD 10_1101-2020_12_31_424971 401 36 . . . 10_1101-2020_12_31_424971 402 1 924 924 CD 10_1101-2020_12_31_424971 402 2 Briefly briefly RB 10_1101-2020_12_31_424971 402 3 , , , 10_1101-2020_12_31_424971 402 4 for for IN 10_1101-2020_12_31_424971 402 5 each each DT 10_1101-2020_12_31_424971 402 6 trajectory trajectory CD 10_1101-2020_12_31_424971 402 7 , , , 10_1101-2020_12_31_424971 402 8 we -PRON- PRP 10_1101-2020_12_31_424971 402 9 extracted extract VBD 10_1101-2020_12_31_424971 402 10 the the DT 10_1101-2020_12_31_424971 402 11 time time NN 10_1101-2020_12_31_424971 402 12 interval interval NN 10_1101-2020_12_31_424971 402 13 during during IN 10_1101-2020_12_31_424971 402 14 which which WDT 10_1101-2020_12_31_424971 402 15 we -PRON- PRP 10_1101-2020_12_31_424971 402 16 were be VBD 10_1101-2020_12_31_424971 402 17 waiting wait VBG 10_1101-2020_12_31_424971 402 18 for for IN 10_1101-2020_12_31_424971 402 19 the the DT 10_1101-2020_12_31_424971 402 20 925 925 CD 10_1101-2020_12_31_424971 402 21 70S 70S NNP 10_1101-2020_12_31_424971 402 22 IC IC NNP 10_1101-2020_12_31_424971 402 23 to to TO 10_1101-2020_12_31_424971 402 24 undergo undergo VB 10_1101-2020_12_31_424971 402 25 a a DT 10_1101-2020_12_31_424971 402 26 transition transition NN 10_1101-2020_12_31_424971 402 27 to to IN 10_1101-2020_12_31_424971 402 28 GS2 GS2 NNP 10_1101-2020_12_31_424971 402 29 and and CC 10_1101-2020_12_31_424971 402 30 used use VBD 10_1101-2020_12_31_424971 402 31 these these DT 10_1101-2020_12_31_424971 402 32 ‘ ' `` 10_1101-2020_12_31_424971 402 33 waiting wait VBG 10_1101-2020_12_31_424971 402 34 times time NNS 10_1101-2020_12_31_424971 402 35 ’ ' '' 10_1101-2020_12_31_424971 402 36 to to TO 10_1101-2020_12_31_424971 402 37 construct construct VB 10_1101-2020_12_31_424971 402 38 a a DT 10_1101-2020_12_31_424971 402 39 70S 70S NNP 10_1101-2020_12_31_424971 402 40 IC IC NNP 10_1101-2020_12_31_424971 402 41 survival survival NN 10_1101-2020_12_31_424971 402 42 926 926 CD 10_1101-2020_12_31_424971 402 43 probability probability NN 10_1101-2020_12_31_424971 402 44 distribution distribution NN 10_1101-2020_12_31_424971 402 45 , , , 10_1101-2020_12_31_424971 402 46 as as IN 10_1101-2020_12_31_424971 402 47 shown show VBN 10_1101-2020_12_31_424971 402 48 in in IN 10_1101-2020_12_31_424971 402 49 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 402 50 Figure Figure NNP 10_1101-2020_12_31_424971 402 51 4 4 CD 10_1101-2020_12_31_424971 402 52 . . . 10_1101-2020_12_31_424971 403 1 All all DT 10_1101-2020_12_31_424971 403 2 70S 70S NNP 10_1101-2020_12_31_424971 403 3 IC IC NNP 10_1101-2020_12_31_424971 403 4 survival survival NN 10_1101-2020_12_31_424971 403 5 probability probability NN 10_1101-2020_12_31_424971 403 6 927 927 CD 10_1101-2020_12_31_424971 403 7 distributions distribution NNS 10_1101-2020_12_31_424971 403 8 were be VBD 10_1101-2020_12_31_424971 403 9 best well RBS 10_1101-2020_12_31_424971 403 10 described describe VBN 10_1101-2020_12_31_424971 403 11 by by IN 10_1101-2020_12_31_424971 403 12 a a DT 10_1101-2020_12_31_424971 403 13 single single JJ 10_1101-2020_12_31_424971 403 14 exponential exponential JJ 10_1101-2020_12_31_424971 403 15 decay decay NN 10_1101-2020_12_31_424971 403 16 function function NN 10_1101-2020_12_31_424971 403 17 of of IN 10_1101-2020_12_31_424971 403 18 the the DT 10_1101-2020_12_31_424971 403 19 type type NN 10_1101-2020_12_31_424971 403 20 928 928 NNP 10_1101-2020_12_31_424971 403 21 𝑌 𝑌 NNP 10_1101-2020_12_31_424971 403 22 = = SYM 10_1101-2020_12_31_424971 403 23 𝐴e("#/𝜏 𝐴e("#/𝜏 NNP 10_1101-2020_12_31_424971 403 24 ! ! . 10_1101-2020_12_31_424971 404 1 " " `` 10_1101-2020_12_31_424971 404 2 # # $ 10_1101-2020_12_31_424971 404 3 % % NN 10_1101-2020_12_31_424971 404 4 & & CC 10_1101-2020_12_31_424971 404 5 ) ) -RRB- 10_1101-2020_12_31_424971 404 6 , , , 10_1101-2020_12_31_424971 404 7 ( ( -LRB- 10_1101-2020_12_31_424971 404 8 1 1 LS 10_1101-2020_12_31_424971 404 9 ) ) -RRB- 10_1101-2020_12_31_424971 404 10 929 929 CD 10_1101-2020_12_31_424971 404 11 where where WRB 10_1101-2020_12_31_424971 404 12 Y Y NNP 10_1101-2020_12_31_424971 404 13 is be VBZ 10_1101-2020_12_31_424971 404 14 survival survival NN 10_1101-2020_12_31_424971 404 15 probability probability NN 10_1101-2020_12_31_424971 404 16 , , , 10_1101-2020_12_31_424971 404 17 A A NNP 10_1101-2020_12_31_424971 404 18 is be VBZ 10_1101-2020_12_31_424971 404 19 the the DT 10_1101-2020_12_31_424971 404 20 initial initial JJ 10_1101-2020_12_31_424971 404 21 population population NN 10_1101-2020_12_31_424971 404 22 of of IN 10_1101-2020_12_31_424971 404 23 70S 70S NNP 10_1101-2020_12_31_424971 404 24 IC IC NNP 10_1101-2020_12_31_424971 404 25 , , , 10_1101-2020_12_31_424971 404 26 t t NNP 10_1101-2020_12_31_424971 404 27 is be VBZ 10_1101-2020_12_31_424971 404 28 time time NN 10_1101-2020_12_31_424971 404 29 , , , 10_1101-2020_12_31_424971 404 30 and and CC 10_1101-2020_12_31_424971 404 31 τ70S τ70S `` 10_1101-2020_12_31_424971 404 32 IC IC NNP 10_1101-2020_12_31_424971 404 33 is be VBZ 10_1101-2020_12_31_424971 404 34 the the DT 10_1101-2020_12_31_424971 404 35 930 930 CD 10_1101-2020_12_31_424971 404 36 time time NN 10_1101-2020_12_31_424971 404 37 constant constant JJ 10_1101-2020_12_31_424971 404 38 with with IN 10_1101-2020_12_31_424971 404 39 which which WDT 10_1101-2020_12_31_424971 404 40 70S 70S NNP 10_1101-2020_12_31_424971 404 41 IC IC NNP 10_1101-2020_12_31_424971 404 42 transitions transition VBZ 10_1101-2020_12_31_424971 404 43 to to IN 10_1101-2020_12_31_424971 404 44 a a DT 10_1101-2020_12_31_424971 404 45 PRE PRE NNP 10_1101-2020_12_31_424971 404 46 complex complex NN 10_1101-2020_12_31_424971 404 47 in in IN 10_1101-2020_12_31_424971 404 48 the the DT 10_1101-2020_12_31_424971 404 49 GS2 GS2 NNP 10_1101-2020_12_31_424971 404 50 state state NN 10_1101-2020_12_31_424971 404 51 . . . 10_1101-2020_12_31_424971 405 1 k70S k70S NNP 10_1101-2020_12_31_424971 405 2 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 405 3 was be VBD 10_1101-2020_12_31_424971 405 4 931 931 CD 10_1101-2020_12_31_424971 405 5 .CC .CC : 10_1101-2020_12_31_424971 405 6 - - HYPH 10_1101-2020_12_31_424971 405 7 BY by IN 10_1101-2020_12_31_424971 405 8 - - HYPH 10_1101-2020_12_31_424971 405 9 NC NC NNP 10_1101-2020_12_31_424971 405 10 - - HYPH 10_1101-2020_12_31_424971 405 11 ND ND NNP 10_1101-2020_12_31_424971 405 12 4.0 4.0 CD 10_1101-2020_12_31_424971 405 13 International International NNP 10_1101-2020_12_31_424971 405 14 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 405 15 under under IN 10_1101-2020_12_31_424971 405 16 a a DT 10_1101-2020_12_31_424971 405 17 ( ( -LRB- 10_1101-2020_12_31_424971 405 18 which which WDT 10_1101-2020_12_31_424971 405 19 was be VBD 10_1101-2020_12_31_424971 405 20 not not RB 10_1101-2020_12_31_424971 405 21 certified certify VBN 10_1101-2020_12_31_424971 405 22 by by IN 10_1101-2020_12_31_424971 405 23 peer peer NN 10_1101-2020_12_31_424971 405 24 review review NN 10_1101-2020_12_31_424971 405 25 ) ) -RRB- 10_1101-2020_12_31_424971 405 26 is be VBZ 10_1101-2020_12_31_424971 405 27 the the DT 10_1101-2020_12_31_424971 405 28 author author NN 10_1101-2020_12_31_424971 405 29 / / SYM 10_1101-2020_12_31_424971 405 30 funder funder NN 10_1101-2020_12_31_424971 405 31 , , , 10_1101-2020_12_31_424971 405 32 who who WP 10_1101-2020_12_31_424971 405 33 has have VBZ 10_1101-2020_12_31_424971 405 34 granted grant VBN 10_1101-2020_12_31_424971 405 35 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 405 36 a a DT 10_1101-2020_12_31_424971 405 37 license license NN 10_1101-2020_12_31_424971 405 38 to to TO 10_1101-2020_12_31_424971 405 39 display display VB 10_1101-2020_12_31_424971 405 40 the the DT 10_1101-2020_12_31_424971 405 41 preprint preprint NN 10_1101-2020_12_31_424971 405 42 in in IN 10_1101-2020_12_31_424971 405 43 perpetuity perpetuity NN 10_1101-2020_12_31_424971 405 44 . . . 10_1101-2020_12_31_424971 406 1 It -PRON- PRP 10_1101-2020_12_31_424971 406 2 is be VBZ 10_1101-2020_12_31_424971 406 3 made make VBN 10_1101-2020_12_31_424971 406 4 The the DT 10_1101-2020_12_31_424971 406 5 copyright copyright NN 10_1101-2020_12_31_424971 406 6 holder holder NN 10_1101-2020_12_31_424971 406 7 for for IN 10_1101-2020_12_31_424971 406 8 this this DT 10_1101-2020_12_31_424971 406 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 406 10 version version NN 10_1101-2020_12_31_424971 406 11 posted post VBD 10_1101-2020_12_31_424971 406 12 January January NNP 10_1101-2020_12_31_424971 406 13 2 2 CD 10_1101-2020_12_31_424971 406 14 , , , 10_1101-2020_12_31_424971 406 15 2021 2021 CD 10_1101-2020_12_31_424971 406 16 . . . 10_1101-2020_12_31_424971 406 17 ; ; : 10_1101-2020_12_31_424971 406 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 406 19 : : : 10_1101-2020_12_31_424971 406 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 406 21 preprint preprint NN 10_1101-2020_12_31_424971 406 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NNP 10_1101-2020_12_31_424971 406 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 406 24 38 38 CD 10_1101-2020_12_31_424971 406 25 then then RB 10_1101-2020_12_31_424971 406 26 calculated calculate VBN 10_1101-2020_12_31_424971 406 27 using use VBG 10_1101-2020_12_31_424971 406 28 the the DT 10_1101-2020_12_31_424971 406 29 equation equation NN 10_1101-2020_12_31_424971 406 30 k70S k70s ADD 10_1101-2020_12_31_424971 406 31 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 406 32 = = SYM 10_1101-2020_12_31_424971 406 33 1 1 CD 10_1101-2020_12_31_424971 406 34 / / SYM 10_1101-2020_12_31_424971 406 35 τ70S τ70s NN 10_1101-2020_12_31_424971 406 36 IC IC NNP 10_1101-2020_12_31_424971 406 37 . . . 10_1101-2020_12_31_424971 407 1 Errors error NNS 10_1101-2020_12_31_424971 407 2 were be VBD 10_1101-2020_12_31_424971 407 3 calculated calculate VBN 10_1101-2020_12_31_424971 407 4 as as IN 10_1101-2020_12_31_424971 407 5 the the DT 10_1101-2020_12_31_424971 407 6 standard standard JJ 10_1101-2020_12_31_424971 407 7 932 932 CD 10_1101-2020_12_31_424971 407 8 deviation deviation NN 10_1101-2020_12_31_424971 407 9 of of IN 10_1101-2020_12_31_424971 407 10 technical technical JJ 10_1101-2020_12_31_424971 407 11 triplicates triplicate NNS 10_1101-2020_12_31_424971 407 12 . . . 10_1101-2020_12_31_424971 408 1 933 933 CD 10_1101-2020_12_31_424971 408 2 Six six CD 10_1101-2020_12_31_424971 408 3 sets set NNS 10_1101-2020_12_31_424971 408 4 of of IN 10_1101-2020_12_31_424971 408 5 kinetic kinetic NN 10_1101-2020_12_31_424971 408 6 and/or and/or CC 10_1101-2020_12_31_424971 408 7 thermodynamic thermodynamic JJ 10_1101-2020_12_31_424971 408 8 parameters parameter NNS 10_1101-2020_12_31_424971 408 9 were be VBD 10_1101-2020_12_31_424971 408 10 calculated calculate VBN 10_1101-2020_12_31_424971 408 11 from from IN 10_1101-2020_12_31_424971 408 12 hidden hide VBN 10_1101-2020_12_31_424971 408 13 Markov Markov NNP 10_1101-2020_12_31_424971 408 14 934 934 CD 10_1101-2020_12_31_424971 408 15 model model NN 10_1101-2020_12_31_424971 408 16 ( ( -LRB- 10_1101-2020_12_31_424971 408 17 HMM HMM NNP 10_1101-2020_12_31_424971 408 18 ) ) -RRB- 10_1101-2020_12_31_424971 408 19 analyses analysis NNS 10_1101-2020_12_31_424971 408 20 of of IN 10_1101-2020_12_31_424971 408 21 the the DT 10_1101-2020_12_31_424971 408 22 recorded record VBN 10_1101-2020_12_31_424971 408 23 movies movie NNS 10_1101-2020_12_31_424971 408 24 . . . 10_1101-2020_12_31_424971 409 1 These these DT 10_1101-2020_12_31_424971 409 2 parameters parameter NNS 10_1101-2020_12_31_424971 409 3 are be VBP 10_1101-2020_12_31_424971 409 4 defined define VBN 10_1101-2020_12_31_424971 409 5 here here RB 10_1101-2020_12_31_424971 409 6 as as IN 10_1101-2020_12_31_424971 409 7 : : : 10_1101-2020_12_31_424971 409 8 ( ( -LRB- 10_1101-2020_12_31_424971 409 9 i i NN 10_1101-2020_12_31_424971 409 10 ) ) -RRB- 10_1101-2020_12_31_424971 409 11 935 935 CD 10_1101-2020_12_31_424971 409 12 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 409 13 , , , 10_1101-2020_12_31_424971 409 14 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 409 15 , , , 10_1101-2020_12_31_424971 409 16 and and CC 10_1101-2020_12_31_424971 409 17 Keq Keq NNP 10_1101-2020_12_31_424971 409 18 from from IN 10_1101-2020_12_31_424971 409 19 the the DT 10_1101-2020_12_31_424971 409 20 pre pre JJ 10_1101-2020_12_31_424971 409 21 - - JJ 10_1101-2020_12_31_424971 409 22 steady steady JJ 10_1101-2020_12_31_424971 409 23 - - HYPH 10_1101-2020_12_31_424971 409 24 state state NN 10_1101-2020_12_31_424971 409 25 and and CC 10_1101-2020_12_31_424971 409 26 steady steady JJ 10_1101-2020_12_31_424971 409 27 - - HYPH 10_1101-2020_12_31_424971 409 28 state state NN 10_1101-2020_12_31_424971 409 29 movies movie NNS 10_1101-2020_12_31_424971 409 30 recorded record VBN 10_1101-2020_12_31_424971 409 31 for for IN 10_1101-2020_12_31_424971 409 32 the the DT 10_1101-2020_12_31_424971 409 33 936 936 CD 10_1101-2020_12_31_424971 409 34 delivery delivery NN 10_1101-2020_12_31_424971 409 35 of of IN 10_1101-2020_12_31_424971 409 36 SufB2- SufB2- NNP 10_1101-2020_12_31_424971 409 37 and and CC 10_1101-2020_12_31_424971 409 38 ProL ProL NNP 10_1101-2020_12_31_424971 409 39 - - HYPH 10_1101-2020_12_31_424971 409 40 TCs tc NNS 10_1101-2020_12_31_424971 409 41 in in IN 10_1101-2020_12_31_424971 409 42 the the DT 10_1101-2020_12_31_424971 409 43 absence absence NN 10_1101-2020_12_31_424971 409 44 of of IN 10_1101-2020_12_31_424971 409 45 EF EF NNP 10_1101-2020_12_31_424971 409 46 - - HYPH 10_1101-2020_12_31_424971 409 47 G G NNP 10_1101-2020_12_31_424971 409 48 ( ( -LRB- 10_1101-2020_12_31_424971 409 49 Figures Figures NNP 10_1101-2020_12_31_424971 409 50 6d 6d NNP 10_1101-2020_12_31_424971 409 51 , , , 10_1101-2020_12_31_424971 409 52 6f 6f RB 10_1101-2020_12_31_424971 409 53 , , , 10_1101-2020_12_31_424971 409 54 and and CC 10_1101-2020_12_31_424971 409 55 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 409 56 937 937 CD 10_1101-2020_12_31_424971 409 57 Figure figure NN 10_1101-2020_12_31_424971 409 58 3 3 CD 10_1101-2020_12_31_424971 409 59 and and CC 10_1101-2020_12_31_424971 409 60 Table table NN 10_1101-2020_12_31_424971 409 61 4 4 CD 10_1101-2020_12_31_424971 409 62 ) ) -RRB- 10_1101-2020_12_31_424971 409 63 ; ; : 10_1101-2020_12_31_424971 409 64 ( ( -LRB- 10_1101-2020_12_31_424971 409 65 ii ii NNP 10_1101-2020_12_31_424971 409 66 ) ) -RRB- 10_1101-2020_12_31_424971 409 67 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 409 68 from from IN 10_1101-2020_12_31_424971 409 69 the the DT 10_1101-2020_12_31_424971 409 70 pre pre JJ 10_1101-2020_12_31_424971 409 71 - - JJ 10_1101-2020_12_31_424971 409 72 steady steady JJ 10_1101-2020_12_31_424971 409 73 - - HYPH 10_1101-2020_12_31_424971 409 74 state state NN 10_1101-2020_12_31_424971 409 75 movie movie NN 10_1101-2020_12_31_424971 409 76 recorded record VBN 10_1101-2020_12_31_424971 409 77 for for IN 10_1101-2020_12_31_424971 409 78 the the DT 10_1101-2020_12_31_424971 409 79 delivery delivery NN 10_1101-2020_12_31_424971 409 80 of of IN 10_1101-2020_12_31_424971 409 81 938 938 CD 10_1101-2020_12_31_424971 409 82 ProL ProL NNP 10_1101-2020_12_31_424971 409 83 - - HYPH 10_1101-2020_12_31_424971 409 84 TC TC NNP 10_1101-2020_12_31_424971 409 85 in in IN 10_1101-2020_12_31_424971 409 86 the the DT 10_1101-2020_12_31_424971 409 87 presence presence NN 10_1101-2020_12_31_424971 409 88 of of IN 10_1101-2020_12_31_424971 409 89 EF EF NNP 10_1101-2020_12_31_424971 409 90 - - HYPH 10_1101-2020_12_31_424971 409 91 G G NNP 10_1101-2020_12_31_424971 409 92 ( ( -LRB- 10_1101-2020_12_31_424971 409 93 Figures Figures NNP 10_1101-2020_12_31_424971 409 94 6e 6e NNP 10_1101-2020_12_31_424971 409 95 , , , 10_1101-2020_12_31_424971 409 96 6 6 CD 10_1101-2020_12_31_424971 409 97 g g NN 10_1101-2020_12_31_424971 409 98 , , , 10_1101-2020_12_31_424971 409 99 and and CC 10_1101-2020_12_31_424971 409 100 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 409 101 Figure Figure NNP 10_1101-2020_12_31_424971 409 102 5 5 CD 10_1101-2020_12_31_424971 409 103 and and CC 10_1101-2020_12_31_424971 409 104 Table table NN 10_1101-2020_12_31_424971 409 105 5 5 CD 10_1101-2020_12_31_424971 409 106 ) ) -RRB- 10_1101-2020_12_31_424971 409 107 ; ; : 10_1101-2020_12_31_424971 409 108 ( ( -LRB- 10_1101-2020_12_31_424971 409 109 iii iii NN 10_1101-2020_12_31_424971 409 110 ) ) -RRB- 10_1101-2020_12_31_424971 409 111 939 939 CD 10_1101-2020_12_31_424971 409 112 the the DT 10_1101-2020_12_31_424971 409 113 fractional fractional JJ 10_1101-2020_12_31_424971 409 114 population population NN 10_1101-2020_12_31_424971 409 115 of of IN 10_1101-2020_12_31_424971 409 116 the the DT 10_1101-2020_12_31_424971 409 117 POST POST NNP 10_1101-2020_12_31_424971 409 118 complex complex NN 10_1101-2020_12_31_424971 409 119 from from IN 10_1101-2020_12_31_424971 409 120 the the DT 10_1101-2020_12_31_424971 409 121 pre pre JJ 10_1101-2020_12_31_424971 409 122 - - JJ 10_1101-2020_12_31_424971 409 123 steady steady JJ 10_1101-2020_12_31_424971 409 124 - - HYPH 10_1101-2020_12_31_424971 409 125 state state NN 10_1101-2020_12_31_424971 409 126 and and CC 10_1101-2020_12_31_424971 409 127 steady steady JJ 10_1101-2020_12_31_424971 409 128 - - HYPH 10_1101-2020_12_31_424971 409 129 state state NN 10_1101-2020_12_31_424971 409 130 movies movie NNS 10_1101-2020_12_31_424971 409 131 940 940 CD 10_1101-2020_12_31_424971 409 132 recorded record VBN 10_1101-2020_12_31_424971 409 133 for for IN 10_1101-2020_12_31_424971 409 134 the the DT 10_1101-2020_12_31_424971 409 135 delivery delivery NN 10_1101-2020_12_31_424971 409 136 of of IN 10_1101-2020_12_31_424971 409 137 SufB2- SufB2- NNP 10_1101-2020_12_31_424971 409 138 and and CC 10_1101-2020_12_31_424971 409 139 ProL ProL NNP 10_1101-2020_12_31_424971 409 140 - - HYPH 10_1101-2020_12_31_424971 409 141 TCs tc NNS 10_1101-2020_12_31_424971 409 142 in in IN 10_1101-2020_12_31_424971 409 143 the the DT 10_1101-2020_12_31_424971 409 144 presence presence NN 10_1101-2020_12_31_424971 409 145 of of IN 10_1101-2020_12_31_424971 409 146 EF EF NNP 10_1101-2020_12_31_424971 409 147 - - HYPH 10_1101-2020_12_31_424971 409 148 G G NNP 10_1101-2020_12_31_424971 409 149 ( ( -LRB- 10_1101-2020_12_31_424971 409 150 Figures Figures NNP 10_1101-2020_12_31_424971 409 151 6e 6e NNP 10_1101-2020_12_31_424971 409 152 , , , 10_1101-2020_12_31_424971 409 153 6 6 CD 10_1101-2020_12_31_424971 409 154 g g NN 10_1101-2020_12_31_424971 409 155 , , , 10_1101-2020_12_31_424971 409 156 and and CC 10_1101-2020_12_31_424971 409 157 941 941 CD 10_1101-2020_12_31_424971 409 158 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 409 159 Figure figure NN 10_1101-2020_12_31_424971 409 160 5 5 CD 10_1101-2020_12_31_424971 409 161 and and CC 10_1101-2020_12_31_424971 409 162 Table table NN 10_1101-2020_12_31_424971 409 163 5 5 CD 10_1101-2020_12_31_424971 409 164 ) ) -RRB- 10_1101-2020_12_31_424971 409 165 ; ; : 10_1101-2020_12_31_424971 409 166 ( ( -LRB- 10_1101-2020_12_31_424971 409 167 iv iv NN 10_1101-2020_12_31_424971 409 168 ) ) -RRB- 10_1101-2020_12_31_424971 409 169 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 409 170 , , , 10_1101-2020_12_31_424971 409 171 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 409 172 , , , 10_1101-2020_12_31_424971 409 173 and and CC 10_1101-2020_12_31_424971 409 174 Keq Keq NNP 10_1101-2020_12_31_424971 409 175 from from IN 10_1101-2020_12_31_424971 409 176 a a DT 10_1101-2020_12_31_424971 409 177 sub sub NN 10_1101-2020_12_31_424971 409 178 - - NN 10_1101-2020_12_31_424971 409 179 population population NN 10_1101-2020_12_31_424971 409 180 of of IN 10_1101-2020_12_31_424971 409 181 942 942 CD 10_1101-2020_12_31_424971 409 182 PRE PRE NNP 10_1101-2020_12_31_424971 409 183 complexes complex NNS 10_1101-2020_12_31_424971 409 184 that that WDT 10_1101-2020_12_31_424971 409 185 lacked lack VBD 10_1101-2020_12_31_424971 409 186 an an DT 10_1101-2020_12_31_424971 409 187 A A NNP 10_1101-2020_12_31_424971 409 188 site site NN 10_1101-2020_12_31_424971 409 189 - - HYPH 10_1101-2020_12_31_424971 409 190 bound bind VBN 10_1101-2020_12_31_424971 409 191 , , , 10_1101-2020_12_31_424971 409 192 deacylated deacylate VBN 10_1101-2020_12_31_424971 409 193 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 409 194 in in IN 10_1101-2020_12_31_424971 409 195 the the DT 10_1101-2020_12_31_424971 409 196 steady steady JJ 10_1101-2020_12_31_424971 409 197 - - HYPH 10_1101-2020_12_31_424971 409 198 state state NN 10_1101-2020_12_31_424971 409 199 movies movie NNS 10_1101-2020_12_31_424971 409 200 943 943 CD 10_1101-2020_12_31_424971 409 201 recorded record VBN 10_1101-2020_12_31_424971 409 202 for for IN 10_1101-2020_12_31_424971 409 203 the the DT 10_1101-2020_12_31_424971 409 204 longer long JJR 10_1101-2020_12_31_424971 409 205 time time NN 10_1101-2020_12_31_424971 409 206 points point NNS 10_1101-2020_12_31_424971 409 207 ( ( -LRB- 10_1101-2020_12_31_424971 409 208 i.e. i.e. FW 10_1101-2020_12_31_424971 409 209 , , , 10_1101-2020_12_31_424971 409 210 3 3 CD 10_1101-2020_12_31_424971 409 211 , , , 10_1101-2020_12_31_424971 409 212 10 10 CD 10_1101-2020_12_31_424971 409 213 , , , 10_1101-2020_12_31_424971 409 214 and and CC 10_1101-2020_12_31_424971 409 215 20 20 CD 10_1101-2020_12_31_424971 409 216 min min NN 10_1101-2020_12_31_424971 409 217 ) ) -RRB- 10_1101-2020_12_31_424971 409 218 after after IN 10_1101-2020_12_31_424971 409 219 the the DT 10_1101-2020_12_31_424971 409 220 delivery delivery NN 10_1101-2020_12_31_424971 409 221 of of IN 10_1101-2020_12_31_424971 409 222 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 409 223 in in IN 10_1101-2020_12_31_424971 409 224 the the DT 10_1101-2020_12_31_424971 409 225 944 944 CD 10_1101-2020_12_31_424971 409 226 presence presence NN 10_1101-2020_12_31_424971 409 227 of of IN 10_1101-2020_12_31_424971 409 228 EF EF NNP 10_1101-2020_12_31_424971 409 229 - - HYPH 10_1101-2020_12_31_424971 409 230 G G NNP 10_1101-2020_12_31_424971 409 231 ( ( -LRB- 10_1101-2020_12_31_424971 409 232 Figures figure NNS 10_1101-2020_12_31_424971 409 233 6 6 CD 10_1101-2020_12_31_424971 409 234 g g NN 10_1101-2020_12_31_424971 409 235 , , , 10_1101-2020_12_31_424971 409 236 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 409 237 Table Table NNP 10_1101-2020_12_31_424971 409 238 6 6 CD 10_1101-2020_12_31_424971 409 239 ) ) -RRB- 10_1101-2020_12_31_424971 409 240 ; ; : 10_1101-2020_12_31_424971 409 241 ( ( -LRB- 10_1101-2020_12_31_424971 409 242 v v NN 10_1101-2020_12_31_424971 409 243 ) ) -RRB- 10_1101-2020_12_31_424971 409 244 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 409 245 , , , 10_1101-2020_12_31_424971 409 246 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 409 247 , , , 10_1101-2020_12_31_424971 409 248 and and CC 10_1101-2020_12_31_424971 409 249 Keq Keq NNP 10_1101-2020_12_31_424971 409 250 from from IN 10_1101-2020_12_31_424971 409 251 the the DT 10_1101-2020_12_31_424971 409 252 945 945 CD 10_1101-2020_12_31_424971 409 253 steady steady JJ 10_1101-2020_12_31_424971 409 254 - - HYPH 10_1101-2020_12_31_424971 409 255 state state NN 10_1101-2020_12_31_424971 409 256 movies movie NNS 10_1101-2020_12_31_424971 409 257 recorded record VBN 10_1101-2020_12_31_424971 409 258 for for IN 10_1101-2020_12_31_424971 409 259 the the DT 10_1101-2020_12_31_424971 409 260 SufB2- sufb2- NN 10_1101-2020_12_31_424971 409 261 and and CC 10_1101-2020_12_31_424971 409 262 ProL ProL NNP 10_1101-2020_12_31_424971 409 263 PRE PRE NNP 10_1101-2020_12_31_424971 409 264 – – : 10_1101-2020_12_31_424971 409 265 A a DT 10_1101-2020_12_31_424971 409 266 complexes complex NNS 10_1101-2020_12_31_424971 409 267 ( ( -LRB- 10_1101-2020_12_31_424971 409 268 Figures figure NNS 10_1101-2020_12_31_424971 409 269 6h 6h CD 10_1101-2020_12_31_424971 409 270 and and CC 10_1101-2020_12_31_424971 409 271 946 946 CD 10_1101-2020_12_31_424971 409 272 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 409 273 Figure Figure NNP 10_1101-2020_12_31_424971 409 274 6 6 CD 10_1101-2020_12_31_424971 409 275 and and CC 10_1101-2020_12_31_424971 409 276 Table table NN 10_1101-2020_12_31_424971 409 277 7 7 CD 10_1101-2020_12_31_424971 409 278 ) ) -RRB- 10_1101-2020_12_31_424971 409 279 ; ; : 10_1101-2020_12_31_424971 409 280 and and CC 10_1101-2020_12_31_424971 409 281 ( ( -LRB- 10_1101-2020_12_31_424971 409 282 vi vi NNP 10_1101-2020_12_31_424971 409 283 ) ) -RRB- 10_1101-2020_12_31_424971 409 284 kGS1→Æ kGS1→Æ NNP 10_1101-2020_12_31_424971 409 285 , , , 10_1101-2020_12_31_424971 409 286 kGS2→Æ kGS2→Æ NNP 10_1101-2020_12_31_424971 409 287 , , , 10_1101-2020_12_31_424971 409 288 kGS2(G)→Æ kGS2(G)→Æ NNP 10_1101-2020_12_31_424971 409 289 , , , 10_1101-2020_12_31_424971 409 290 and and CC 10_1101-2020_12_31_424971 409 291 kPOST→Æ kpost→æ NN 10_1101-2020_12_31_424971 409 292 from from IN 10_1101-2020_12_31_424971 409 293 the the DT 10_1101-2020_12_31_424971 409 294 947 947 CD 10_1101-2020_12_31_424971 409 295 movies movie NNS 10_1101-2020_12_31_424971 409 296 described describe VBN 10_1101-2020_12_31_424971 409 297 in in IN 10_1101-2020_12_31_424971 409 298 ( ( -LRB- 10_1101-2020_12_31_424971 409 299 i)-(v i)-(v NN 10_1101-2020_12_31_424971 409 300 ) ) -RRB- 10_1101-2020_12_31_424971 409 301 ( ( -LRB- 10_1101-2020_12_31_424971 409 302 Figures figure NNS 10_1101-2020_12_31_424971 409 303 6d-6h 6d-6h CD 10_1101-2020_12_31_424971 409 304 , , , 10_1101-2020_12_31_424971 409 305 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 409 306 Figures Figures NNP 10_1101-2020_12_31_424971 409 307 3 3 CD 10_1101-2020_12_31_424971 409 308 , , , 10_1101-2020_12_31_424971 409 309 5 5 CD 10_1101-2020_12_31_424971 409 310 , , , 10_1101-2020_12_31_424971 409 311 and and CC 10_1101-2020_12_31_424971 409 312 6 6 CD 10_1101-2020_12_31_424971 409 313 , , , 10_1101-2020_12_31_424971 409 314 and and CC 10_1101-2020_12_31_424971 409 315 reported report VBD 10_1101-2020_12_31_424971 409 316 two two CD 10_1101-2020_12_31_424971 409 317 948 948 CD 10_1101-2020_12_31_424971 409 318 paragraphs paragraph NNS 10_1101-2020_12_31_424971 409 319 above above RB 10_1101-2020_12_31_424971 409 320 ) ) -RRB- 10_1101-2020_12_31_424971 409 321 . . . 10_1101-2020_12_31_424971 410 1 To to TO 10_1101-2020_12_31_424971 410 2 calculate calculate VB 10_1101-2020_12_31_424971 410 3 these these DT 10_1101-2020_12_31_424971 410 4 parameters parameter NNS 10_1101-2020_12_31_424971 410 5 , , , 10_1101-2020_12_31_424971 410 6 we -PRON- PRP 10_1101-2020_12_31_424971 410 7 extended extend VBD 10_1101-2020_12_31_424971 410 8 the the DT 10_1101-2020_12_31_424971 410 9 variational variational JJ 10_1101-2020_12_31_424971 410 10 Bayes Bayes NNP 10_1101-2020_12_31_424971 410 11 approach approach VBP 10_1101-2020_12_31_424971 410 12 949 949 CD 10_1101-2020_12_31_424971 410 13 we -PRON- PRP 10_1101-2020_12_31_424971 410 14 introduced introduce VBD 10_1101-2020_12_31_424971 410 15 in in IN 10_1101-2020_12_31_424971 410 16 the the DT 10_1101-2020_12_31_424971 410 17 vbFRET vbFRET NNP 10_1101-2020_12_31_424971 410 18 algorithm68 algorithm68 NNP 10_1101-2020_12_31_424971 410 19 to to TO 10_1101-2020_12_31_424971 410 20 estimate estimate VB 10_1101-2020_12_31_424971 410 21 a a DT 10_1101-2020_12_31_424971 410 22 ‘ ' `` 10_1101-2020_12_31_424971 410 23 consensus consensus NN 10_1101-2020_12_31_424971 410 24 ’ ' '' 10_1101-2020_12_31_424971 410 25 ( ( -LRB- 10_1101-2020_12_31_424971 410 26 i.e. i.e. FW 10_1101-2020_12_31_424971 410 27 , , , 10_1101-2020_12_31_424971 410 28 ‘ ' `` 10_1101-2020_12_31_424971 410 29 global global JJ 10_1101-2020_12_31_424971 410 30 ’ ' '' 10_1101-2020_12_31_424971 410 31 ) ) -RRB- 10_1101-2020_12_31_424971 410 32 HMM HMM NNP 10_1101-2020_12_31_424971 410 33 of of IN 10_1101-2020_12_31_424971 410 34 the the DT 10_1101-2020_12_31_424971 410 35 950 950 CD 10_1101-2020_12_31_424971 410 36 EFRET EFRET NNP 10_1101-2020_12_31_424971 410 37 vs. vs. NN 10_1101-2020_12_31_424971 410 38 time time NN 10_1101-2020_12_31_424971 410 39 trajectories trajectory NNS 10_1101-2020_12_31_424971 410 40 . . . 10_1101-2020_12_31_424971 411 1 In in IN 10_1101-2020_12_31_424971 411 2 this this DT 10_1101-2020_12_31_424971 411 3 approach approach NN 10_1101-2020_12_31_424971 411 4 , , , 10_1101-2020_12_31_424971 411 5 we -PRON- PRP 10_1101-2020_12_31_424971 411 6 use use VBP 10_1101-2020_12_31_424971 411 7 Bayesian bayesian JJ 10_1101-2020_12_31_424971 411 8 inference inference NN 10_1101-2020_12_31_424971 411 9 to to TO 10_1101-2020_12_31_424971 411 10 estimate estimate VB 10_1101-2020_12_31_424971 411 11 a a DT 10_1101-2020_12_31_424971 411 12 single single JJ 10_1101-2020_12_31_424971 411 13 , , , 10_1101-2020_12_31_424971 411 14 951 951 CD 10_1101-2020_12_31_424971 411 15 consensus consensus NN 10_1101-2020_12_31_424971 411 16 HMM HMM NNP 10_1101-2020_12_31_424971 411 17 that that WDT 10_1101-2020_12_31_424971 411 18 is be VBZ 10_1101-2020_12_31_424971 411 19 most most RBS 10_1101-2020_12_31_424971 411 20 consistent consistent JJ 10_1101-2020_12_31_424971 411 21 with with IN 10_1101-2020_12_31_424971 411 22 all all PDT 10_1101-2020_12_31_424971 411 23 the the DT 10_1101-2020_12_31_424971 411 24 EFRET EFRET NNP 10_1101-2020_12_31_424971 411 25 vs. vs. NN 10_1101-2020_12_31_424971 411 26 time time NN 10_1101-2020_12_31_424971 411 27 trajectories trajectory NNS 10_1101-2020_12_31_424971 411 28 in in IN 10_1101-2020_12_31_424971 411 29 a a DT 10_1101-2020_12_31_424971 411 30 movie movie NN 10_1101-2020_12_31_424971 411 31 , , , 10_1101-2020_12_31_424971 411 32 rather rather RB 10_1101-2020_12_31_424971 411 33 952 952 CD 10_1101-2020_12_31_424971 411 34 than than IN 10_1101-2020_12_31_424971 411 35 to to TO 10_1101-2020_12_31_424971 411 36 estimate estimate VB 10_1101-2020_12_31_424971 411 37 a a DT 10_1101-2020_12_31_424971 411 38 separate separate JJ 10_1101-2020_12_31_424971 411 39 HMM HMM NNP 10_1101-2020_12_31_424971 411 40 for for IN 10_1101-2020_12_31_424971 411 41 each each DT 10_1101-2020_12_31_424971 411 42 trajectory trajectory NN 10_1101-2020_12_31_424971 411 43 in in IN 10_1101-2020_12_31_424971 411 44 the the DT 10_1101-2020_12_31_424971 411 45 movie movie NN 10_1101-2020_12_31_424971 411 46 . . . 10_1101-2020_12_31_424971 412 1 To to TO 10_1101-2020_12_31_424971 412 2 estimate estimate VB 10_1101-2020_12_31_424971 412 3 such such PDT 10_1101-2020_12_31_424971 412 4 a a DT 10_1101-2020_12_31_424971 412 5 consensus consensus NN 10_1101-2020_12_31_424971 412 6 953 953 CD 10_1101-2020_12_31_424971 412 7 HMM HMM NNP 10_1101-2020_12_31_424971 412 8 , , , 10_1101-2020_12_31_424971 412 9 we -PRON- PRP 10_1101-2020_12_31_424971 412 10 assume assume VBP 10_1101-2020_12_31_424971 412 11 each each DT 10_1101-2020_12_31_424971 412 12 trajectory trajectory CD 10_1101-2020_12_31_424971 412 13 is be VBZ 10_1101-2020_12_31_424971 412 14 independent independent JJ 10_1101-2020_12_31_424971 412 15 and and CC 10_1101-2020_12_31_424971 412 16 identically identically RB 10_1101-2020_12_31_424971 412 17 distributed distribute VBN 10_1101-2020_12_31_424971 412 18 , , , 10_1101-2020_12_31_424971 412 19 thereby thereby RB 10_1101-2020_12_31_424971 412 20 enabling enable VBG 10_1101-2020_12_31_424971 412 21 us -PRON- PRP 10_1101-2020_12_31_424971 412 22 954 954 CD 10_1101-2020_12_31_424971 412 23 to to TO 10_1101-2020_12_31_424971 412 24 perform perform VB 10_1101-2020_12_31_424971 412 25 the the DT 10_1101-2020_12_31_424971 412 26 inference inference NN 10_1101-2020_12_31_424971 412 27 using use VBG 10_1101-2020_12_31_424971 412 28 the the DT 10_1101-2020_12_31_424971 412 29 likelihood likelihood NN 10_1101-2020_12_31_424971 412 30 function function NN 10_1101-2020_12_31_424971 412 31 955 955 CD 10_1101-2020_12_31_424971 412 32 ℒ ℒ NNP 10_1101-2020_12_31_424971 412 33 = = SYM 10_1101-2020_12_31_424971 412 34 ∏ ∏ NNP 10_1101-2020_12_31_424971 412 35 ℒ ℒ NNP 10_1101-2020_12_31_424971 412 36 & & CC 10_1101-2020_12_31_424971 412 37 & & CC 10_1101-2020_12_31_424971 412 38 ∈ ∈ NNP 10_1101-2020_12_31_424971 412 39 ) ) -RRB- 10_1101-2020_12_31_424971 412 40 * * NFP 10_1101-2020_12_31_424971 412 41 + + SYM 10_1101-2020_12_31_424971 412 42 , , , 10_1101-2020_12_31_424971 412 43 - - : 10_1101-2020_12_31_424971 412 44 . . . 10_1101-2020_12_31_424971 413 1 ) ) -RRB- 10_1101-2020_12_31_424971 413 2 /*0 /*0 NFP 10_1101-2020_12_31_424971 413 3 - - SYM 10_1101-2020_12_31_424971 413 4 1 1 CD 10_1101-2020_12_31_424971 413 5 , , , 10_1101-2020_12_31_424971 413 6 ( ( -LRB- 10_1101-2020_12_31_424971 413 7 2 2 LS 10_1101-2020_12_31_424971 413 8 ) ) -RRB- 10_1101-2020_12_31_424971 413 9 956 956 CD 10_1101-2020_12_31_424971 413 10 .CC .CC : 10_1101-2020_12_31_424971 413 11 - - HYPH 10_1101-2020_12_31_424971 413 12 BY by IN 10_1101-2020_12_31_424971 413 13 - - HYPH 10_1101-2020_12_31_424971 413 14 NC NC NNP 10_1101-2020_12_31_424971 413 15 - - HYPH 10_1101-2020_12_31_424971 413 16 ND ND NNP 10_1101-2020_12_31_424971 413 17 4.0 4.0 CD 10_1101-2020_12_31_424971 413 18 International International NNP 10_1101-2020_12_31_424971 413 19 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 413 20 under under IN 10_1101-2020_12_31_424971 413 21 a a DT 10_1101-2020_12_31_424971 413 22 ( ( -LRB- 10_1101-2020_12_31_424971 413 23 which which WDT 10_1101-2020_12_31_424971 413 24 was be VBD 10_1101-2020_12_31_424971 413 25 not not RB 10_1101-2020_12_31_424971 413 26 certified certify VBN 10_1101-2020_12_31_424971 413 27 by by IN 10_1101-2020_12_31_424971 413 28 peer peer NN 10_1101-2020_12_31_424971 413 29 review review NN 10_1101-2020_12_31_424971 413 30 ) ) -RRB- 10_1101-2020_12_31_424971 413 31 is be VBZ 10_1101-2020_12_31_424971 413 32 the the DT 10_1101-2020_12_31_424971 413 33 author author NN 10_1101-2020_12_31_424971 413 34 / / SYM 10_1101-2020_12_31_424971 413 35 funder funder NN 10_1101-2020_12_31_424971 413 36 , , , 10_1101-2020_12_31_424971 413 37 who who WP 10_1101-2020_12_31_424971 413 38 has have VBZ 10_1101-2020_12_31_424971 413 39 granted grant VBN 10_1101-2020_12_31_424971 413 40 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 413 41 a a DT 10_1101-2020_12_31_424971 413 42 license license NN 10_1101-2020_12_31_424971 413 43 to to TO 10_1101-2020_12_31_424971 413 44 display display VB 10_1101-2020_12_31_424971 413 45 the the DT 10_1101-2020_12_31_424971 413 46 preprint preprint NN 10_1101-2020_12_31_424971 413 47 in in IN 10_1101-2020_12_31_424971 413 48 perpetuity perpetuity NN 10_1101-2020_12_31_424971 413 49 . . . 10_1101-2020_12_31_424971 414 1 It -PRON- PRP 10_1101-2020_12_31_424971 414 2 is be VBZ 10_1101-2020_12_31_424971 414 3 made make VBN 10_1101-2020_12_31_424971 414 4 The the DT 10_1101-2020_12_31_424971 414 5 copyright copyright NN 10_1101-2020_12_31_424971 414 6 holder holder NN 10_1101-2020_12_31_424971 414 7 for for IN 10_1101-2020_12_31_424971 414 8 this this DT 10_1101-2020_12_31_424971 414 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 414 10 version version NN 10_1101-2020_12_31_424971 414 11 posted post VBD 10_1101-2020_12_31_424971 414 12 January January NNP 10_1101-2020_12_31_424971 414 13 2 2 CD 10_1101-2020_12_31_424971 414 14 , , , 10_1101-2020_12_31_424971 414 15 2021 2021 CD 10_1101-2020_12_31_424971 414 16 . . . 10_1101-2020_12_31_424971 414 17 ; ; : 10_1101-2020_12_31_424971 414 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 414 19 : : : 10_1101-2020_12_31_424971 414 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 414 21 preprint preprint NN 10_1101-2020_12_31_424971 414 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 414 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 414 24 39 39 CD 10_1101-2020_12_31_424971 414 25 where where WRB 10_1101-2020_12_31_424971 414 26 ℒ ℒ NNP 10_1101-2020_12_31_424971 414 27 & & CC 10_1101-2020_12_31_424971 414 28 is be VBZ 10_1101-2020_12_31_424971 414 29 the the DT 10_1101-2020_12_31_424971 414 30 variational variational JJ 10_1101-2020_12_31_424971 414 31 approximation approximation NN 10_1101-2020_12_31_424971 414 32 of of IN 10_1101-2020_12_31_424971 414 33 the the DT 10_1101-2020_12_31_424971 414 34 likelihood likelihood NN 10_1101-2020_12_31_424971 414 35 function function NN 10_1101-2020_12_31_424971 414 36 for for IN 10_1101-2020_12_31_424971 414 37 a a DT 10_1101-2020_12_31_424971 414 38 single single JJ 10_1101-2020_12_31_424971 414 39 trajectory trajectory NN 10_1101-2020_12_31_424971 414 40 . . . 10_1101-2020_12_31_424971 415 1 957 957 CD 10_1101-2020_12_31_424971 415 2 Subsequently subsequently RB 10_1101-2020_12_31_424971 415 3 , , , 10_1101-2020_12_31_424971 415 4 the the DT 10_1101-2020_12_31_424971 415 5 single single JJ 10_1101-2020_12_31_424971 415 6 , , , 10_1101-2020_12_31_424971 415 7 consensus consensus NN 10_1101-2020_12_31_424971 415 8 HMM HMM NNP 10_1101-2020_12_31_424971 415 9 that that WDT 10_1101-2020_12_31_424971 415 10 is be VBZ 10_1101-2020_12_31_424971 415 11 most most RBS 10_1101-2020_12_31_424971 415 12 consistent consistent JJ 10_1101-2020_12_31_424971 415 13 with with IN 10_1101-2020_12_31_424971 415 14 all all DT 10_1101-2020_12_31_424971 415 15 of of IN 10_1101-2020_12_31_424971 415 16 the the DT 10_1101-2020_12_31_424971 415 17 trajectories trajectory NNS 10_1101-2020_12_31_424971 415 18 is be VBZ 10_1101-2020_12_31_424971 415 19 958 958 CD 10_1101-2020_12_31_424971 415 20 estimated estimate VBN 10_1101-2020_12_31_424971 415 21 using use VBG 10_1101-2020_12_31_424971 415 22 the the DT 10_1101-2020_12_31_424971 415 23 expectation expectation NN 10_1101-2020_12_31_424971 415 24 - - HYPH 10_1101-2020_12_31_424971 415 25 maximization maximization NN 10_1101-2020_12_31_424971 415 26 algorithm algorithm NN 10_1101-2020_12_31_424971 415 27 that that IN 10_1101-2020_12_31_424971 415 28 we -PRON- PRP 10_1101-2020_12_31_424971 415 29 have have VBP 10_1101-2020_12_31_424971 415 30 previously previously RB 10_1101-2020_12_31_424971 415 31 described68 described68 NFP 10_1101-2020_12_31_424971 415 32 . . . 10_1101-2020_12_31_424971 416 1 959 959 CD 10_1101-2020_12_31_424971 416 2 Viterbi Viterbi NNP 10_1101-2020_12_31_424971 416 3 paths path NNS 10_1101-2020_12_31_424971 416 4 ( ( -LRB- 10_1101-2020_12_31_424971 416 5 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 416 6 Figures Figures NNPS 10_1101-2020_12_31_424971 416 7 3 3 CD 10_1101-2020_12_31_424971 416 8 , , , 10_1101-2020_12_31_424971 416 9 5 5 CD 10_1101-2020_12_31_424971 416 10 , , , 10_1101-2020_12_31_424971 416 11 and and CC 10_1101-2020_12_31_424971 416 12 6 6 CD 10_1101-2020_12_31_424971 416 13 ) ) -RRB- 10_1101-2020_12_31_424971 416 14 , , , 10_1101-2020_12_31_424971 416 15 representing represent VBG 10_1101-2020_12_31_424971 416 16 the the DT 10_1101-2020_12_31_424971 416 17 most most RBS 10_1101-2020_12_31_424971 416 18 probable probable JJ 10_1101-2020_12_31_424971 416 19 hidden hide VBN 10_1101-2020_12_31_424971 416 20 - - HYPH 10_1101-2020_12_31_424971 416 21 state state NN 10_1101-2020_12_31_424971 416 22 960 960 CD 10_1101-2020_12_31_424971 416 23 trajectory trajectory CD 10_1101-2020_12_31_424971 416 24 , , , 10_1101-2020_12_31_424971 416 25 were be VBD 10_1101-2020_12_31_424971 416 26 then then RB 10_1101-2020_12_31_424971 416 27 calculated calculate VBN 10_1101-2020_12_31_424971 416 28 from from IN 10_1101-2020_12_31_424971 416 29 the the DT 10_1101-2020_12_31_424971 416 30 HMM HMM NNP 10_1101-2020_12_31_424971 416 31 using use VBG 10_1101-2020_12_31_424971 416 32 the the DT 10_1101-2020_12_31_424971 416 33 Viterbi Viterbi NNP 10_1101-2020_12_31_424971 416 34 algorithm69 algorithm69 NNP 10_1101-2020_12_31_424971 416 35 . . . 10_1101-2020_12_31_424971 417 1 Based base VBN 10_1101-2020_12_31_424971 417 2 on on IN 10_1101-2020_12_31_424971 417 3 extensive extensive JJ 10_1101-2020_12_31_424971 417 4 961 961 CD 10_1101-2020_12_31_424971 417 5 smFRET smFRET NNP 10_1101-2020_12_31_424971 417 6 studies study NNS 10_1101-2020_12_31_424971 417 7 of of IN 10_1101-2020_12_31_424971 417 8 translation translation NN 10_1101-2020_12_31_424971 417 9 elongation elongation NN 10_1101-2020_12_31_424971 417 10 using use VBG 10_1101-2020_12_31_424971 417 11 the the DT 10_1101-2020_12_31_424971 417 12 bL9(Cy3)-uL1(Cy5 bl9(cy3)-ul1(cy5 NN 10_1101-2020_12_31_424971 417 13 ) ) -RRB- 10_1101-2020_12_31_424971 417 14 smFRET smfret $ 10_1101-2020_12_31_424971 417 15 signal35,36,38 signal35,36,38 CD 10_1101-2020_12_31_424971 417 16 , , , 10_1101-2020_12_31_424971 417 17 we -PRON- PRP 10_1101-2020_12_31_424971 417 18 962 962 CD 10_1101-2020_12_31_424971 417 19 selected select VBD 10_1101-2020_12_31_424971 417 20 a a DT 10_1101-2020_12_31_424971 417 21 consensus consensus NN 10_1101-2020_12_31_424971 417 22 HMM HMM NNP 10_1101-2020_12_31_424971 417 23 composed compose VBN 10_1101-2020_12_31_424971 417 24 of of IN 10_1101-2020_12_31_424971 417 25 three three CD 10_1101-2020_12_31_424971 417 26 states state NNS 10_1101-2020_12_31_424971 417 27 for for IN 10_1101-2020_12_31_424971 417 28 further further JJ 10_1101-2020_12_31_424971 417 29 analysis analysis NN 10_1101-2020_12_31_424971 417 30 of of IN 10_1101-2020_12_31_424971 417 31 the the DT 10_1101-2020_12_31_424971 417 32 data datum NNS 10_1101-2020_12_31_424971 417 33 . . . 10_1101-2020_12_31_424971 418 1 For for IN 10_1101-2020_12_31_424971 418 2 963 963 CD 10_1101-2020_12_31_424971 418 3 calculation calculation NN 10_1101-2020_12_31_424971 418 4 of of IN 10_1101-2020_12_31_424971 418 5 the the DT 10_1101-2020_12_31_424971 418 6 kinetic kinetic JJ 10_1101-2020_12_31_424971 418 7 and/or and/or CC 10_1101-2020_12_31_424971 418 8 thermodynamic thermodynamic JJ 10_1101-2020_12_31_424971 418 9 parameters parameter NNS 10_1101-2020_12_31_424971 418 10 in in IN 10_1101-2020_12_31_424971 418 11 ( ( -LRB- 10_1101-2020_12_31_424971 418 12 i i NN 10_1101-2020_12_31_424971 418 13 ) ) -RRB- 10_1101-2020_12_31_424971 418 14 , , , 10_1101-2020_12_31_424971 418 15 ( ( -LRB- 10_1101-2020_12_31_424971 418 16 iv iv LS 10_1101-2020_12_31_424971 418 17 ) ) -RRB- 10_1101-2020_12_31_424971 418 18 , , , 10_1101-2020_12_31_424971 418 19 and and CC 10_1101-2020_12_31_424971 418 20 ( ( -LRB- 10_1101-2020_12_31_424971 418 21 v v NN 10_1101-2020_12_31_424971 418 22 ) ) -RRB- 10_1101-2020_12_31_424971 418 23 , , , 10_1101-2020_12_31_424971 418 24 the the DT 10_1101-2020_12_31_424971 418 25 three three CD 10_1101-2020_12_31_424971 418 26 states state NNS 10_1101-2020_12_31_424971 418 27 964 964 CD 10_1101-2020_12_31_424971 418 28 corresponded correspond VBD 10_1101-2020_12_31_424971 418 29 to to IN 10_1101-2020_12_31_424971 418 30 GS1 GS1 NNP 10_1101-2020_12_31_424971 418 31 , , , 10_1101-2020_12_31_424971 418 32 GS2 GS2 NNP 10_1101-2020_12_31_424971 418 33 , , , 10_1101-2020_12_31_424971 418 34 and and CC 10_1101-2020_12_31_424971 418 35 Æ Æ NNP 10_1101-2020_12_31_424971 418 36 and and CC 10_1101-2020_12_31_424971 418 37 for for IN 10_1101-2020_12_31_424971 418 38 calculation calculation NN 10_1101-2020_12_31_424971 418 39 of of IN 10_1101-2020_12_31_424971 418 40 the the DT 10_1101-2020_12_31_424971 418 41 kinetic kinetic NN 10_1101-2020_12_31_424971 418 42 and/or and/or CC 10_1101-2020_12_31_424971 418 43 thermodynamic thermodynamic NN 10_1101-2020_12_31_424971 418 44 965 965 CD 10_1101-2020_12_31_424971 418 45 parameters parameter NNS 10_1101-2020_12_31_424971 418 46 in in IN 10_1101-2020_12_31_424971 418 47 ( ( -LRB- 10_1101-2020_12_31_424971 418 48 ii ii NNP 10_1101-2020_12_31_424971 418 49 ) ) -RRB- 10_1101-2020_12_31_424971 418 50 and and CC 10_1101-2020_12_31_424971 418 51 ( ( -LRB- 10_1101-2020_12_31_424971 418 52 iii iii NNP 10_1101-2020_12_31_424971 418 53 ) ) -RRB- 10_1101-2020_12_31_424971 418 54 , , , 10_1101-2020_12_31_424971 418 55 the the DT 10_1101-2020_12_31_424971 418 56 three three CD 10_1101-2020_12_31_424971 418 57 states state NNS 10_1101-2020_12_31_424971 418 58 corresponded correspond VBD 10_1101-2020_12_31_424971 418 59 to to IN 10_1101-2020_12_31_424971 418 60 EF EF NNP 10_1101-2020_12_31_424971 418 61 - - HYPH 10_1101-2020_12_31_424971 418 62 G G NNP 10_1101-2020_12_31_424971 418 63 - - HYPH 10_1101-2020_12_31_424971 418 64 bound bind VBN 10_1101-2020_12_31_424971 418 65 GS2-like GS2-like NNP 10_1101-2020_12_31_424971 418 66 , , , 10_1101-2020_12_31_424971 418 67 POST POST NNP 10_1101-2020_12_31_424971 418 68 , , , 10_1101-2020_12_31_424971 418 69 and and CC 10_1101-2020_12_31_424971 418 70 Æ. æ. NN 10_1101-2020_12_31_424971 419 1 966 966 CD 10_1101-2020_12_31_424971 419 2 The the DT 10_1101-2020_12_31_424971 419 3 transition transition NN 10_1101-2020_12_31_424971 419 4 matrix matrix NN 10_1101-2020_12_31_424971 419 5 of of IN 10_1101-2020_12_31_424971 419 6 the the DT 10_1101-2020_12_31_424971 419 7 consensus consensus NN 10_1101-2020_12_31_424971 419 8 HMM HMM NNP 10_1101-2020_12_31_424971 419 9 was be VBD 10_1101-2020_12_31_424971 419 10 then then RB 10_1101-2020_12_31_424971 419 11 used use VBN 10_1101-2020_12_31_424971 419 12 to to TO 10_1101-2020_12_31_424971 419 13 calculate calculate VB 10_1101-2020_12_31_424971 419 14 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 419 15 and and CC 10_1101-2020_12_31_424971 419 16 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 419 17 in in IN 10_1101-2020_12_31_424971 419 18 967 967 CD 10_1101-2020_12_31_424971 419 19 ( ( -LRB- 10_1101-2020_12_31_424971 419 20 i i NN 10_1101-2020_12_31_424971 419 21 ) ) -RRB- 10_1101-2020_12_31_424971 419 22 , , , 10_1101-2020_12_31_424971 419 23 ( ( -LRB- 10_1101-2020_12_31_424971 419 24 iv iv LS 10_1101-2020_12_31_424971 419 25 ) ) -RRB- 10_1101-2020_12_31_424971 419 26 , , , 10_1101-2020_12_31_424971 419 27 and and CC 10_1101-2020_12_31_424971 419 28 ( ( -LRB- 10_1101-2020_12_31_424971 419 29 v v NN 10_1101-2020_12_31_424971 419 30 ) ) -RRB- 10_1101-2020_12_31_424971 419 31 ; ; : 10_1101-2020_12_31_424971 419 32 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 419 33 in in IN 10_1101-2020_12_31_424971 419 34 ( ( -LRB- 10_1101-2020_12_31_424971 419 35 ii ii NNP 10_1101-2020_12_31_424971 419 36 ) ) -RRB- 10_1101-2020_12_31_424971 419 37 ; ; : 10_1101-2020_12_31_424971 419 38 kGS1→Æ kGS1→Æ NNP 10_1101-2020_12_31_424971 419 39 , , , 10_1101-2020_12_31_424971 419 40 kGS2→Æ kGS2→Æ NNP 10_1101-2020_12_31_424971 419 41 , , , 10_1101-2020_12_31_424971 419 42 kGS2(G)→Æ kGS2(G)→Æ NNP 10_1101-2020_12_31_424971 419 43 , , , 10_1101-2020_12_31_424971 419 44 and and CC 10_1101-2020_12_31_424971 419 45 kPOST→Æ kpost→æ NN 10_1101-2020_12_31_424971 419 46 in in RP 10_1101-2020_12_31_424971 419 47 ( ( -LRB- 10_1101-2020_12_31_424971 419 48 vi vi NNP 10_1101-2020_12_31_424971 419 49 ) ) -RRB- 10_1101-2020_12_31_424971 419 50 ; ; : 10_1101-2020_12_31_424971 419 51 and and CC 10_1101-2020_12_31_424971 419 52 the the DT 10_1101-2020_12_31_424971 419 53 errors error NNS 10_1101-2020_12_31_424971 419 54 968 968 CD 10_1101-2020_12_31_424971 419 55 corresponding correspond VBG 10_1101-2020_12_31_424971 419 56 to to IN 10_1101-2020_12_31_424971 419 57 each each DT 10_1101-2020_12_31_424971 419 58 of of IN 10_1101-2020_12_31_424971 419 59 these these DT 10_1101-2020_12_31_424971 419 60 parameters parameter NNS 10_1101-2020_12_31_424971 419 61 . . . 10_1101-2020_12_31_424971 420 1 This this DT 10_1101-2020_12_31_424971 420 2 transition transition NN 10_1101-2020_12_31_424971 420 3 matrix matrix NN 10_1101-2020_12_31_424971 420 4 consists consist VBZ 10_1101-2020_12_31_424971 420 5 of of IN 10_1101-2020_12_31_424971 420 6 a a DT 10_1101-2020_12_31_424971 420 7 3 3 CD 10_1101-2020_12_31_424971 420 8 x x SYM 10_1101-2020_12_31_424971 420 9 3 3 CD 10_1101-2020_12_31_424971 420 10 matrix matrix NN 10_1101-2020_12_31_424971 420 11 in in IN 10_1101-2020_12_31_424971 420 12 969 969 CD 10_1101-2020_12_31_424971 420 13 which which WDT 10_1101-2020_12_31_424971 420 14 the the DT 10_1101-2020_12_31_424971 420 15 off off JJ 10_1101-2020_12_31_424971 420 16 - - HYPH 10_1101-2020_12_31_424971 420 17 diagonal diagonal JJ 10_1101-2020_12_31_424971 420 18 elements element NNS 10_1101-2020_12_31_424971 420 19 correspond correspond VBP 10_1101-2020_12_31_424971 420 20 to to IN 10_1101-2020_12_31_424971 420 21 the the DT 10_1101-2020_12_31_424971 420 22 number number NN 10_1101-2020_12_31_424971 420 23 of of IN 10_1101-2020_12_31_424971 420 24 times time NNS 10_1101-2020_12_31_424971 420 25 a a DT 10_1101-2020_12_31_424971 420 26 transition transition NN 10_1101-2020_12_31_424971 420 27 takes take VBZ 10_1101-2020_12_31_424971 420 28 place place NN 10_1101-2020_12_31_424971 420 29 970 970 CD 10_1101-2020_12_31_424971 420 30 between between IN 10_1101-2020_12_31_424971 420 31 each each DT 10_1101-2020_12_31_424971 420 32 pair pair NN 10_1101-2020_12_31_424971 420 33 of of IN 10_1101-2020_12_31_424971 420 34 the the DT 10_1101-2020_12_31_424971 420 35 GS1 GS1 NNP 10_1101-2020_12_31_424971 420 36 , , , 10_1101-2020_12_31_424971 420 37 GS2 GS2 NNP 10_1101-2020_12_31_424971 420 38 , , , 10_1101-2020_12_31_424971 420 39 and and CC 10_1101-2020_12_31_424971 420 40 Æ æ NN 10_1101-2020_12_31_424971 420 41 states state NNS 10_1101-2020_12_31_424971 420 42 ( ( -LRB- 10_1101-2020_12_31_424971 420 43 in in IN 10_1101-2020_12_31_424971 420 44 ( ( -LRB- 10_1101-2020_12_31_424971 420 45 i i NN 10_1101-2020_12_31_424971 420 46 ) ) -RRB- 10_1101-2020_12_31_424971 420 47 , , , 10_1101-2020_12_31_424971 420 48 ( ( -LRB- 10_1101-2020_12_31_424971 420 49 iv iv NN 10_1101-2020_12_31_424971 420 50 ) ) -RRB- 10_1101-2020_12_31_424971 420 51 , , , 10_1101-2020_12_31_424971 420 52 ( ( -LRB- 10_1101-2020_12_31_424971 420 53 v v NN 10_1101-2020_12_31_424971 420 54 ) ) -RRB- 10_1101-2020_12_31_424971 420 55 , , , 10_1101-2020_12_31_424971 420 56 and and CC 10_1101-2020_12_31_424971 420 57 ( ( -LRB- 10_1101-2020_12_31_424971 420 58 vi vi NNP 10_1101-2020_12_31_424971 420 59 ) ) -RRB- 10_1101-2020_12_31_424971 420 60 ) ) -RRB- 10_1101-2020_12_31_424971 420 61 or or CC 10_1101-2020_12_31_424971 420 62 each each DT 10_1101-2020_12_31_424971 420 63 pair pair NN 10_1101-2020_12_31_424971 420 64 of of IN 10_1101-2020_12_31_424971 420 65 the the DT 10_1101-2020_12_31_424971 420 66 EF-971 EF-971 NNP 10_1101-2020_12_31_424971 420 67 G G NNP 10_1101-2020_12_31_424971 420 68 - - HYPH 10_1101-2020_12_31_424971 420 69 bound bind VBN 10_1101-2020_12_31_424971 420 70 GS2-like GS2-like NNP 10_1101-2020_12_31_424971 420 71 , , , 10_1101-2020_12_31_424971 420 72 POST POST NNP 10_1101-2020_12_31_424971 420 73 , , , 10_1101-2020_12_31_424971 420 74 and and CC 10_1101-2020_12_31_424971 420 75 Æ Æ NNP 10_1101-2020_12_31_424971 420 76 states state NNS 10_1101-2020_12_31_424971 420 77 ( ( -LRB- 10_1101-2020_12_31_424971 420 78 in in IN 10_1101-2020_12_31_424971 420 79 ( ( -LRB- 10_1101-2020_12_31_424971 420 80 ii ii NNP 10_1101-2020_12_31_424971 420 81 ) ) -RRB- 10_1101-2020_12_31_424971 420 82 and and CC 10_1101-2020_12_31_424971 420 83 ( ( -LRB- 10_1101-2020_12_31_424971 420 84 vi vi NNP 10_1101-2020_12_31_424971 420 85 ) ) -RRB- 10_1101-2020_12_31_424971 420 86 ) ) -RRB- 10_1101-2020_12_31_424971 420 87 and and CC 10_1101-2020_12_31_424971 420 88 the the DT 10_1101-2020_12_31_424971 420 89 on on IN 10_1101-2020_12_31_424971 420 90 - - HYPH 10_1101-2020_12_31_424971 420 91 diagonal diagonal JJ 10_1101-2020_12_31_424971 420 92 elements element NNS 10_1101-2020_12_31_424971 420 93 correspond correspond VBP 10_1101-2020_12_31_424971 420 94 972 972 CD 10_1101-2020_12_31_424971 420 95 to to IN 10_1101-2020_12_31_424971 420 96 the the DT 10_1101-2020_12_31_424971 420 97 number number NN 10_1101-2020_12_31_424971 420 98 of of IN 10_1101-2020_12_31_424971 420 99 times time NNS 10_1101-2020_12_31_424971 420 100 a a DT 10_1101-2020_12_31_424971 420 101 transition transition NN 10_1101-2020_12_31_424971 420 102 does do VBZ 10_1101-2020_12_31_424971 420 103 not not RB 10_1101-2020_12_31_424971 420 104 take take VB 10_1101-2020_12_31_424971 420 105 place place NN 10_1101-2020_12_31_424971 420 106 out out IN 10_1101-2020_12_31_424971 420 107 of of IN 10_1101-2020_12_31_424971 420 108 the the DT 10_1101-2020_12_31_424971 420 109 GS1 GS1 NNP 10_1101-2020_12_31_424971 420 110 , , , 10_1101-2020_12_31_424971 420 111 GS2 GS2 NNP 10_1101-2020_12_31_424971 420 112 , , , 10_1101-2020_12_31_424971 420 113 and and CC 10_1101-2020_12_31_424971 420 114 Æ æ NN 10_1101-2020_12_31_424971 420 115 states state NNS 10_1101-2020_12_31_424971 420 116 ( ( -LRB- 10_1101-2020_12_31_424971 420 117 in in IN 10_1101-2020_12_31_424971 420 118 ( ( -LRB- 10_1101-2020_12_31_424971 420 119 i i NN 10_1101-2020_12_31_424971 420 120 ) ) -RRB- 10_1101-2020_12_31_424971 420 121 , , , 10_1101-2020_12_31_424971 420 122 973 973 CD 10_1101-2020_12_31_424971 420 123 ( ( -LRB- 10_1101-2020_12_31_424971 420 124 iv iv NN 10_1101-2020_12_31_424971 420 125 ) ) -RRB- 10_1101-2020_12_31_424971 420 126 , , , 10_1101-2020_12_31_424971 420 127 ( ( -LRB- 10_1101-2020_12_31_424971 420 128 v v NN 10_1101-2020_12_31_424971 420 129 ) ) -RRB- 10_1101-2020_12_31_424971 420 130 , , , 10_1101-2020_12_31_424971 420 131 and and CC 10_1101-2020_12_31_424971 420 132 ( ( -LRB- 10_1101-2020_12_31_424971 420 133 vi vi NNP 10_1101-2020_12_31_424971 420 134 ) ) -RRB- 10_1101-2020_12_31_424971 420 135 ) ) -RRB- 10_1101-2020_12_31_424971 420 136 or or CC 10_1101-2020_12_31_424971 420 137 out out IN 10_1101-2020_12_31_424971 420 138 of of IN 10_1101-2020_12_31_424971 420 139 the the DT 10_1101-2020_12_31_424971 420 140 EF EF NNP 10_1101-2020_12_31_424971 420 141 - - HYPH 10_1101-2020_12_31_424971 420 142 G G NNP 10_1101-2020_12_31_424971 420 143 - - HYPH 10_1101-2020_12_31_424971 420 144 bound bind VBN 10_1101-2020_12_31_424971 420 145 GS2-like GS2-like NNP 10_1101-2020_12_31_424971 420 146 , , , 10_1101-2020_12_31_424971 420 147 POST POST NNP 10_1101-2020_12_31_424971 420 148 , , , 10_1101-2020_12_31_424971 420 149 and and CC 10_1101-2020_12_31_424971 420 150 Æ Æ NNP 10_1101-2020_12_31_424971 420 151 states state NNS 10_1101-2020_12_31_424971 420 152 ( ( -LRB- 10_1101-2020_12_31_424971 420 153 in in IN 10_1101-2020_12_31_424971 420 154 ( ( -LRB- 10_1101-2020_12_31_424971 420 155 ii ii NNP 10_1101-2020_12_31_424971 420 156 ) ) -RRB- 10_1101-2020_12_31_424971 420 157 and and CC 10_1101-2020_12_31_424971 420 158 ( ( -LRB- 10_1101-2020_12_31_424971 420 159 vi vi NNP 10_1101-2020_12_31_424971 420 160 ) ) -RRB- 10_1101-2020_12_31_424971 420 161 ) ) -RRB- 10_1101-2020_12_31_424971 420 162 . . . 10_1101-2020_12_31_424971 421 1 Each each DT 10_1101-2020_12_31_424971 421 2 974 974 CD 10_1101-2020_12_31_424971 421 3 element element NN 10_1101-2020_12_31_424971 421 4 of of IN 10_1101-2020_12_31_424971 421 5 this this DT 10_1101-2020_12_31_424971 421 6 matrix matrix NN 10_1101-2020_12_31_424971 421 7 parameterizes parameterize VBZ 10_1101-2020_12_31_424971 421 8 a a DT 10_1101-2020_12_31_424971 421 9 Dirichlet Dirichlet NNP 10_1101-2020_12_31_424971 421 10 distribution distribution NN 10_1101-2020_12_31_424971 421 11 , , , 10_1101-2020_12_31_424971 421 12 from from IN 10_1101-2020_12_31_424971 421 13 which which WDT 10_1101-2020_12_31_424971 421 14 we -PRON- PRP 10_1101-2020_12_31_424971 421 15 calculated calculate VBD 10_1101-2020_12_31_424971 421 16 the the DT 10_1101-2020_12_31_424971 421 17 mean mean NN 10_1101-2020_12_31_424971 421 18 975 975 CD 10_1101-2020_12_31_424971 421 19 and and CC 10_1101-2020_12_31_424971 421 20 the the DT 10_1101-2020_12_31_424971 421 21 square square JJ 10_1101-2020_12_31_424971 421 22 root root NN 10_1101-2020_12_31_424971 421 23 of of IN 10_1101-2020_12_31_424971 421 24 the the DT 10_1101-2020_12_31_424971 421 25 variance variance NN 10_1101-2020_12_31_424971 421 26 for for IN 10_1101-2020_12_31_424971 421 27 four four CD 10_1101-2020_12_31_424971 421 28 transition transition NN 10_1101-2020_12_31_424971 421 29 probabilities probability NNS 10_1101-2020_12_31_424971 421 30 pGS1→GS2 pgs1→gs2 CD 10_1101-2020_12_31_424971 421 31 , , , 10_1101-2020_12_31_424971 421 32 pGS2→GS1 pGS2→GS1 NNP 10_1101-2020_12_31_424971 421 33 , , , 10_1101-2020_12_31_424971 421 34 pGS1→Æ pGS1→Æ NNP 10_1101-2020_12_31_424971 421 35 , , , 10_1101-2020_12_31_424971 421 36 and and CC 10_1101-2020_12_31_424971 421 37 976 976 CD 10_1101-2020_12_31_424971 421 38 pGS2→Æ pgs2→æ NN 10_1101-2020_12_31_424971 421 39 ( ( -LRB- 10_1101-2020_12_31_424971 421 40 in in FW 10_1101-2020_12_31_424971 421 41 ( ( -LRB- 10_1101-2020_12_31_424971 421 42 i i NN 10_1101-2020_12_31_424971 421 43 ) ) -RRB- 10_1101-2020_12_31_424971 421 44 , , , 10_1101-2020_12_31_424971 421 45 ( ( -LRB- 10_1101-2020_12_31_424971 421 46 iv iv NN 10_1101-2020_12_31_424971 421 47 ) ) -RRB- 10_1101-2020_12_31_424971 421 48 , , , 10_1101-2020_12_31_424971 421 49 ( ( -LRB- 10_1101-2020_12_31_424971 421 50 v v NN 10_1101-2020_12_31_424971 421 51 ) ) -RRB- 10_1101-2020_12_31_424971 421 52 , , , 10_1101-2020_12_31_424971 421 53 and and CC 10_1101-2020_12_31_424971 421 54 ( ( -LRB- 10_1101-2020_12_31_424971 421 55 vi vi NNP 10_1101-2020_12_31_424971 421 56 ) ) -RRB- 10_1101-2020_12_31_424971 421 57 ) ) -RRB- 10_1101-2020_12_31_424971 421 58 or or CC 10_1101-2020_12_31_424971 421 59 for for IN 10_1101-2020_12_31_424971 421 60 three three CD 10_1101-2020_12_31_424971 421 61 transition transition NN 10_1101-2020_12_31_424971 421 62 probabilities probability NNS 10_1101-2020_12_31_424971 421 63 pGS2→POST pGS2→POST NNS 10_1101-2020_12_31_424971 421 64 , , , 10_1101-2020_12_31_424971 421 65 pGS2(G)→Æ pGS2(G)→Æ NNS 10_1101-2020_12_31_424971 421 66 , , , 10_1101-2020_12_31_424971 421 67 and and CC 10_1101-2020_12_31_424971 421 68 pPOST→977 pPOST→977 NNP 10_1101-2020_12_31_424971 421 69 Æ Æ NNP 10_1101-2020_12_31_424971 421 70 ( ( -LRB- 10_1101-2020_12_31_424971 421 71 in in IN 10_1101-2020_12_31_424971 421 72 ( ( -LRB- 10_1101-2020_12_31_424971 421 73 ii ii NNP 10_1101-2020_12_31_424971 421 74 ) ) -RRB- 10_1101-2020_12_31_424971 421 75 and and CC 10_1101-2020_12_31_424971 421 76 ( ( -LRB- 10_1101-2020_12_31_424971 421 77 vi vi NNP 10_1101-2020_12_31_424971 421 78 ) ) -RRB- 10_1101-2020_12_31_424971 421 79 ) ) -RRB- 10_1101-2020_12_31_424971 421 80 . . . 10_1101-2020_12_31_424971 422 1 These these DT 10_1101-2020_12_31_424971 422 2 transition transition NN 10_1101-2020_12_31_424971 422 3 probabilities probability NNS 10_1101-2020_12_31_424971 422 4 were be VBD 10_1101-2020_12_31_424971 422 5 then then RB 10_1101-2020_12_31_424971 422 6 used use VBN 10_1101-2020_12_31_424971 422 7 to to TO 10_1101-2020_12_31_424971 422 8 calculate calculate VB 10_1101-2020_12_31_424971 422 9 the the DT 10_1101-2020_12_31_424971 422 10 corresponding correspond VBG 10_1101-2020_12_31_424971 422 11 978 978 CD 10_1101-2020_12_31_424971 422 12 four four CD 10_1101-2020_12_31_424971 422 13 rate rate NN 10_1101-2020_12_31_424971 422 14 constants constant NNS 10_1101-2020_12_31_424971 422 15 , , , 10_1101-2020_12_31_424971 422 16 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 422 17 , , , 10_1101-2020_12_31_424971 422 18 kGS2→GS1 kGS2→GS1 NNP 10_1101-2020_12_31_424971 422 19 , , , 10_1101-2020_12_31_424971 422 20 kGS1→Æ kGS1→Æ NNP 10_1101-2020_12_31_424971 422 21 , , , 10_1101-2020_12_31_424971 422 22 and and CC 10_1101-2020_12_31_424971 422 23 kGS2→Æ kGS2→Æ NNP 10_1101-2020_12_31_424971 422 24 ( ( -LRB- 10_1101-2020_12_31_424971 422 25 in in IN 10_1101-2020_12_31_424971 422 26 ( ( -LRB- 10_1101-2020_12_31_424971 422 27 i i NN 10_1101-2020_12_31_424971 422 28 ) ) -RRB- 10_1101-2020_12_31_424971 422 29 , , , 10_1101-2020_12_31_424971 422 30 ( ( -LRB- 10_1101-2020_12_31_424971 422 31 iv iv NN 10_1101-2020_12_31_424971 422 32 ) ) -RRB- 10_1101-2020_12_31_424971 422 33 , , , 10_1101-2020_12_31_424971 422 34 ( ( -LRB- 10_1101-2020_12_31_424971 422 35 v v NN 10_1101-2020_12_31_424971 422 36 ) ) -RRB- 10_1101-2020_12_31_424971 422 37 , , , 10_1101-2020_12_31_424971 422 38 and and CC 10_1101-2020_12_31_424971 422 39 ( ( -LRB- 10_1101-2020_12_31_424971 422 40 vi vi NNP 10_1101-2020_12_31_424971 422 41 ) ) -RRB- 10_1101-2020_12_31_424971 422 42 ) ) -RRB- 10_1101-2020_12_31_424971 422 43 or or CC 10_1101-2020_12_31_424971 422 44 three three CD 10_1101-2020_12_31_424971 422 45 rate rate NN 10_1101-2020_12_31_424971 422 46 979 979 CD 10_1101-2020_12_31_424971 422 47 constants constant NNS 10_1101-2020_12_31_424971 422 48 , , , 10_1101-2020_12_31_424971 422 49 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 422 50 , , , 10_1101-2020_12_31_424971 422 51 kGS2(G)→Æ kGS2(G)→Æ NNP 10_1101-2020_12_31_424971 422 52 , , , 10_1101-2020_12_31_424971 422 53 and and CC 10_1101-2020_12_31_424971 422 54 kPOST→Æ kPOST→Æ NNP 10_1101-2020_12_31_424971 422 55 ( ( -LRB- 10_1101-2020_12_31_424971 422 56 in in IN 10_1101-2020_12_31_424971 422 57 ( ( -LRB- 10_1101-2020_12_31_424971 422 58 ii ii NNP 10_1101-2020_12_31_424971 422 59 ) ) -RRB- 10_1101-2020_12_31_424971 422 60 and and CC 10_1101-2020_12_31_424971 422 61 ( ( -LRB- 10_1101-2020_12_31_424971 422 62 vi vi NNP 10_1101-2020_12_31_424971 422 63 ) ) -RRB- 10_1101-2020_12_31_424971 422 64 ) ) -RRB- 10_1101-2020_12_31_424971 422 65 using use VBG 10_1101-2020_12_31_424971 422 66 the the DT 10_1101-2020_12_31_424971 422 67 equation equation NN 10_1101-2020_12_31_424971 422 68 980 980 CD 10_1101-2020_12_31_424971 422 69 .CC .CC , 10_1101-2020_12_31_424971 422 70 - - HYPH 10_1101-2020_12_31_424971 422 71 BY by IN 10_1101-2020_12_31_424971 422 72 - - HYPH 10_1101-2020_12_31_424971 422 73 NC NC NNP 10_1101-2020_12_31_424971 422 74 - - HYPH 10_1101-2020_12_31_424971 422 75 ND ND NNP 10_1101-2020_12_31_424971 422 76 4.0 4.0 CD 10_1101-2020_12_31_424971 422 77 International International NNP 10_1101-2020_12_31_424971 422 78 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 422 79 under under IN 10_1101-2020_12_31_424971 422 80 a a DT 10_1101-2020_12_31_424971 422 81 ( ( -LRB- 10_1101-2020_12_31_424971 422 82 which which WDT 10_1101-2020_12_31_424971 422 83 was be VBD 10_1101-2020_12_31_424971 422 84 not not RB 10_1101-2020_12_31_424971 422 85 certified certify VBN 10_1101-2020_12_31_424971 422 86 by by IN 10_1101-2020_12_31_424971 422 87 peer peer NN 10_1101-2020_12_31_424971 422 88 review review NN 10_1101-2020_12_31_424971 422 89 ) ) -RRB- 10_1101-2020_12_31_424971 422 90 is be VBZ 10_1101-2020_12_31_424971 422 91 the the DT 10_1101-2020_12_31_424971 422 92 author author NN 10_1101-2020_12_31_424971 422 93 / / SYM 10_1101-2020_12_31_424971 422 94 funder funder NN 10_1101-2020_12_31_424971 422 95 , , , 10_1101-2020_12_31_424971 422 96 who who WP 10_1101-2020_12_31_424971 422 97 has have VBZ 10_1101-2020_12_31_424971 422 98 granted grant VBN 10_1101-2020_12_31_424971 422 99 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 422 100 a a DT 10_1101-2020_12_31_424971 422 101 license license NN 10_1101-2020_12_31_424971 422 102 to to TO 10_1101-2020_12_31_424971 422 103 display display VB 10_1101-2020_12_31_424971 422 104 the the DT 10_1101-2020_12_31_424971 422 105 preprint preprint NN 10_1101-2020_12_31_424971 422 106 in in IN 10_1101-2020_12_31_424971 422 107 perpetuity perpetuity NN 10_1101-2020_12_31_424971 422 108 . . . 10_1101-2020_12_31_424971 423 1 It -PRON- PRP 10_1101-2020_12_31_424971 423 2 is be VBZ 10_1101-2020_12_31_424971 423 3 made make VBN 10_1101-2020_12_31_424971 423 4 The the DT 10_1101-2020_12_31_424971 423 5 copyright copyright NN 10_1101-2020_12_31_424971 423 6 holder holder NN 10_1101-2020_12_31_424971 423 7 for for IN 10_1101-2020_12_31_424971 423 8 this this DT 10_1101-2020_12_31_424971 423 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 423 10 version version NN 10_1101-2020_12_31_424971 423 11 posted post VBD 10_1101-2020_12_31_424971 423 12 January January NNP 10_1101-2020_12_31_424971 423 13 2 2 CD 10_1101-2020_12_31_424971 423 14 , , , 10_1101-2020_12_31_424971 423 15 2021 2021 CD 10_1101-2020_12_31_424971 423 16 . . . 10_1101-2020_12_31_424971 423 17 ; ; : 10_1101-2020_12_31_424971 423 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 423 19 : : : 10_1101-2020_12_31_424971 423 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 423 21 preprint preprint NN 10_1101-2020_12_31_424971 423 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NNP 10_1101-2020_12_31_424971 423 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 423 24 40 40 CD 10_1101-2020_12_31_424971 423 25 𝑘 𝑘 NNS 10_1101-2020_12_31_424971 423 26 = = SYM 10_1101-2020_12_31_424971 423 27 − − NNP 10_1101-2020_12_31_424971 423 28 ln(1 ln(1 NN 10_1101-2020_12_31_424971 423 29 − − NNP 10_1101-2020_12_31_424971 423 30 𝑝 𝑝 NNP 10_1101-2020_12_31_424971 423 31 ) ) -RRB- 10_1101-2020_12_31_424971 423 32 𝑡 𝑡 NN 10_1101-2020_12_31_424971 423 33 , , , 10_1101-2020_12_31_424971 423 34 ( ( -LRB- 10_1101-2020_12_31_424971 423 35 3 3 CD 10_1101-2020_12_31_424971 423 36 ) ) -RRB- 10_1101-2020_12_31_424971 423 37 981 981 CD 10_1101-2020_12_31_424971 423 38 where where WRB 10_1101-2020_12_31_424971 423 39 t t NN 10_1101-2020_12_31_424971 423 40 is be VBZ 10_1101-2020_12_31_424971 423 41 the the DT 10_1101-2020_12_31_424971 423 42 time time NN 10_1101-2020_12_31_424971 423 43 interval interval NN 10_1101-2020_12_31_424971 423 44 between between IN 10_1101-2020_12_31_424971 423 45 data datum NNS 10_1101-2020_12_31_424971 423 46 points point NNS 10_1101-2020_12_31_424971 423 47 ( ( -LRB- 10_1101-2020_12_31_424971 423 48 t t NNP 10_1101-2020_12_31_424971 423 49 = = SYM 10_1101-2020_12_31_424971 423 50 0.1 0.1 CD 10_1101-2020_12_31_424971 423 51 s s NN 10_1101-2020_12_31_424971 423 52 ) ) -RRB- 10_1101-2020_12_31_424971 423 53 . . . 10_1101-2020_12_31_424971 424 1 We -PRON- PRP 10_1101-2020_12_31_424971 424 2 propagated propagate VBD 10_1101-2020_12_31_424971 424 3 the the DT 10_1101-2020_12_31_424971 424 4 error error NN 10_1101-2020_12_31_424971 424 5 for for IN 10_1101-2020_12_31_424971 424 6 the the DT 10_1101-2020_12_31_424971 424 7 982 982 CD 10_1101-2020_12_31_424971 424 8 transition transition NN 10_1101-2020_12_31_424971 424 9 probabilities probability NNS 10_1101-2020_12_31_424971 424 10 into into IN 10_1101-2020_12_31_424971 424 11 the the DT 10_1101-2020_12_31_424971 424 12 error error NN 10_1101-2020_12_31_424971 424 13 for for IN 10_1101-2020_12_31_424971 424 14 the the DT 10_1101-2020_12_31_424971 424 15 rate rate NN 10_1101-2020_12_31_424971 424 16 constants constant NNS 10_1101-2020_12_31_424971 424 17 using use VBG 10_1101-2020_12_31_424971 424 18 the the DT 10_1101-2020_12_31_424971 424 19 equation equation NN 10_1101-2020_12_31_424971 424 20 983 983 CD 10_1101-2020_12_31_424971 424 21 𝜎2 𝜎2 NN 10_1101-2020_12_31_424971 424 22 = = NFP 10_1101-2020_12_31_424971 424 23 𝜎3 𝜎3 NN 10_1101-2020_12_31_424971 424 24 ( ( -LRB- 10_1101-2020_12_31_424971 424 25 1 1 CD 10_1101-2020_12_31_424971 424 26 − − NNP 10_1101-2020_12_31_424971 424 27 𝑝 𝑝 NNP 10_1101-2020_12_31_424971 424 28 ) ) -RRB- 10_1101-2020_12_31_424971 424 29 × × NN 10_1101-2020_12_31_424971 424 30 𝑡 𝑡 XX 10_1101-2020_12_31_424971 424 31 , , , 10_1101-2020_12_31_424971 424 32 ( ( -LRB- 10_1101-2020_12_31_424971 424 33 4 4 CD 10_1101-2020_12_31_424971 424 34 ) ) -RRB- 10_1101-2020_12_31_424971 424 35 984 984 CD 10_1101-2020_12_31_424971 424 36 where where WRB 10_1101-2020_12_31_424971 424 37 𝜎3 𝜎3 NNP 10_1101-2020_12_31_424971 424 38 is be VBZ 10_1101-2020_12_31_424971 424 39 the the DT 10_1101-2020_12_31_424971 424 40 standard standard JJ 10_1101-2020_12_31_424971 424 41 deviation deviation NN 10_1101-2020_12_31_424971 424 42 of of IN 10_1101-2020_12_31_424971 424 43 the the DT 10_1101-2020_12_31_424971 424 44 variance variance NN 10_1101-2020_12_31_424971 424 45 of of IN 10_1101-2020_12_31_424971 424 46 p p NN 10_1101-2020_12_31_424971 424 47 and and CC 10_1101-2020_12_31_424971 424 48 𝜎2 𝜎2 NNP 10_1101-2020_12_31_424971 424 49 is be VBZ 10_1101-2020_12_31_424971 424 50 the the DT 10_1101-2020_12_31_424971 424 51 standard standard JJ 10_1101-2020_12_31_424971 424 52 deviation deviation NN 10_1101-2020_12_31_424971 424 53 of of IN 10_1101-2020_12_31_424971 424 54 the the DT 10_1101-2020_12_31_424971 424 55 985 985 CD 10_1101-2020_12_31_424971 424 56 variance variance NN 10_1101-2020_12_31_424971 424 57 of of IN 10_1101-2020_12_31_424971 424 58 k. k. NNP 10_1101-2020_12_31_424971 424 59 Keq Keq NNP 10_1101-2020_12_31_424971 424 60 in in IN 10_1101-2020_12_31_424971 424 61 ( ( -LRB- 10_1101-2020_12_31_424971 424 62 i i NN 10_1101-2020_12_31_424971 424 63 ) ) -RRB- 10_1101-2020_12_31_424971 424 64 , , , 10_1101-2020_12_31_424971 424 65 ( ( -LRB- 10_1101-2020_12_31_424971 424 66 iv iv LS 10_1101-2020_12_31_424971 424 67 ) ) -RRB- 10_1101-2020_12_31_424971 424 68 , , , 10_1101-2020_12_31_424971 424 69 and and CC 10_1101-2020_12_31_424971 424 70 ( ( -LRB- 10_1101-2020_12_31_424971 424 71 v v NN 10_1101-2020_12_31_424971 424 72 ) ) -RRB- 10_1101-2020_12_31_424971 424 73 was be VBD 10_1101-2020_12_31_424971 424 74 determined determine VBN 10_1101-2020_12_31_424971 424 75 using use VBG 10_1101-2020_12_31_424971 424 76 the the DT 10_1101-2020_12_31_424971 424 77 equation equation NN 10_1101-2020_12_31_424971 424 78 Keq Keq NNP 10_1101-2020_12_31_424971 424 79 = = SYM 10_1101-2020_12_31_424971 424 80 kGS1→GS2 kGS1→GS2 NNP 10_1101-2020_12_31_424971 424 81 / / SYM 10_1101-2020_12_31_424971 424 82 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 424 83 . . . 10_1101-2020_12_31_424971 425 1 986 986 CD 10_1101-2020_12_31_424971 425 2 The the DT 10_1101-2020_12_31_424971 425 3 fractional fractional JJ 10_1101-2020_12_31_424971 425 4 populations population NNS 10_1101-2020_12_31_424971 425 5 of of IN 10_1101-2020_12_31_424971 425 6 the the DT 10_1101-2020_12_31_424971 425 7 POST POST NNP 10_1101-2020_12_31_424971 425 8 complex complex NN 10_1101-2020_12_31_424971 425 9 in in IN 10_1101-2020_12_31_424971 425 10 ( ( -LRB- 10_1101-2020_12_31_424971 425 11 iii iii NN 10_1101-2020_12_31_424971 425 12 ) ) -RRB- 10_1101-2020_12_31_424971 425 13 and and CC 10_1101-2020_12_31_424971 425 14 the the DT 10_1101-2020_12_31_424971 425 15 corresponding corresponding JJ 10_1101-2020_12_31_424971 425 16 errors error NNS 10_1101-2020_12_31_424971 425 17 were be VBD 10_1101-2020_12_31_424971 425 18 987 987 CD 10_1101-2020_12_31_424971 425 19 calculated calculate VBN 10_1101-2020_12_31_424971 425 20 by by IN 10_1101-2020_12_31_424971 425 21 marginalizing marginalize VBG 10_1101-2020_12_31_424971 425 22 , , , 10_1101-2020_12_31_424971 425 23 which which WDT 10_1101-2020_12_31_424971 425 24 in in IN 10_1101-2020_12_31_424971 425 25 this this DT 10_1101-2020_12_31_424971 425 26 case case NN 10_1101-2020_12_31_424971 425 27 simply simply RB 10_1101-2020_12_31_424971 425 28 amounts amount VBZ 10_1101-2020_12_31_424971 425 29 to to IN 10_1101-2020_12_31_424971 425 30 calculating calculate VBG 10_1101-2020_12_31_424971 425 31 the the DT 10_1101-2020_12_31_424971 425 32 mean mean NN 10_1101-2020_12_31_424971 425 33 and and CC 10_1101-2020_12_31_424971 425 34 the the DT 10_1101-2020_12_31_424971 425 35 988 988 CD 10_1101-2020_12_31_424971 425 36 standard standard JJ 10_1101-2020_12_31_424971 425 37 error error NN 10_1101-2020_12_31_424971 425 38 of of IN 10_1101-2020_12_31_424971 425 39 the the DT 10_1101-2020_12_31_424971 425 40 mean mean NN 10_1101-2020_12_31_424971 425 41 , , , 10_1101-2020_12_31_424971 425 42 for for IN 10_1101-2020_12_31_424971 425 43 the the DT 10_1101-2020_12_31_424971 425 44 conditional conditional JJ 10_1101-2020_12_31_424971 425 45 probabilities probability NNS 10_1101-2020_12_31_424971 425 46 of of IN 10_1101-2020_12_31_424971 425 47 each each DT 10_1101-2020_12_31_424971 425 48 EFRET EFRET NNP 10_1101-2020_12_31_424971 425 49 data datum NNS 10_1101-2020_12_31_424971 425 50 point point NN 10_1101-2020_12_31_424971 425 51 given give VBN 10_1101-2020_12_31_424971 425 52 each each DT 10_1101-2020_12_31_424971 425 53 989 989 CD 10_1101-2020_12_31_424971 425 54 hidden hide VBN 10_1101-2020_12_31_424971 425 55 state state NN 10_1101-2020_12_31_424971 425 56 . . . 10_1101-2020_12_31_424971 426 1 Because because IN 10_1101-2020_12_31_424971 426 2 the the DT 10_1101-2020_12_31_424971 426 3 data data NN 10_1101-2020_12_31_424971 426 4 points point NNS 10_1101-2020_12_31_424971 426 5 preceding precede VBG 10_1101-2020_12_31_424971 426 6 the the DT 10_1101-2020_12_31_424971 426 7 initial initial JJ 10_1101-2020_12_31_424971 426 8 70S 70S NNP 10_1101-2020_12_31_424971 426 9 IC→GS2 ic→gs2 NN 10_1101-2020_12_31_424971 426 10 transition transition NN 10_1101-2020_12_31_424971 426 11 in in IN 10_1101-2020_12_31_424971 426 12 the the DT 10_1101-2020_12_31_424971 426 13 pre-990 pre-990 NNP 10_1101-2020_12_31_424971 426 14 steady steady JJ 10_1101-2020_12_31_424971 426 15 - - HYPH 10_1101-2020_12_31_424971 426 16 state state NN 10_1101-2020_12_31_424971 426 17 movies movie NNS 10_1101-2020_12_31_424971 426 18 do do VBP 10_1101-2020_12_31_424971 426 19 not not RB 10_1101-2020_12_31_424971 426 20 contribute contribute VB 10_1101-2020_12_31_424971 426 21 to to IN 10_1101-2020_12_31_424971 426 22 the the DT 10_1101-2020_12_31_424971 426 23 kinetic kinetic JJ 10_1101-2020_12_31_424971 426 24 and/or and/or CC 10_1101-2020_12_31_424971 426 25 thermodynamic thermodynamic JJ 10_1101-2020_12_31_424971 426 26 parameters parameter NNS 10_1101-2020_12_31_424971 426 27 in in IN 10_1101-2020_12_31_424971 426 28 ( ( -LRB- 10_1101-2020_12_31_424971 426 29 i)-(vi i)-(vi NNP 10_1101-2020_12_31_424971 426 30 ) ) -RRB- 10_1101-2020_12_31_424971 426 31 , , , 10_1101-2020_12_31_424971 426 32 991 991 CD 10_1101-2020_12_31_424971 426 33 these these DT 10_1101-2020_12_31_424971 426 34 data datum NNS 10_1101-2020_12_31_424971 426 35 points point NNS 10_1101-2020_12_31_424971 426 36 were be VBD 10_1101-2020_12_31_424971 426 37 not not RB 10_1101-2020_12_31_424971 426 38 included include VBN 10_1101-2020_12_31_424971 426 39 in in IN 10_1101-2020_12_31_424971 426 40 the the DT 10_1101-2020_12_31_424971 426 41 analyses analysis NNS 10_1101-2020_12_31_424971 426 42 that that WDT 10_1101-2020_12_31_424971 426 43 were be VBD 10_1101-2020_12_31_424971 426 44 used use VBN 10_1101-2020_12_31_424971 426 45 to to TO 10_1101-2020_12_31_424971 426 46 determine determine VB 10_1101-2020_12_31_424971 426 47 these these DT 10_1101-2020_12_31_424971 426 48 992 992 CD 10_1101-2020_12_31_424971 426 49 thermodynamic thermodynamic JJ 10_1101-2020_12_31_424971 426 50 parameters parameter NNS 10_1101-2020_12_31_424971 426 51 . . . 10_1101-2020_12_31_424971 427 1 993 993 CD 10_1101-2020_12_31_424971 427 2 994 994 CD 10_1101-2020_12_31_424971 427 3 QUANTIFICATION QUANTIFICATION NNS 10_1101-2020_12_31_424971 427 4 AND and CC 10_1101-2020_12_31_424971 427 5 STATISTICAL statistical NN 10_1101-2020_12_31_424971 427 6 ANALYSES analyse VBZ 10_1101-2020_12_31_424971 427 7 995 995 CD 10_1101-2020_12_31_424971 427 8 All all DT 10_1101-2020_12_31_424971 427 9 ensemble ensemble JJ 10_1101-2020_12_31_424971 427 10 biochemical biochemical JJ 10_1101-2020_12_31_424971 427 11 experiments experiment NNS 10_1101-2020_12_31_424971 427 12 and and CC 10_1101-2020_12_31_424971 427 13 cell cell NN 10_1101-2020_12_31_424971 427 14 - - HYPH 10_1101-2020_12_31_424971 427 15 based base VBN 10_1101-2020_12_31_424971 427 16 reporter reporter NN 10_1101-2020_12_31_424971 427 17 assays assays RB 10_1101-2020_12_31_424971 427 18 were be VBD 10_1101-2020_12_31_424971 427 19 repeated repeat VBN 10_1101-2020_12_31_424971 427 20 at at IN 10_1101-2020_12_31_424971 427 21 least least RBS 10_1101-2020_12_31_424971 427 22 996 996 CD 10_1101-2020_12_31_424971 427 23 three three CD 10_1101-2020_12_31_424971 427 24 times time NNS 10_1101-2020_12_31_424971 427 25 and and CC 10_1101-2020_12_31_424971 427 26 the the DT 10_1101-2020_12_31_424971 427 27 mean mean JJ 10_1101-2020_12_31_424971 427 28 values value NNS 10_1101-2020_12_31_424971 427 29 and and CC 10_1101-2020_12_31_424971 427 30 standard standard JJ 10_1101-2020_12_31_424971 427 31 deviations deviation NNS 10_1101-2020_12_31_424971 427 32 for for IN 10_1101-2020_12_31_424971 427 33 each each DT 10_1101-2020_12_31_424971 427 34 experiment experiment NN 10_1101-2020_12_31_424971 427 35 or or CC 10_1101-2020_12_31_424971 427 36 assay assay NNP 10_1101-2020_12_31_424971 427 37 are be VBP 10_1101-2020_12_31_424971 427 38 997 997 CD 10_1101-2020_12_31_424971 427 39 reported report VBN 10_1101-2020_12_31_424971 427 40 . . . 10_1101-2020_12_31_424971 428 1 Technical technical JJ 10_1101-2020_12_31_424971 428 2 replicates replicate NNS 10_1101-2020_12_31_424971 428 3 of of IN 10_1101-2020_12_31_424971 428 4 all all DT 10_1101-2020_12_31_424971 428 5 smFRET smfret NN 10_1101-2020_12_31_424971 428 6 experiments experiment NNS 10_1101-2020_12_31_424971 428 7 were be VBD 10_1101-2020_12_31_424971 428 8 repeated repeat VBN 10_1101-2020_12_31_424971 428 9 at at IN 10_1101-2020_12_31_424971 428 10 least least RBS 10_1101-2020_12_31_424971 428 11 three three CD 10_1101-2020_12_31_424971 428 12 times time NNS 10_1101-2020_12_31_424971 428 13 and and CC 10_1101-2020_12_31_424971 428 14 998 998 CD 10_1101-2020_12_31_424971 428 15 trajectories trajectory NNS 10_1101-2020_12_31_424971 428 16 from from IN 10_1101-2020_12_31_424971 428 17 all all DT 10_1101-2020_12_31_424971 428 18 of of IN 10_1101-2020_12_31_424971 428 19 the the DT 10_1101-2020_12_31_424971 428 20 technical technical JJ 10_1101-2020_12_31_424971 428 21 replicates replicate NNS 10_1101-2020_12_31_424971 428 22 for for IN 10_1101-2020_12_31_424971 428 23 each each DT 10_1101-2020_12_31_424971 428 24 experiment experiment NN 10_1101-2020_12_31_424971 428 25 were be VBD 10_1101-2020_12_31_424971 428 26 combined combine VBN 10_1101-2020_12_31_424971 428 27 prior prior RB 10_1101-2020_12_31_424971 428 28 to to IN 10_1101-2020_12_31_424971 428 29 999 999 CD 10_1101-2020_12_31_424971 428 30 generating generate VBG 10_1101-2020_12_31_424971 428 31 the the DT 10_1101-2020_12_31_424971 428 32 surface surface NN 10_1101-2020_12_31_424971 428 33 contour contour NN 10_1101-2020_12_31_424971 428 34 plot plot NN 10_1101-2020_12_31_424971 428 35 of of IN 10_1101-2020_12_31_424971 428 36 the the DT 10_1101-2020_12_31_424971 428 37 time time NN 10_1101-2020_12_31_424971 428 38 evolution evolution NN 10_1101-2020_12_31_424971 428 39 of of IN 10_1101-2020_12_31_424971 428 40 population population NN 10_1101-2020_12_31_424971 428 41 FRET FRET NNS 10_1101-2020_12_31_424971 428 42 and and CC 10_1101-2020_12_31_424971 428 43 modeling model VBG 10_1101-2020_12_31_424971 428 44 with with IN 10_1101-2020_12_31_424971 428 45 1000 1000 CD 10_1101-2020_12_31_424971 428 46 the the DT 10_1101-2020_12_31_424971 428 47 HMM HMM NNP 10_1101-2020_12_31_424971 428 48 . . . 10_1101-2020_12_31_424971 429 1 Mean mean JJ 10_1101-2020_12_31_424971 429 2 values value NNS 10_1101-2020_12_31_424971 429 3 and and CC 10_1101-2020_12_31_424971 429 4 errors error NNS 10_1101-2020_12_31_424971 429 5 for for IN 10_1101-2020_12_31_424971 429 6 the the DT 10_1101-2020_12_31_424971 429 7 transition transition NN 10_1101-2020_12_31_424971 429 8 rates rate NNS 10_1101-2020_12_31_424971 429 9 and and CC 10_1101-2020_12_31_424971 429 10 fractional fractional JJ 10_1101-2020_12_31_424971 429 11 populations population NNS 10_1101-2020_12_31_424971 429 12 determined determine VBD 10_1101-2020_12_31_424971 429 13 1001 1001 CD 10_1101-2020_12_31_424971 429 14 from from IN 10_1101-2020_12_31_424971 429 15 modeling model VBG 10_1101-2020_12_31_424971 429 16 with with IN 10_1101-2020_12_31_424971 429 17 an an DT 10_1101-2020_12_31_424971 429 18 HMM HMM NNP 10_1101-2020_12_31_424971 429 19 are be VBP 10_1101-2020_12_31_424971 429 20 reported report VBN 10_1101-2020_12_31_424971 429 21 ( ( -LRB- 10_1101-2020_12_31_424971 429 22 for for IN 10_1101-2020_12_31_424971 429 23 details detail NNS 10_1101-2020_12_31_424971 429 24 see see VBP 10_1101-2020_12_31_424971 429 25 “ " `` 10_1101-2020_12_31_424971 429 26 Analysis analysis NN 10_1101-2020_12_31_424971 429 27 of of IN 10_1101-2020_12_31_424971 429 28 smFRET smFRET NNP 10_1101-2020_12_31_424971 429 29 experiments experiment NNS 10_1101-2020_12_31_424971 429 30 ” " '' 10_1101-2020_12_31_424971 429 31 in in IN 10_1101-2020_12_31_424971 429 32 1002 1002 CD 10_1101-2020_12_31_424971 429 33 Methods Methods NNPS 10_1101-2020_12_31_424971 429 34 ) ) -RRB- 10_1101-2020_12_31_424971 429 35 . . . 10_1101-2020_12_31_424971 430 1 Mean mean JJ 10_1101-2020_12_31_424971 430 2 values value NNS 10_1101-2020_12_31_424971 430 3 and and CC 10_1101-2020_12_31_424971 430 4 standard standard JJ 10_1101-2020_12_31_424971 430 5 deviations deviation NNS 10_1101-2020_12_31_424971 430 6 for for IN 10_1101-2020_12_31_424971 430 7 the the DT 10_1101-2020_12_31_424971 430 8 k70S k70s XX 10_1101-2020_12_31_424971 430 9 IC→GS2s IC→GS2s NNP 10_1101-2020_12_31_424971 430 10 were be VBD 10_1101-2020_12_31_424971 430 11 determined determine VBN 10_1101-2020_12_31_424971 430 12 from from IN 10_1101-2020_12_31_424971 430 13 1003 1003 CD 10_1101-2020_12_31_424971 430 14 technical technical JJ 10_1101-2020_12_31_424971 430 15 triplicates triplicate NNS 10_1101-2020_12_31_424971 430 16 of of IN 10_1101-2020_12_31_424971 430 17 the the DT 10_1101-2020_12_31_424971 430 18 survival survival NN 10_1101-2020_12_31_424971 430 19 plots plot NNS 10_1101-2020_12_31_424971 430 20 analysis analysis NN 10_1101-2020_12_31_424971 430 21 for for IN 10_1101-2020_12_31_424971 430 22 each each DT 10_1101-2020_12_31_424971 430 23 experiment experiment NN 10_1101-2020_12_31_424971 430 24 and and CC 10_1101-2020_12_31_424971 430 25 are be VBP 10_1101-2020_12_31_424971 430 26 reported report VBN 10_1101-2020_12_31_424971 430 27 . . . 10_1101-2020_12_31_424971 431 1 1004 1004 CD 10_1101-2020_12_31_424971 431 2 1005 1005 CD 10_1101-2020_12_31_424971 431 3 .CC .cc SYM 10_1101-2020_12_31_424971 431 4 - - : 10_1101-2020_12_31_424971 431 5 BY by IN 10_1101-2020_12_31_424971 431 6 - - HYPH 10_1101-2020_12_31_424971 431 7 NC NC NNP 10_1101-2020_12_31_424971 431 8 - - HYPH 10_1101-2020_12_31_424971 431 9 ND ND NNP 10_1101-2020_12_31_424971 431 10 4.0 4.0 CD 10_1101-2020_12_31_424971 431 11 International International NNP 10_1101-2020_12_31_424971 431 12 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 431 13 under under IN 10_1101-2020_12_31_424971 431 14 a a DT 10_1101-2020_12_31_424971 431 15 ( ( -LRB- 10_1101-2020_12_31_424971 431 16 which which WDT 10_1101-2020_12_31_424971 431 17 was be VBD 10_1101-2020_12_31_424971 431 18 not not RB 10_1101-2020_12_31_424971 431 19 certified certify VBN 10_1101-2020_12_31_424971 431 20 by by IN 10_1101-2020_12_31_424971 431 21 peer peer NN 10_1101-2020_12_31_424971 431 22 review review NN 10_1101-2020_12_31_424971 431 23 ) ) -RRB- 10_1101-2020_12_31_424971 431 24 is be VBZ 10_1101-2020_12_31_424971 431 25 the the DT 10_1101-2020_12_31_424971 431 26 author author NN 10_1101-2020_12_31_424971 431 27 / / SYM 10_1101-2020_12_31_424971 431 28 funder funder NN 10_1101-2020_12_31_424971 431 29 , , , 10_1101-2020_12_31_424971 431 30 who who WP 10_1101-2020_12_31_424971 431 31 has have VBZ 10_1101-2020_12_31_424971 431 32 granted grant VBN 10_1101-2020_12_31_424971 431 33 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 431 34 a a DT 10_1101-2020_12_31_424971 431 35 license license NN 10_1101-2020_12_31_424971 431 36 to to TO 10_1101-2020_12_31_424971 431 37 display display VB 10_1101-2020_12_31_424971 431 38 the the DT 10_1101-2020_12_31_424971 431 39 preprint preprint NN 10_1101-2020_12_31_424971 431 40 in in IN 10_1101-2020_12_31_424971 431 41 perpetuity perpetuity NN 10_1101-2020_12_31_424971 431 42 . . . 10_1101-2020_12_31_424971 432 1 It -PRON- PRP 10_1101-2020_12_31_424971 432 2 is be VBZ 10_1101-2020_12_31_424971 432 3 made make VBN 10_1101-2020_12_31_424971 432 4 The the DT 10_1101-2020_12_31_424971 432 5 copyright copyright NN 10_1101-2020_12_31_424971 432 6 holder holder NN 10_1101-2020_12_31_424971 432 7 for for IN 10_1101-2020_12_31_424971 432 8 this this DT 10_1101-2020_12_31_424971 432 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 432 10 version version NN 10_1101-2020_12_31_424971 432 11 posted post VBD 10_1101-2020_12_31_424971 432 12 January January NNP 10_1101-2020_12_31_424971 432 13 2 2 CD 10_1101-2020_12_31_424971 432 14 , , , 10_1101-2020_12_31_424971 432 15 2021 2021 CD 10_1101-2020_12_31_424971 432 16 . . . 10_1101-2020_12_31_424971 432 17 ; ; : 10_1101-2020_12_31_424971 432 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 432 19 : : : 10_1101-2020_12_31_424971 432 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 432 21 preprint preprint NN 10_1101-2020_12_31_424971 432 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 432 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 432 24 41 41 CD 10_1101-2020_12_31_424971 432 25 DATA datum NNS 10_1101-2020_12_31_424971 432 26 AND and CC 10_1101-2020_12_31_424971 432 27 CODE code NN 10_1101-2020_12_31_424971 432 28 AVAILABILITY AVAILABILITY NNS 10_1101-2020_12_31_424971 432 29 1006 1006 CD 10_1101-2020_12_31_424971 432 30 1007 1007 CD 10_1101-2020_12_31_424971 432 31 Data Data NNP 10_1101-2020_12_31_424971 432 32 Availability Availability NNP 10_1101-2020_12_31_424971 432 33 1008 1008 CD 10_1101-2020_12_31_424971 432 34 With with IN 10_1101-2020_12_31_424971 432 35 the the DT 10_1101-2020_12_31_424971 432 36 exception exception NN 10_1101-2020_12_31_424971 432 37 of of IN 10_1101-2020_12_31_424971 432 38 the the DT 10_1101-2020_12_31_424971 432 39 smFRET smFRET NNP 10_1101-2020_12_31_424971 432 40 data datum NNS 10_1101-2020_12_31_424971 432 41 , , , 10_1101-2020_12_31_424971 432 42 all all DT 10_1101-2020_12_31_424971 432 43 other other JJ 10_1101-2020_12_31_424971 432 44 data datum NNS 10_1101-2020_12_31_424971 432 45 supporting support VBG 10_1101-2020_12_31_424971 432 46 the the DT 10_1101-2020_12_31_424971 432 47 findings finding NNS 10_1101-2020_12_31_424971 432 48 of of IN 10_1101-2020_12_31_424971 432 49 this this DT 10_1101-2020_12_31_424971 432 50 study study NN 10_1101-2020_12_31_424971 432 51 are be VBP 10_1101-2020_12_31_424971 432 52 1009 1009 CD 10_1101-2020_12_31_424971 432 53 presented present VBN 10_1101-2020_12_31_424971 432 54 within within IN 10_1101-2020_12_31_424971 432 55 this this DT 10_1101-2020_12_31_424971 432 56 article article NN 10_1101-2020_12_31_424971 432 57 . . . 10_1101-2020_12_31_424971 433 1 Due due IN 10_1101-2020_12_31_424971 433 2 to to IN 10_1101-2020_12_31_424971 433 3 the the DT 10_1101-2020_12_31_424971 433 4 lack lack NN 10_1101-2020_12_31_424971 433 5 of of IN 10_1101-2020_12_31_424971 433 6 a a DT 10_1101-2020_12_31_424971 433 7 public public JJ 10_1101-2020_12_31_424971 433 8 repository repository NN 10_1101-2020_12_31_424971 433 9 for for IN 10_1101-2020_12_31_424971 433 10 smFRET smFRET NNP 10_1101-2020_12_31_424971 433 11 data datum NNS 10_1101-2020_12_31_424971 433 12 , , , 10_1101-2020_12_31_424971 433 13 the the DT 10_1101-2020_12_31_424971 433 14 smFRET smfret $ 10_1101-2020_12_31_424971 433 15 1010 1010 CD 10_1101-2020_12_31_424971 433 16 data datum NNS 10_1101-2020_12_31_424971 433 17 supporting support VBG 10_1101-2020_12_31_424971 433 18 the the DT 10_1101-2020_12_31_424971 433 19 findings finding NNS 10_1101-2020_12_31_424971 433 20 of of IN 10_1101-2020_12_31_424971 433 21 this this DT 10_1101-2020_12_31_424971 433 22 study study NN 10_1101-2020_12_31_424971 433 23 are be VBP 10_1101-2020_12_31_424971 433 24 available available JJ 10_1101-2020_12_31_424971 433 25 from from IN 10_1101-2020_12_31_424971 433 26 the the DT 10_1101-2020_12_31_424971 433 27 corresponding correspond VBG 10_1101-2020_12_31_424971 433 28 authors author NNS 10_1101-2020_12_31_424971 433 29 upon upon IN 10_1101-2020_12_31_424971 433 30 1011 1011 CD 10_1101-2020_12_31_424971 433 31 request request NN 10_1101-2020_12_31_424971 433 32 . . . 10_1101-2020_12_31_424971 434 1 Source source NN 10_1101-2020_12_31_424971 434 2 data datum NNS 10_1101-2020_12_31_424971 434 3 are be VBP 10_1101-2020_12_31_424971 434 4 provided provide VBN 10_1101-2020_12_31_424971 434 5 with with IN 10_1101-2020_12_31_424971 434 6 this this DT 10_1101-2020_12_31_424971 434 7 paper paper NN 10_1101-2020_12_31_424971 434 8 . . . 10_1101-2020_12_31_424971 435 1 1012 1012 CD 10_1101-2020_12_31_424971 435 2 Code Code NNP 10_1101-2020_12_31_424971 435 3 Availability Availability NNP 10_1101-2020_12_31_424971 435 4 1013 1013 CD 10_1101-2020_12_31_424971 435 5 The the DT 10_1101-2020_12_31_424971 435 6 code code NN 10_1101-2020_12_31_424971 435 7 used use VBD 10_1101-2020_12_31_424971 435 8 to to TO 10_1101-2020_12_31_424971 435 9 analyze analyze VB 10_1101-2020_12_31_424971 435 10 the the DT 10_1101-2020_12_31_424971 435 11 TIRF tirf NN 10_1101-2020_12_31_424971 435 12 movies movie NNS 10_1101-2020_12_31_424971 435 13 in in IN 10_1101-2020_12_31_424971 435 14 this this DT 10_1101-2020_12_31_424971 435 15 study study NN 10_1101-2020_12_31_424971 435 16 is be VBZ 10_1101-2020_12_31_424971 435 17 described describe VBN 10_1101-2020_12_31_424971 435 18 in in IN 10_1101-2020_12_31_424971 435 19 a a DT 10_1101-2020_12_31_424971 435 20 manuscript manuscript NN 10_1101-2020_12_31_424971 435 21 in in IN 10_1101-2020_12_31_424971 435 22 preparation preparation NN 10_1101-2020_12_31_424971 435 23 1014 1014 CD 10_1101-2020_12_31_424971 435 24 ( ( -LRB- 10_1101-2020_12_31_424971 435 25 Jason Jason NNP 10_1101-2020_12_31_424971 435 26 Hon Hon NNP 10_1101-2020_12_31_424971 435 27 , , , 10_1101-2020_12_31_424971 435 28 Colin Colin NNP 10_1101-2020_12_31_424971 435 29 Kinz Kinz NNP 10_1101-2020_12_31_424971 435 30 - - HYPH 10_1101-2020_12_31_424971 435 31 Thompson Thompson NNP 10_1101-2020_12_31_424971 435 32 , , , 10_1101-2020_12_31_424971 435 33 Ruben Ruben NNP 10_1101-2020_12_31_424971 435 34 L. L. NNP 10_1101-2020_12_31_424971 435 35 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 435 36 ) ) -RRB- 10_1101-2020_12_31_424971 435 37 , , , 10_1101-2020_12_31_424971 435 38 where where WRB 10_1101-2020_12_31_424971 435 39 R.L.G. R.L.G. NNP 10_1101-2020_12_31_424971 436 1 is be VBZ 10_1101-2020_12_31_424971 436 2 the the DT 10_1101-2020_12_31_424971 436 3 corresponding corresponding JJ 10_1101-2020_12_31_424971 436 4 1015 1015 CD 10_1101-2020_12_31_424971 436 5 author author NN 10_1101-2020_12_31_424971 436 6 . . . 10_1101-2020_12_31_424971 437 1 Therefore therefore RB 10_1101-2020_12_31_424971 437 2 , , , 10_1101-2020_12_31_424971 437 3 the the DT 10_1101-2020_12_31_424971 437 4 code code NN 10_1101-2020_12_31_424971 437 5 is be VBZ 10_1101-2020_12_31_424971 437 6 available available JJ 10_1101-2020_12_31_424971 437 7 from from IN 10_1101-2020_12_31_424971 437 8 R.L.G R.L.G NNP 10_1101-2020_12_31_424971 437 9 , , , 10_1101-2020_12_31_424971 437 10 upon upon IN 10_1101-2020_12_31_424971 437 11 request request NN 10_1101-2020_12_31_424971 437 12 . . . 10_1101-2020_12_31_424971 438 1 1016 1016 CD 10_1101-2020_12_31_424971 438 2 1017 1017 CD 10_1101-2020_12_31_424971 438 3 1018 1018 CD 10_1101-2020_12_31_424971 438 4 1019 1019 CD 10_1101-2020_12_31_424971 438 5 .CC .cc SYM 10_1101-2020_12_31_424971 438 6 - - HYPH 10_1101-2020_12_31_424971 438 7 BY by IN 10_1101-2020_12_31_424971 438 8 - - HYPH 10_1101-2020_12_31_424971 438 9 NC NC NNP 10_1101-2020_12_31_424971 438 10 - - HYPH 10_1101-2020_12_31_424971 438 11 ND ND NNP 10_1101-2020_12_31_424971 438 12 4.0 4.0 CD 10_1101-2020_12_31_424971 438 13 International International NNP 10_1101-2020_12_31_424971 438 14 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 438 15 under under IN 10_1101-2020_12_31_424971 438 16 a a DT 10_1101-2020_12_31_424971 438 17 ( ( -LRB- 10_1101-2020_12_31_424971 438 18 which which WDT 10_1101-2020_12_31_424971 438 19 was be VBD 10_1101-2020_12_31_424971 438 20 not not RB 10_1101-2020_12_31_424971 438 21 certified certify VBN 10_1101-2020_12_31_424971 438 22 by by IN 10_1101-2020_12_31_424971 438 23 peer peer NN 10_1101-2020_12_31_424971 438 24 review review NN 10_1101-2020_12_31_424971 438 25 ) ) -RRB- 10_1101-2020_12_31_424971 438 26 is be VBZ 10_1101-2020_12_31_424971 438 27 the the DT 10_1101-2020_12_31_424971 438 28 author author NN 10_1101-2020_12_31_424971 438 29 / / SYM 10_1101-2020_12_31_424971 438 30 funder funder NN 10_1101-2020_12_31_424971 438 31 , , , 10_1101-2020_12_31_424971 438 32 who who WP 10_1101-2020_12_31_424971 438 33 has have VBZ 10_1101-2020_12_31_424971 438 34 granted grant VBN 10_1101-2020_12_31_424971 438 35 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 438 36 a a DT 10_1101-2020_12_31_424971 438 37 license license NN 10_1101-2020_12_31_424971 438 38 to to TO 10_1101-2020_12_31_424971 438 39 display display VB 10_1101-2020_12_31_424971 438 40 the the DT 10_1101-2020_12_31_424971 438 41 preprint preprint NN 10_1101-2020_12_31_424971 438 42 in in IN 10_1101-2020_12_31_424971 438 43 perpetuity perpetuity NN 10_1101-2020_12_31_424971 438 44 . . . 10_1101-2020_12_31_424971 439 1 It -PRON- PRP 10_1101-2020_12_31_424971 439 2 is be VBZ 10_1101-2020_12_31_424971 439 3 made make VBN 10_1101-2020_12_31_424971 439 4 The the DT 10_1101-2020_12_31_424971 439 5 copyright copyright NN 10_1101-2020_12_31_424971 439 6 holder holder NN 10_1101-2020_12_31_424971 439 7 for for IN 10_1101-2020_12_31_424971 439 8 this this DT 10_1101-2020_12_31_424971 439 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 439 10 version version NN 10_1101-2020_12_31_424971 439 11 posted post VBD 10_1101-2020_12_31_424971 439 12 January January NNP 10_1101-2020_12_31_424971 439 13 2 2 CD 10_1101-2020_12_31_424971 439 14 , , , 10_1101-2020_12_31_424971 439 15 2021 2021 CD 10_1101-2020_12_31_424971 439 16 . . . 10_1101-2020_12_31_424971 439 17 ; ; : 10_1101-2020_12_31_424971 439 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 439 19 : : : 10_1101-2020_12_31_424971 439 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 439 21 preprint preprint NN 10_1101-2020_12_31_424971 439 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 439 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 439 24 42 42 CD 10_1101-2020_12_31_424971 439 25 REFERENCES reference NNS 10_1101-2020_12_31_424971 439 26 1020 1020 CD 10_1101-2020_12_31_424971 439 27 1 1 CD 10_1101-2020_12_31_424971 439 28 . . . 10_1101-2020_12_31_424971 440 1 Wang Wang NNP 10_1101-2020_12_31_424971 440 2 , , , 10_1101-2020_12_31_424971 440 3 K. K. NNP 10_1101-2020_12_31_424971 440 4 , , , 10_1101-2020_12_31_424971 440 5 Schmied Schmied NNP 10_1101-2020_12_31_424971 440 6 , , , 10_1101-2020_12_31_424971 440 7 W.H. W.H. NNP 10_1101-2020_12_31_424971 441 1 & & CC 10_1101-2020_12_31_424971 441 2 Chin Chin NNP 10_1101-2020_12_31_424971 441 3 , , , 10_1101-2020_12_31_424971 441 4 J.W. J.W. NNP 10_1101-2020_12_31_424971 442 1 Reprogramming reprogramme VBG 10_1101-2020_12_31_424971 442 2 the the DT 10_1101-2020_12_31_424971 442 3 genetic genetic JJ 10_1101-2020_12_31_424971 442 4 code code NN 10_1101-2020_12_31_424971 442 5 : : : 10_1101-2020_12_31_424971 442 6 from from IN 10_1101-2020_12_31_424971 442 7 triplet triplet NN 10_1101-2020_12_31_424971 442 8 to to IN 10_1101-2020_12_31_424971 442 9 1021 1021 CD 10_1101-2020_12_31_424971 442 10 quadruplet quadruplet NN 10_1101-2020_12_31_424971 442 11 codes code NNS 10_1101-2020_12_31_424971 442 12 . . . 10_1101-2020_12_31_424971 443 1 Angew angew VB 10_1101-2020_12_31_424971 443 2 Chem Chem NNP 10_1101-2020_12_31_424971 443 3 Int Int NNP 10_1101-2020_12_31_424971 443 4 Ed Ed NNP 10_1101-2020_12_31_424971 443 5 Engl Engl NNP 10_1101-2020_12_31_424971 443 6 51 51 CD 10_1101-2020_12_31_424971 443 7 , , , 10_1101-2020_12_31_424971 443 8 2288 2288 CD 10_1101-2020_12_31_424971 443 9 - - SYM 10_1101-2020_12_31_424971 443 10 97 97 CD 10_1101-2020_12_31_424971 443 11 ( ( -LRB- 10_1101-2020_12_31_424971 443 12 2012 2012 CD 10_1101-2020_12_31_424971 443 13 ) ) -RRB- 10_1101-2020_12_31_424971 443 14 . . . 10_1101-2020_12_31_424971 444 1 1022 1022 CD 10_1101-2020_12_31_424971 444 2 2 2 CD 10_1101-2020_12_31_424971 444 3 . . . 10_1101-2020_12_31_424971 445 1 Chen Chen NNP 10_1101-2020_12_31_424971 445 2 , , , 10_1101-2020_12_31_424971 445 3 Y. Y. NNP 10_1101-2020_12_31_424971 445 4 et et NNP 10_1101-2020_12_31_424971 445 5 al al NNP 10_1101-2020_12_31_424971 445 6 . . . 10_1101-2020_12_31_424971 446 1 Controlling control VBG 10_1101-2020_12_31_424971 446 2 the the DT 10_1101-2020_12_31_424971 446 3 Replication Replication NNP 10_1101-2020_12_31_424971 446 4 of of IN 10_1101-2020_12_31_424971 446 5 a a DT 10_1101-2020_12_31_424971 446 6 Genomically genomically RB 10_1101-2020_12_31_424971 446 7 Recoded recode VBN 10_1101-2020_12_31_424971 446 8 HIV-1 HIV-1 NNP 10_1101-2020_12_31_424971 446 9 with with IN 10_1101-2020_12_31_424971 446 10 a a DT 10_1101-2020_12_31_424971 446 11 1023 1023 CD 10_1101-2020_12_31_424971 446 12 Functional Functional NNP 10_1101-2020_12_31_424971 446 13 Quadruplet Quadruplet NNP 10_1101-2020_12_31_424971 446 14 Codon Codon NNP 10_1101-2020_12_31_424971 446 15 in in IN 10_1101-2020_12_31_424971 446 16 Mammalian Mammalian NNP 10_1101-2020_12_31_424971 446 17 Cells Cells NNPS 10_1101-2020_12_31_424971 446 18 . . . 10_1101-2020_12_31_424971 447 1 ACS ACS NNP 10_1101-2020_12_31_424971 447 2 Synth Synth NNP 10_1101-2020_12_31_424971 447 3 Biol Biol NNP 10_1101-2020_12_31_424971 447 4 7 7 CD 10_1101-2020_12_31_424971 447 5 , , , 10_1101-2020_12_31_424971 447 6 1612 1612 CD 10_1101-2020_12_31_424971 447 7 - - SYM 10_1101-2020_12_31_424971 447 8 1617 1617 CD 10_1101-2020_12_31_424971 447 9 ( ( -LRB- 10_1101-2020_12_31_424971 447 10 2018 2018 CD 10_1101-2020_12_31_424971 447 11 ) ) -RRB- 10_1101-2020_12_31_424971 447 12 . . . 10_1101-2020_12_31_424971 448 1 1024 1024 CD 10_1101-2020_12_31_424971 448 2 3 3 CD 10_1101-2020_12_31_424971 448 3 . . . 10_1101-2020_12_31_424971 449 1 Lee Lee NNP 10_1101-2020_12_31_424971 449 2 , , , 10_1101-2020_12_31_424971 449 3 B.S. B.S. NNP 10_1101-2020_12_31_424971 449 4 , , , 10_1101-2020_12_31_424971 449 5 Kim Kim NNP 10_1101-2020_12_31_424971 449 6 , , , 10_1101-2020_12_31_424971 449 7 S. S. NNP 10_1101-2020_12_31_424971 449 8 , , , 10_1101-2020_12_31_424971 449 9 Ko Ko NNP 10_1101-2020_12_31_424971 449 10 , , , 10_1101-2020_12_31_424971 449 11 B.J. B.J. NNP 10_1101-2020_12_31_424971 450 1 & & CC 10_1101-2020_12_31_424971 450 2 Yoo Yoo NNP 10_1101-2020_12_31_424971 450 3 , , , 10_1101-2020_12_31_424971 450 4 T.H. T.H. NNP 10_1101-2020_12_31_424971 451 1 An an DT 10_1101-2020_12_31_424971 451 2 efficient efficient JJ 10_1101-2020_12_31_424971 451 3 system system NN 10_1101-2020_12_31_424971 451 4 for for IN 10_1101-2020_12_31_424971 451 5 incorporation incorporation NN 10_1101-2020_12_31_424971 451 6 of of IN 10_1101-2020_12_31_424971 451 7 unnatural unnatural JJ 10_1101-2020_12_31_424971 451 8 1025 1025 CD 10_1101-2020_12_31_424971 451 9 amino amino JJ 10_1101-2020_12_31_424971 451 10 acids acid NNS 10_1101-2020_12_31_424971 451 11 in in IN 10_1101-2020_12_31_424971 451 12 response response NN 10_1101-2020_12_31_424971 451 13 to to IN 10_1101-2020_12_31_424971 451 14 the the DT 10_1101-2020_12_31_424971 451 15 four four CD 10_1101-2020_12_31_424971 451 16 - - HYPH 10_1101-2020_12_31_424971 451 17 base base NN 10_1101-2020_12_31_424971 451 18 codon codon NN 10_1101-2020_12_31_424971 451 19 AGGA AGGA NNP 10_1101-2020_12_31_424971 451 20 in in IN 10_1101-2020_12_31_424971 451 21 Escherichia Escherichia NNP 10_1101-2020_12_31_424971 451 22 coli coli NNS 10_1101-2020_12_31_424971 451 23 . . . 10_1101-2020_12_31_424971 452 1 Biochim Biochim NNP 10_1101-2020_12_31_424971 452 2 1026 1026 CD 10_1101-2020_12_31_424971 452 3 Biophys Biophys NNP 10_1101-2020_12_31_424971 452 4 Acta Acta NNP 10_1101-2020_12_31_424971 452 5 1861 1861 CD 10_1101-2020_12_31_424971 452 6 , , , 10_1101-2020_12_31_424971 452 7 3016 3016 CD 10_1101-2020_12_31_424971 452 8 - - SYM 10_1101-2020_12_31_424971 452 9 3023 3023 CD 10_1101-2020_12_31_424971 452 10 ( ( -LRB- 10_1101-2020_12_31_424971 452 11 2017 2017 CD 10_1101-2020_12_31_424971 452 12 ) ) -RRB- 10_1101-2020_12_31_424971 452 13 . . . 10_1101-2020_12_31_424971 453 1 1027 1027 CD 10_1101-2020_12_31_424971 453 2 4 4 CD 10_1101-2020_12_31_424971 453 3 . . . 10_1101-2020_12_31_424971 454 1 Chatterjee Chatterjee NNP 10_1101-2020_12_31_424971 454 2 , , , 10_1101-2020_12_31_424971 454 3 A. A. NNP 10_1101-2020_12_31_424971 454 4 , , , 10_1101-2020_12_31_424971 454 5 Lajoie Lajoie NNP 10_1101-2020_12_31_424971 454 6 , , , 10_1101-2020_12_31_424971 454 7 M.J. M.J. NNP 10_1101-2020_12_31_424971 454 8 , , , 10_1101-2020_12_31_424971 454 9 Xiao Xiao NNP 10_1101-2020_12_31_424971 454 10 , , , 10_1101-2020_12_31_424971 454 11 H. H. NNP 10_1101-2020_12_31_424971 454 12 , , , 10_1101-2020_12_31_424971 454 13 Church Church NNP 10_1101-2020_12_31_424971 454 14 , , , 10_1101-2020_12_31_424971 454 15 G.M. G.M. NNP 10_1101-2020_12_31_424971 455 1 & & CC 10_1101-2020_12_31_424971 455 2 Schultz Schultz NNP 10_1101-2020_12_31_424971 455 3 , , , 10_1101-2020_12_31_424971 455 4 P.G. P.G. NNP 10_1101-2020_12_31_424971 456 1 A a DT 10_1101-2020_12_31_424971 456 2 bacterial bacterial JJ 10_1101-2020_12_31_424971 456 3 strain strain NN 10_1101-2020_12_31_424971 456 4 with with IN 10_1101-2020_12_31_424971 456 5 1028 1028 CD 10_1101-2020_12_31_424971 456 6 a a DT 10_1101-2020_12_31_424971 456 7 unique unique JJ 10_1101-2020_12_31_424971 456 8 quadruplet quadruplet NN 10_1101-2020_12_31_424971 456 9 codon codon NN 10_1101-2020_12_31_424971 456 10 specifying specify VBG 10_1101-2020_12_31_424971 456 11 non non JJ 10_1101-2020_12_31_424971 456 12 - - JJ 10_1101-2020_12_31_424971 456 13 native native JJ 10_1101-2020_12_31_424971 456 14 amino amino NN 10_1101-2020_12_31_424971 456 15 acids acid NNS 10_1101-2020_12_31_424971 456 16 . . . 10_1101-2020_12_31_424971 457 1 Chembiochem Chembiochem NNP 10_1101-2020_12_31_424971 457 2 15 15 CD 10_1101-2020_12_31_424971 457 3 , , , 10_1101-2020_12_31_424971 457 4 1782 1782 CD 10_1101-2020_12_31_424971 457 5 - - SYM 10_1101-2020_12_31_424971 457 6 6 6 CD 10_1101-2020_12_31_424971 457 7 1029 1029 CD 10_1101-2020_12_31_424971 457 8 ( ( -LRB- 10_1101-2020_12_31_424971 457 9 2014 2014 CD 10_1101-2020_12_31_424971 457 10 ) ) -RRB- 10_1101-2020_12_31_424971 457 11 . . . 10_1101-2020_12_31_424971 458 1 1030 1030 CD 10_1101-2020_12_31_424971 458 2 5 5 CD 10_1101-2020_12_31_424971 458 3 . . . 10_1101-2020_12_31_424971 459 1 Niu niu UH 10_1101-2020_12_31_424971 459 2 , , , 10_1101-2020_12_31_424971 459 3 W. W. NNP 10_1101-2020_12_31_424971 459 4 , , , 10_1101-2020_12_31_424971 459 5 Schultz Schultz NNP 10_1101-2020_12_31_424971 459 6 , , , 10_1101-2020_12_31_424971 459 7 P.G. P.G. NNP 10_1101-2020_12_31_424971 460 1 & & CC 10_1101-2020_12_31_424971 460 2 Guo Guo NNP 10_1101-2020_12_31_424971 460 3 , , , 10_1101-2020_12_31_424971 460 4 J. J. NNP 10_1101-2020_12_31_424971 461 1 An an DT 10_1101-2020_12_31_424971 461 2 expanded expand VBN 10_1101-2020_12_31_424971 461 3 genetic genetic JJ 10_1101-2020_12_31_424971 461 4 code code NN 10_1101-2020_12_31_424971 461 5 in in IN 10_1101-2020_12_31_424971 461 6 mammalian mammalian JJ 10_1101-2020_12_31_424971 461 7 cells cell NNS 10_1101-2020_12_31_424971 461 8 with with IN 10_1101-2020_12_31_424971 461 9 a a DT 10_1101-2020_12_31_424971 461 10 1031 1031 CD 10_1101-2020_12_31_424971 461 11 functional functional JJ 10_1101-2020_12_31_424971 461 12 quadruplet quadruplet NN 10_1101-2020_12_31_424971 461 13 codon codon NNP 10_1101-2020_12_31_424971 461 14 . . . 10_1101-2020_12_31_424971 462 1 ACS ACS NNP 10_1101-2020_12_31_424971 462 2 Chem Chem NNP 10_1101-2020_12_31_424971 462 3 Biol Biol NNP 10_1101-2020_12_31_424971 462 4 8 8 CD 10_1101-2020_12_31_424971 462 5 , , , 10_1101-2020_12_31_424971 462 6 1640 1640 CD 10_1101-2020_12_31_424971 462 7 - - SYM 10_1101-2020_12_31_424971 462 8 5 5 CD 10_1101-2020_12_31_424971 462 9 ( ( -LRB- 10_1101-2020_12_31_424971 462 10 2013 2013 CD 10_1101-2020_12_31_424971 462 11 ) ) -RRB- 10_1101-2020_12_31_424971 462 12 . . . 10_1101-2020_12_31_424971 463 1 1032 1032 CD 10_1101-2020_12_31_424971 463 2 6 6 CD 10_1101-2020_12_31_424971 463 3 . . . 10_1101-2020_12_31_424971 464 1 Wang Wang NNP 10_1101-2020_12_31_424971 464 2 , , , 10_1101-2020_12_31_424971 464 3 N. N. NNP 10_1101-2020_12_31_424971 464 4 , , , 10_1101-2020_12_31_424971 464 5 Shang Shang NNP 10_1101-2020_12_31_424971 464 6 , , , 10_1101-2020_12_31_424971 464 7 X. X. NNP 10_1101-2020_12_31_424971 464 8 , , , 10_1101-2020_12_31_424971 464 9 Cerny Cerny NNP 10_1101-2020_12_31_424971 464 10 , , , 10_1101-2020_12_31_424971 464 11 R. R. NNP 10_1101-2020_12_31_424971 464 12 , , , 10_1101-2020_12_31_424971 464 13 Niu Niu NNP 10_1101-2020_12_31_424971 464 14 , , , 10_1101-2020_12_31_424971 464 15 W. W. NNP 10_1101-2020_12_31_424971 464 16 & & CC 10_1101-2020_12_31_424971 464 17 Guo Guo NNP 10_1101-2020_12_31_424971 464 18 , , , 10_1101-2020_12_31_424971 464 19 J. J. NNP 10_1101-2020_12_31_424971 465 1 Systematic Systematic NNP 10_1101-2020_12_31_424971 465 2 Evolution Evolution NNP 10_1101-2020_12_31_424971 465 3 and and CC 10_1101-2020_12_31_424971 465 4 Study Study NNP 10_1101-2020_12_31_424971 465 5 of of IN 10_1101-2020_12_31_424971 465 6 1033 1033 CD 10_1101-2020_12_31_424971 465 7 UAGN UAGN NNP 10_1101-2020_12_31_424971 465 8 Decoding Decoding NNP 10_1101-2020_12_31_424971 465 9 tRNAs trnas CD 10_1101-2020_12_31_424971 465 10 in in IN 10_1101-2020_12_31_424971 465 11 a a DT 10_1101-2020_12_31_424971 465 12 Genomically genomically RB 10_1101-2020_12_31_424971 465 13 Recoded recode VBN 10_1101-2020_12_31_424971 465 14 Bacteria Bacteria NNS 10_1101-2020_12_31_424971 465 15 . . . 10_1101-2020_12_31_424971 466 1 Sci Sci NNP 10_1101-2020_12_31_424971 466 2 Rep Rep NNP 10_1101-2020_12_31_424971 466 3 6 6 CD 10_1101-2020_12_31_424971 466 4 , , , 10_1101-2020_12_31_424971 466 5 21898 21898 CD 10_1101-2020_12_31_424971 466 6 ( ( -LRB- 10_1101-2020_12_31_424971 466 7 2016 2016 CD 10_1101-2020_12_31_424971 466 8 ) ) -RRB- 10_1101-2020_12_31_424971 466 9 . . . 10_1101-2020_12_31_424971 467 1 1034 1034 CD 10_1101-2020_12_31_424971 467 2 7 7 CD 10_1101-2020_12_31_424971 467 3 . . . 10_1101-2020_12_31_424971 468 1 Neumann Neumann NNP 10_1101-2020_12_31_424971 468 2 , , , 10_1101-2020_12_31_424971 468 3 H. H. NNP 10_1101-2020_12_31_424971 468 4 , , , 10_1101-2020_12_31_424971 468 5 Wang Wang NNP 10_1101-2020_12_31_424971 468 6 , , , 10_1101-2020_12_31_424971 468 7 K. K. NNP 10_1101-2020_12_31_424971 468 8 , , , 10_1101-2020_12_31_424971 468 9 Davis Davis NNP 10_1101-2020_12_31_424971 468 10 , , , 10_1101-2020_12_31_424971 468 11 L. L. NNP 10_1101-2020_12_31_424971 468 12 , , , 10_1101-2020_12_31_424971 468 13 Garcia Garcia NNP 10_1101-2020_12_31_424971 468 14 - - HYPH 10_1101-2020_12_31_424971 468 15 Alai Alai NNP 10_1101-2020_12_31_424971 468 16 , , , 10_1101-2020_12_31_424971 468 17 M. M. NNP 10_1101-2020_12_31_424971 468 18 & & CC 10_1101-2020_12_31_424971 468 19 Chin Chin NNP 10_1101-2020_12_31_424971 468 20 , , , 10_1101-2020_12_31_424971 468 21 J.W. J.W. NNP 10_1101-2020_12_31_424971 469 1 Encoding encode VBG 10_1101-2020_12_31_424971 469 2 multiple multiple JJ 10_1101-2020_12_31_424971 469 3 1035 1035 CD 10_1101-2020_12_31_424971 469 4 unnatural unnatural JJ 10_1101-2020_12_31_424971 469 5 amino amino NN 10_1101-2020_12_31_424971 469 6 acids acid NNS 10_1101-2020_12_31_424971 469 7 via via IN 10_1101-2020_12_31_424971 469 8 evolution evolution NN 10_1101-2020_12_31_424971 469 9 of of IN 10_1101-2020_12_31_424971 469 10 a a DT 10_1101-2020_12_31_424971 469 11 quadruplet quadruplet NN 10_1101-2020_12_31_424971 469 12 - - HYPH 10_1101-2020_12_31_424971 469 13 decoding decode VBG 10_1101-2020_12_31_424971 469 14 ribosome ribosome NN 10_1101-2020_12_31_424971 469 15 . . . 10_1101-2020_12_31_424971 470 1 Nature nature NN 10_1101-2020_12_31_424971 470 2 464 464 CD 10_1101-2020_12_31_424971 470 3 , , , 10_1101-2020_12_31_424971 470 4 1036 1036 CD 10_1101-2020_12_31_424971 470 5 441 441 CD 10_1101-2020_12_31_424971 470 6 - - SYM 10_1101-2020_12_31_424971 470 7 4 4 CD 10_1101-2020_12_31_424971 470 8 ( ( -LRB- 10_1101-2020_12_31_424971 470 9 2010 2010 CD 10_1101-2020_12_31_424971 470 10 ) ) -RRB- 10_1101-2020_12_31_424971 470 11 . . . 10_1101-2020_12_31_424971 471 1 1037 1037 CD 10_1101-2020_12_31_424971 471 2 8 8 CD 10_1101-2020_12_31_424971 471 3 . . . 10_1101-2020_12_31_424971 472 1 Wang Wang NNP 10_1101-2020_12_31_424971 472 2 , , , 10_1101-2020_12_31_424971 472 3 K. K. NNP 10_1101-2020_12_31_424971 472 4 et et NNP 10_1101-2020_12_31_424971 472 5 al al NNP 10_1101-2020_12_31_424971 472 6 . . . 10_1101-2020_12_31_424971 473 1 Optimized optimized JJ 10_1101-2020_12_31_424971 473 2 orthogonal orthogonal JJ 10_1101-2020_12_31_424971 473 3 translation translation NN 10_1101-2020_12_31_424971 473 4 of of IN 10_1101-2020_12_31_424971 473 5 unnatural unnatural JJ 10_1101-2020_12_31_424971 473 6 amino amino NN 10_1101-2020_12_31_424971 473 7 acids acid NNS 10_1101-2020_12_31_424971 473 8 enables enable VBZ 10_1101-2020_12_31_424971 473 9 1038 1038 CD 10_1101-2020_12_31_424971 473 10 spontaneous spontaneous JJ 10_1101-2020_12_31_424971 473 11 protein protein NN 10_1101-2020_12_31_424971 473 12 double double JJ 10_1101-2020_12_31_424971 473 13 - - HYPH 10_1101-2020_12_31_424971 473 14 labelling labelling NN 10_1101-2020_12_31_424971 473 15 and and CC 10_1101-2020_12_31_424971 473 16 FRET FRET NNS 10_1101-2020_12_31_424971 473 17 . . . 10_1101-2020_12_31_424971 474 1 Nat Nat NNP 10_1101-2020_12_31_424971 474 2 Chem Chem NNP 10_1101-2020_12_31_424971 474 3 6 6 CD 10_1101-2020_12_31_424971 474 4 , , , 10_1101-2020_12_31_424971 474 5 393 393 CD 10_1101-2020_12_31_424971 474 6 - - SYM 10_1101-2020_12_31_424971 474 7 403 403 CD 10_1101-2020_12_31_424971 474 8 ( ( -LRB- 10_1101-2020_12_31_424971 474 9 2014 2014 CD 10_1101-2020_12_31_424971 474 10 ) ) -RRB- 10_1101-2020_12_31_424971 474 11 . . . 10_1101-2020_12_31_424971 475 1 1039 1039 CD 10_1101-2020_12_31_424971 475 2 9 9 CD 10_1101-2020_12_31_424971 475 3 . . . 10_1101-2020_12_31_424971 476 1 Atkins Atkins NNP 10_1101-2020_12_31_424971 476 2 , , , 10_1101-2020_12_31_424971 476 3 J.F. J.F. NNP 10_1101-2020_12_31_424971 476 4 , , , 10_1101-2020_12_31_424971 476 5 Loughran Loughran NNP 10_1101-2020_12_31_424971 476 6 , , , 10_1101-2020_12_31_424971 476 7 G. G. NNP 10_1101-2020_12_31_424971 476 8 , , , 10_1101-2020_12_31_424971 476 9 Bhatt Bhatt NNP 10_1101-2020_12_31_424971 476 10 , , , 10_1101-2020_12_31_424971 476 11 P.R. P.R. NNP 10_1101-2020_12_31_424971 476 12 , , , 10_1101-2020_12_31_424971 476 13 Firth Firth NNP 10_1101-2020_12_31_424971 476 14 , , , 10_1101-2020_12_31_424971 476 15 A.E. A.E. NNP 10_1101-2020_12_31_424971 477 1 & & CC 10_1101-2020_12_31_424971 477 2 Baranov Baranov NNP 10_1101-2020_12_31_424971 477 3 , , , 10_1101-2020_12_31_424971 477 4 P.V. P.V. NNP 10_1101-2020_12_31_424971 478 1 Ribosomal Ribosomal NNP 10_1101-2020_12_31_424971 478 2 1040 1040 CD 10_1101-2020_12_31_424971 478 3 frameshifting frameshifting NN 10_1101-2020_12_31_424971 478 4 and and CC 10_1101-2020_12_31_424971 478 5 transcriptional transcriptional JJ 10_1101-2020_12_31_424971 478 6 slippage slippage NN 10_1101-2020_12_31_424971 478 7 : : : 10_1101-2020_12_31_424971 478 8 From from IN 10_1101-2020_12_31_424971 478 9 genetic genetic JJ 10_1101-2020_12_31_424971 478 10 steganography steganography NN 10_1101-2020_12_31_424971 478 11 and and CC 10_1101-2020_12_31_424971 478 12 1041 1041 CD 10_1101-2020_12_31_424971 478 13 cryptography cryptography NN 10_1101-2020_12_31_424971 478 14 to to IN 10_1101-2020_12_31_424971 478 15 adventitious adventitious JJ 10_1101-2020_12_31_424971 478 16 use use NN 10_1101-2020_12_31_424971 478 17 . . . 10_1101-2020_12_31_424971 479 1 Nucleic Nucleic NNP 10_1101-2020_12_31_424971 479 2 Acids Acids NNPS 10_1101-2020_12_31_424971 479 3 Res Res NNP 10_1101-2020_12_31_424971 479 4 44 44 CD 10_1101-2020_12_31_424971 479 5 , , , 10_1101-2020_12_31_424971 479 6 7007 7007 CD 10_1101-2020_12_31_424971 479 7 - - SYM 10_1101-2020_12_31_424971 479 8 78 78 CD 10_1101-2020_12_31_424971 479 9 ( ( -LRB- 10_1101-2020_12_31_424971 479 10 2016 2016 CD 10_1101-2020_12_31_424971 479 11 ) ) -RRB- 10_1101-2020_12_31_424971 479 12 . . . 10_1101-2020_12_31_424971 480 1 1042 1042 CD 10_1101-2020_12_31_424971 480 2 10 10 CD 10_1101-2020_12_31_424971 480 3 . . . 10_1101-2020_12_31_424971 481 1 Atkins Atkins NNP 10_1101-2020_12_31_424971 481 2 , , , 10_1101-2020_12_31_424971 481 3 J.F. J.F. NNP 10_1101-2020_12_31_424971 482 1 & & CC 10_1101-2020_12_31_424971 482 2 Bjork Bjork NNP 10_1101-2020_12_31_424971 482 3 , , , 10_1101-2020_12_31_424971 482 4 G.R. G.R. NNP 10_1101-2020_12_31_424971 483 1 A a DT 10_1101-2020_12_31_424971 483 2 gripping grip VBG 10_1101-2020_12_31_424971 483 3 tale tale NN 10_1101-2020_12_31_424971 483 4 of of IN 10_1101-2020_12_31_424971 483 5 ribosomal ribosomal NN 10_1101-2020_12_31_424971 483 6 frameshifting frameshifting NN 10_1101-2020_12_31_424971 483 7 : : : 10_1101-2020_12_31_424971 483 8 extragenic extragenic NNP 10_1101-2020_12_31_424971 483 9 1043 1043 CD 10_1101-2020_12_31_424971 483 10 suppressors suppressor NNS 10_1101-2020_12_31_424971 483 11 of of IN 10_1101-2020_12_31_424971 483 12 frameshift frameshift NN 10_1101-2020_12_31_424971 483 13 mutations mutation NNS 10_1101-2020_12_31_424971 483 14 spotlight spotlight NNP 10_1101-2020_12_31_424971 483 15 P P NNP 10_1101-2020_12_31_424971 483 16 - - HYPH 10_1101-2020_12_31_424971 483 17 site site NN 10_1101-2020_12_31_424971 483 18 realignment realignment NN 10_1101-2020_12_31_424971 483 19 . . . 10_1101-2020_12_31_424971 484 1 Microbiol Microbiol NNP 10_1101-2020_12_31_424971 484 2 Mol Mol NNP 10_1101-2020_12_31_424971 484 3 Biol Biol NNP 10_1101-2020_12_31_424971 484 4 Rev Rev NNP 10_1101-2020_12_31_424971 484 5 1044 1044 CD 10_1101-2020_12_31_424971 484 6 73 73 CD 10_1101-2020_12_31_424971 484 7 , , , 10_1101-2020_12_31_424971 484 8 178 178 CD 10_1101-2020_12_31_424971 484 9 - - HYPH 10_1101-2020_12_31_424971 484 10 210 210 CD 10_1101-2020_12_31_424971 484 11 ( ( -LRB- 10_1101-2020_12_31_424971 484 12 2009 2009 CD 10_1101-2020_12_31_424971 484 13 ) ) -RRB- 10_1101-2020_12_31_424971 484 14 . . . 10_1101-2020_12_31_424971 485 1 1045 1045 CD 10_1101-2020_12_31_424971 485 2 11 11 CD 10_1101-2020_12_31_424971 485 3 . . . 10_1101-2020_12_31_424971 486 1 Roth Roth NNP 10_1101-2020_12_31_424971 486 2 , , , 10_1101-2020_12_31_424971 486 3 J.R. J.R. NNP 10_1101-2020_12_31_424971 486 4 Frameshift Frameshift NNP 10_1101-2020_12_31_424971 486 5 suppression suppression NN 10_1101-2020_12_31_424971 486 6 . . . 10_1101-2020_12_31_424971 487 1 Cell Cell NNP 10_1101-2020_12_31_424971 487 2 24 24 CD 10_1101-2020_12_31_424971 487 3 , , , 10_1101-2020_12_31_424971 487 4 601 601 CD 10_1101-2020_12_31_424971 487 5 - - HYPH 10_1101-2020_12_31_424971 487 6 2 2 CD 10_1101-2020_12_31_424971 487 7 ( ( -LRB- 10_1101-2020_12_31_424971 487 8 1981 1981 CD 10_1101-2020_12_31_424971 487 9 ) ) -RRB- 10_1101-2020_12_31_424971 487 10 . . . 10_1101-2020_12_31_424971 488 1 1046 1046 CD 10_1101-2020_12_31_424971 488 2 12 12 CD 10_1101-2020_12_31_424971 488 3 . . . 10_1101-2020_12_31_424971 489 1 Bossi Bossi NNP 10_1101-2020_12_31_424971 489 2 , , , 10_1101-2020_12_31_424971 489 3 L. L. NNP 10_1101-2020_12_31_424971 489 4 & & CC 10_1101-2020_12_31_424971 489 5 Roth Roth NNP 10_1101-2020_12_31_424971 489 6 , , , 10_1101-2020_12_31_424971 489 7 J.R. J.R. NNP 10_1101-2020_12_31_424971 489 8 Four four CD 10_1101-2020_12_31_424971 489 9 - - HYPH 10_1101-2020_12_31_424971 489 10 base base NN 10_1101-2020_12_31_424971 489 11 codons codon NNS 10_1101-2020_12_31_424971 489 12 ACCA ACCA NNP 10_1101-2020_12_31_424971 489 13 , , , 10_1101-2020_12_31_424971 489 14 ACCU ACCU NNP 10_1101-2020_12_31_424971 489 15 and and CC 10_1101-2020_12_31_424971 489 16 ACCC ACCC NNP 10_1101-2020_12_31_424971 489 17 are be VBP 10_1101-2020_12_31_424971 489 18 recognized recognize VBN 10_1101-2020_12_31_424971 489 19 by by IN 10_1101-2020_12_31_424971 489 20 1047 1047 CD 10_1101-2020_12_31_424971 489 21 frameshift frameshift NN 10_1101-2020_12_31_424971 489 22 suppressor suppressor NN 10_1101-2020_12_31_424971 489 23 sufJ. sufJ. NNP 10_1101-2020_12_31_424971 490 1 Cell cell NN 10_1101-2020_12_31_424971 490 2 25 25 CD 10_1101-2020_12_31_424971 490 3 , , , 10_1101-2020_12_31_424971 490 4 489 489 CD 10_1101-2020_12_31_424971 490 5 - - SYM 10_1101-2020_12_31_424971 490 6 96 96 CD 10_1101-2020_12_31_424971 490 7 ( ( -LRB- 10_1101-2020_12_31_424971 490 8 1981 1981 CD 10_1101-2020_12_31_424971 490 9 ) ) -RRB- 10_1101-2020_12_31_424971 490 10 . . . 10_1101-2020_12_31_424971 491 1 1048 1048 CD 10_1101-2020_12_31_424971 491 2 13 13 CD 10_1101-2020_12_31_424971 491 3 . . . 10_1101-2020_12_31_424971 492 1 Qian Qian NNP 10_1101-2020_12_31_424971 492 2 , , , 10_1101-2020_12_31_424971 492 3 Q. Q. NNP 10_1101-2020_12_31_424971 492 4 et et NNP 10_1101-2020_12_31_424971 492 5 al al NNP 10_1101-2020_12_31_424971 492 6 . . . 10_1101-2020_12_31_424971 493 1 A a DT 10_1101-2020_12_31_424971 493 2 new new JJ 10_1101-2020_12_31_424971 493 3 model model NN 10_1101-2020_12_31_424971 493 4 for for IN 10_1101-2020_12_31_424971 493 5 phenotypic phenotypic JJ 10_1101-2020_12_31_424971 493 6 suppression suppression NN 10_1101-2020_12_31_424971 493 7 of of IN 10_1101-2020_12_31_424971 493 8 frameshift frameshift NN 10_1101-2020_12_31_424971 493 9 mutations mutation NNS 10_1101-2020_12_31_424971 493 10 by by IN 10_1101-2020_12_31_424971 493 11 1049 1049 CD 10_1101-2020_12_31_424971 493 12 mutant mutant JJ 10_1101-2020_12_31_424971 493 13 tRNAs tRNAs '' 10_1101-2020_12_31_424971 493 14 . . . 10_1101-2020_12_31_424971 494 1 Mol Mol NNP 10_1101-2020_12_31_424971 494 2 Cell Cell NNP 10_1101-2020_12_31_424971 494 3 1 1 CD 10_1101-2020_12_31_424971 494 4 , , , 10_1101-2020_12_31_424971 494 5 471 471 CD 10_1101-2020_12_31_424971 494 6 - - SYM 10_1101-2020_12_31_424971 494 7 82 82 CD 10_1101-2020_12_31_424971 494 8 ( ( -LRB- 10_1101-2020_12_31_424971 494 9 1998 1998 CD 10_1101-2020_12_31_424971 494 10 ) ) -RRB- 10_1101-2020_12_31_424971 494 11 . . . 10_1101-2020_12_31_424971 495 1 1050 1050 CD 10_1101-2020_12_31_424971 495 2 14 14 CD 10_1101-2020_12_31_424971 495 3 . . . 10_1101-2020_12_31_424971 496 1 Weiss Weiss NNP 10_1101-2020_12_31_424971 496 2 , , , 10_1101-2020_12_31_424971 496 3 R.B. R.B. NNP 10_1101-2020_12_31_424971 496 4 , , , 10_1101-2020_12_31_424971 496 5 Dunn Dunn NNP 10_1101-2020_12_31_424971 496 6 , , , 10_1101-2020_12_31_424971 496 7 D.M. D.M. NNP 10_1101-2020_12_31_424971 496 8 , , , 10_1101-2020_12_31_424971 496 9 Shuh Shuh NNP 10_1101-2020_12_31_424971 496 10 , , , 10_1101-2020_12_31_424971 496 11 M. M. NNP 10_1101-2020_12_31_424971 496 12 , , , 10_1101-2020_12_31_424971 496 13 Atkins Atkins NNP 10_1101-2020_12_31_424971 496 14 , , , 10_1101-2020_12_31_424971 496 15 J.F. J.F. NNP 10_1101-2020_12_31_424971 497 1 & & CC 10_1101-2020_12_31_424971 497 2 Gesteland Gesteland NNP 10_1101-2020_12_31_424971 497 3 , , , 10_1101-2020_12_31_424971 497 4 R.F. R.F. NNP 10_1101-2020_12_31_424971 498 1 E. E. NNP 10_1101-2020_12_31_424971 498 2 coli coli NNS 10_1101-2020_12_31_424971 498 3 ribosomes ribosome VBZ 10_1101-2020_12_31_424971 498 4 re-1051 re-1051 NNPS 10_1101-2020_12_31_424971 498 5 phase phase NN 10_1101-2020_12_31_424971 498 6 on on IN 10_1101-2020_12_31_424971 498 7 retroviral retroviral JJ 10_1101-2020_12_31_424971 498 8 frameshift frameshift NN 10_1101-2020_12_31_424971 498 9 signals signal NNS 10_1101-2020_12_31_424971 498 10 at at IN 10_1101-2020_12_31_424971 498 11 rates rate NNS 10_1101-2020_12_31_424971 498 12 ranging range VBG 10_1101-2020_12_31_424971 498 13 from from IN 10_1101-2020_12_31_424971 498 14 2 2 CD 10_1101-2020_12_31_424971 498 15 to to IN 10_1101-2020_12_31_424971 498 16 50 50 CD 10_1101-2020_12_31_424971 498 17 percent percent NN 10_1101-2020_12_31_424971 498 18 . . . 10_1101-2020_12_31_424971 499 1 New New NNP 10_1101-2020_12_31_424971 499 2 Biol Biol NNP 10_1101-2020_12_31_424971 499 3 1 1 CD 10_1101-2020_12_31_424971 499 4 , , , 10_1101-2020_12_31_424971 499 5 1052 1052 CD 10_1101-2020_12_31_424971 499 6 159 159 CD 10_1101-2020_12_31_424971 499 7 - - SYM 10_1101-2020_12_31_424971 499 8 69 69 CD 10_1101-2020_12_31_424971 499 9 ( ( -LRB- 10_1101-2020_12_31_424971 499 10 1989 1989 CD 10_1101-2020_12_31_424971 499 11 ) ) -RRB- 10_1101-2020_12_31_424971 499 12 . . . 10_1101-2020_12_31_424971 500 1 1053 1053 CD 10_1101-2020_12_31_424971 500 2 15 15 CD 10_1101-2020_12_31_424971 500 3 . . . 10_1101-2020_12_31_424971 501 1 Jager Jager NNP 10_1101-2020_12_31_424971 501 2 , , , 10_1101-2020_12_31_424971 501 3 G. G. NNP 10_1101-2020_12_31_424971 501 4 , , , 10_1101-2020_12_31_424971 501 5 Nilsson Nilsson NNP 10_1101-2020_12_31_424971 501 6 , , , 10_1101-2020_12_31_424971 501 7 K. K. NNP 10_1101-2020_12_31_424971 501 8 & & CC 10_1101-2020_12_31_424971 501 9 Bjork Bjork NNP 10_1101-2020_12_31_424971 501 10 , , , 10_1101-2020_12_31_424971 501 11 G.R. G.R. NNP 10_1101-2020_12_31_424971 502 1 The the DT 10_1101-2020_12_31_424971 502 2 phenotype phenotype NN 10_1101-2020_12_31_424971 502 3 of of IN 10_1101-2020_12_31_424971 502 4 many many JJ 10_1101-2020_12_31_424971 502 5 independently independently RB 10_1101-2020_12_31_424971 502 6 isolated isolate VBN 10_1101-2020_12_31_424971 502 7 +1 +1 NNS 10_1101-2020_12_31_424971 502 8 1054 1054 CD 10_1101-2020_12_31_424971 502 9 frameshift frameshift NN 10_1101-2020_12_31_424971 502 10 suppressor suppressor NN 10_1101-2020_12_31_424971 502 11 mutants mutant NNS 10_1101-2020_12_31_424971 502 12 supports support VBZ 10_1101-2020_12_31_424971 502 13 a a DT 10_1101-2020_12_31_424971 502 14 pivotal pivotal JJ 10_1101-2020_12_31_424971 502 15 role role NN 10_1101-2020_12_31_424971 502 16 of of IN 10_1101-2020_12_31_424971 502 17 the the DT 10_1101-2020_12_31_424971 502 18 P p NN 10_1101-2020_12_31_424971 502 19 - - HYPH 10_1101-2020_12_31_424971 502 20 site site NN 10_1101-2020_12_31_424971 502 21 in in IN 10_1101-2020_12_31_424971 502 22 reading read VBG 10_1101-2020_12_31_424971 502 23 frame frame NN 10_1101-2020_12_31_424971 502 24 1055 1055 CD 10_1101-2020_12_31_424971 502 25 maintenance maintenance NN 10_1101-2020_12_31_424971 502 26 . . . 10_1101-2020_12_31_424971 503 1 PLoS plos ADD 10_1101-2020_12_31_424971 503 2 One one CD 10_1101-2020_12_31_424971 503 3 8 8 CD 10_1101-2020_12_31_424971 503 4 , , , 10_1101-2020_12_31_424971 503 5 e60246 e60246 NNP 10_1101-2020_12_31_424971 503 6 ( ( -LRB- 10_1101-2020_12_31_424971 503 7 2013 2013 CD 10_1101-2020_12_31_424971 503 8 ) ) -RRB- 10_1101-2020_12_31_424971 503 9 . . . 10_1101-2020_12_31_424971 504 1 1056 1056 CD 10_1101-2020_12_31_424971 504 2 16 16 CD 10_1101-2020_12_31_424971 504 3 . . . 10_1101-2020_12_31_424971 505 1 Fagan Fagan NNP 10_1101-2020_12_31_424971 505 2 , , , 10_1101-2020_12_31_424971 505 3 C.E. C.E. NNP 10_1101-2020_12_31_424971 505 4 , , , 10_1101-2020_12_31_424971 505 5 Maehigashi Maehigashi NNP 10_1101-2020_12_31_424971 505 6 , , , 10_1101-2020_12_31_424971 505 7 T. T. NNP 10_1101-2020_12_31_424971 505 8 , , , 10_1101-2020_12_31_424971 505 9 Dunkle Dunkle NNP 10_1101-2020_12_31_424971 505 10 , , , 10_1101-2020_12_31_424971 505 11 J.A. J.A. NNP 10_1101-2020_12_31_424971 505 12 , , , 10_1101-2020_12_31_424971 505 13 Miles Miles NNP 10_1101-2020_12_31_424971 505 14 , , , 10_1101-2020_12_31_424971 505 15 S.J. S.J. NNP 10_1101-2020_12_31_424971 506 1 & & CC 10_1101-2020_12_31_424971 506 2 Dunham Dunham NNP 10_1101-2020_12_31_424971 506 3 , , , 10_1101-2020_12_31_424971 506 4 C.M. C.M. NNP 10_1101-2020_12_31_424971 507 1 Structural structural JJ 10_1101-2020_12_31_424971 507 2 1057 1057 CD 10_1101-2020_12_31_424971 507 3 insights insight NNS 10_1101-2020_12_31_424971 507 4 into into IN 10_1101-2020_12_31_424971 507 5 translational translational JJ 10_1101-2020_12_31_424971 507 6 recoding recoding NN 10_1101-2020_12_31_424971 507 7 by by IN 10_1101-2020_12_31_424971 507 8 frameshift frameshift NN 10_1101-2020_12_31_424971 507 9 suppressor suppressor NN 10_1101-2020_12_31_424971 507 10 tRNASufJ. trnasufj. NN 10_1101-2020_12_31_424971 508 1 RNA RNA NNP 10_1101-2020_12_31_424971 508 2 20 20 CD 10_1101-2020_12_31_424971 508 3 , , , 10_1101-2020_12_31_424971 508 4 1944 1944 CD 10_1101-2020_12_31_424971 508 5 - - SYM 10_1101-2020_12_31_424971 508 6 54 54 CD 10_1101-2020_12_31_424971 508 7 1058 1058 CD 10_1101-2020_12_31_424971 508 8 ( ( -LRB- 10_1101-2020_12_31_424971 508 9 2014 2014 CD 10_1101-2020_12_31_424971 508 10 ) ) -RRB- 10_1101-2020_12_31_424971 508 11 . . . 10_1101-2020_12_31_424971 509 1 1059 1059 CD 10_1101-2020_12_31_424971 509 2 17 17 CD 10_1101-2020_12_31_424971 509 3 . . . 10_1101-2020_12_31_424971 510 1 Maehigashi Maehigashi NNP 10_1101-2020_12_31_424971 510 2 , , , 10_1101-2020_12_31_424971 510 3 T. T. NNP 10_1101-2020_12_31_424971 510 4 , , , 10_1101-2020_12_31_424971 510 5 Dunkle Dunkle NNP 10_1101-2020_12_31_424971 510 6 , , , 10_1101-2020_12_31_424971 510 7 J.A. J.A. NNP 10_1101-2020_12_31_424971 510 8 , , , 10_1101-2020_12_31_424971 510 9 Miles Miles NNP 10_1101-2020_12_31_424971 510 10 , , , 10_1101-2020_12_31_424971 510 11 S.J. S.J. NNP 10_1101-2020_12_31_424971 511 1 & & CC 10_1101-2020_12_31_424971 511 2 Dunham Dunham NNP 10_1101-2020_12_31_424971 511 3 , , , 10_1101-2020_12_31_424971 511 4 C.M. C.M. NNP 10_1101-2020_12_31_424971 512 1 Structural structural JJ 10_1101-2020_12_31_424971 512 2 insights insight NNS 10_1101-2020_12_31_424971 512 3 into into IN 10_1101-2020_12_31_424971 512 4 +1 +1 NNS 10_1101-2020_12_31_424971 512 5 1060 1060 CD 10_1101-2020_12_31_424971 512 6 frameshifting frameshifting NN 10_1101-2020_12_31_424971 512 7 promoted promote VBN 10_1101-2020_12_31_424971 512 8 by by IN 10_1101-2020_12_31_424971 512 9 expanded expanded JJ 10_1101-2020_12_31_424971 512 10 or or CC 10_1101-2020_12_31_424971 512 11 modification modification NN 10_1101-2020_12_31_424971 512 12 - - HYPH 10_1101-2020_12_31_424971 512 13 deficient deficient JJ 10_1101-2020_12_31_424971 512 14 anticodon anticodon NN 10_1101-2020_12_31_424971 512 15 stem stem NN 10_1101-2020_12_31_424971 512 16 loops loop NNS 10_1101-2020_12_31_424971 512 17 . . . 10_1101-2020_12_31_424971 513 1 1061 1061 CD 10_1101-2020_12_31_424971 513 2 Proc Proc NNP 10_1101-2020_12_31_424971 513 3 Natl Natl NNP 10_1101-2020_12_31_424971 513 4 Acad Acad NNP 10_1101-2020_12_31_424971 513 5 Sci Sci NNP 10_1101-2020_12_31_424971 513 6 U U NNP 10_1101-2020_12_31_424971 513 7 S S NNP 10_1101-2020_12_31_424971 513 8 A A NNP 10_1101-2020_12_31_424971 513 9 111 111 CD 10_1101-2020_12_31_424971 513 10 , , , 10_1101-2020_12_31_424971 513 11 12740 12740 CD 10_1101-2020_12_31_424971 513 12 - - SYM 10_1101-2020_12_31_424971 513 13 5 5 CD 10_1101-2020_12_31_424971 513 14 ( ( -LRB- 10_1101-2020_12_31_424971 513 15 2014 2014 CD 10_1101-2020_12_31_424971 513 16 ) ) -RRB- 10_1101-2020_12_31_424971 513 17 . . . 10_1101-2020_12_31_424971 514 1 1062 1062 CD 10_1101-2020_12_31_424971 514 2 18 18 CD 10_1101-2020_12_31_424971 514 3 . . . 10_1101-2020_12_31_424971 515 1 Dunham Dunham NNP 10_1101-2020_12_31_424971 515 2 , , , 10_1101-2020_12_31_424971 515 3 C.M. C.M. NNP 10_1101-2020_12_31_424971 516 1 et et NNP 10_1101-2020_12_31_424971 516 2 al al NNP 10_1101-2020_12_31_424971 516 3 . . . 10_1101-2020_12_31_424971 517 1 Structures structure NNS 10_1101-2020_12_31_424971 517 2 of of IN 10_1101-2020_12_31_424971 517 3 tRNAs trnas CD 10_1101-2020_12_31_424971 517 4 with with IN 10_1101-2020_12_31_424971 517 5 an an DT 10_1101-2020_12_31_424971 517 6 expanded expand VBN 10_1101-2020_12_31_424971 517 7 anticodon anticodon NNP 10_1101-2020_12_31_424971 517 8 loop loop NN 10_1101-2020_12_31_424971 517 9 in in IN 10_1101-2020_12_31_424971 517 10 the the DT 10_1101-2020_12_31_424971 517 11 1063 1063 CD 10_1101-2020_12_31_424971 517 12 decoding decode VBG 10_1101-2020_12_31_424971 517 13 center center NN 10_1101-2020_12_31_424971 517 14 of of IN 10_1101-2020_12_31_424971 517 15 the the DT 10_1101-2020_12_31_424971 517 16 30S 30s CD 10_1101-2020_12_31_424971 517 17 ribosomal ribosomal NN 10_1101-2020_12_31_424971 517 18 subunit subunit NN 10_1101-2020_12_31_424971 517 19 . . . 10_1101-2020_12_31_424971 518 1 RNA RNA NNP 10_1101-2020_12_31_424971 518 2 13 13 CD 10_1101-2020_12_31_424971 518 3 , , , 10_1101-2020_12_31_424971 518 4 817 817 CD 10_1101-2020_12_31_424971 518 5 - - SYM 10_1101-2020_12_31_424971 518 6 23 23 CD 10_1101-2020_12_31_424971 518 7 ( ( -LRB- 10_1101-2020_12_31_424971 518 8 2007 2007 CD 10_1101-2020_12_31_424971 518 9 ) ) -RRB- 10_1101-2020_12_31_424971 518 10 . . . 10_1101-2020_12_31_424971 519 1 1064 1064 CD 10_1101-2020_12_31_424971 519 2 19 19 CD 10_1101-2020_12_31_424971 519 3 . . . 10_1101-2020_12_31_424971 520 1 Hong Hong NNP 10_1101-2020_12_31_424971 520 2 , , , 10_1101-2020_12_31_424971 520 3 S. S. NNP 10_1101-2020_12_31_424971 520 4 et et NNP 10_1101-2020_12_31_424971 520 5 al al NNP 10_1101-2020_12_31_424971 520 6 . . . 10_1101-2020_12_31_424971 521 1 Mechanism mechanism NN 10_1101-2020_12_31_424971 521 2 of of IN 10_1101-2020_12_31_424971 521 3 tRNA trna NN 10_1101-2020_12_31_424971 521 4 - - HYPH 10_1101-2020_12_31_424971 521 5 mediated mediate VBN 10_1101-2020_12_31_424971 521 6 +1 +1 NNS 10_1101-2020_12_31_424971 521 7 ribosomal ribosomal NN 10_1101-2020_12_31_424971 521 8 frameshifting frameshifting NN 10_1101-2020_12_31_424971 521 9 . . . 10_1101-2020_12_31_424971 522 1 Proc Proc NNP 10_1101-2020_12_31_424971 522 2 Natl Natl NNP 10_1101-2020_12_31_424971 522 3 Acad Acad NNP 10_1101-2020_12_31_424971 522 4 1065 1065 CD 10_1101-2020_12_31_424971 522 5 Sci Sci NNP 10_1101-2020_12_31_424971 522 6 U U NNP 10_1101-2020_12_31_424971 522 7 S S NNP 10_1101-2020_12_31_424971 522 8 A A NNP 10_1101-2020_12_31_424971 522 9 115 115 CD 10_1101-2020_12_31_424971 522 10 , , , 10_1101-2020_12_31_424971 522 11 11226 11226 CD 10_1101-2020_12_31_424971 522 12 - - SYM 10_1101-2020_12_31_424971 522 13 11231 11231 CD 10_1101-2020_12_31_424971 522 14 ( ( -LRB- 10_1101-2020_12_31_424971 522 15 2018 2018 CD 10_1101-2020_12_31_424971 522 16 ) ) -RRB- 10_1101-2020_12_31_424971 522 17 . . . 10_1101-2020_12_31_424971 523 1 1066 1066 CD 10_1101-2020_12_31_424971 523 2 20 20 CD 10_1101-2020_12_31_424971 523 3 . . . 10_1101-2020_12_31_424971 524 1 Sroga Sroga NNP 10_1101-2020_12_31_424971 524 2 , , , 10_1101-2020_12_31_424971 524 3 G.E. G.E. NNP 10_1101-2020_12_31_424971 524 4 , , , 10_1101-2020_12_31_424971 524 5 Nemoto Nemoto NNP 10_1101-2020_12_31_424971 524 6 , , , 10_1101-2020_12_31_424971 524 7 F. F. NNP 10_1101-2020_12_31_424971 524 8 , , , 10_1101-2020_12_31_424971 524 9 Kuchino Kuchino NNP 10_1101-2020_12_31_424971 524 10 , , , 10_1101-2020_12_31_424971 524 11 Y. Y. NNP 10_1101-2020_12_31_424971 525 1 & & CC 10_1101-2020_12_31_424971 525 2 Bjork Bjork NNP 10_1101-2020_12_31_424971 525 3 , , , 10_1101-2020_12_31_424971 525 4 G.R. G.R. NNP 10_1101-2020_12_31_424971 526 1 Insertion insertion NN 10_1101-2020_12_31_424971 526 2 ( ( -LRB- 10_1101-2020_12_31_424971 526 3 sufB sufB NNP 10_1101-2020_12_31_424971 526 4 ) ) -RRB- 10_1101-2020_12_31_424971 526 5 in in IN 10_1101-2020_12_31_424971 526 6 the the DT 10_1101-2020_12_31_424971 526 7 anticodon anticodon NN 10_1101-2020_12_31_424971 526 8 loop loop NN 10_1101-2020_12_31_424971 526 9 1067 1067 CD 10_1101-2020_12_31_424971 526 10 or or CC 10_1101-2020_12_31_424971 526 11 base base NN 10_1101-2020_12_31_424971 526 12 substitution substitution NN 10_1101-2020_12_31_424971 526 13 ( ( -LRB- 10_1101-2020_12_31_424971 526 14 sufC sufC NFP 10_1101-2020_12_31_424971 526 15 ) ) -RRB- 10_1101-2020_12_31_424971 526 16 in in IN 10_1101-2020_12_31_424971 526 17 the the DT 10_1101-2020_12_31_424971 526 18 anticodon anticodon NN 10_1101-2020_12_31_424971 526 19 stem stem NN 10_1101-2020_12_31_424971 526 20 of of IN 10_1101-2020_12_31_424971 526 21 tRNA(Pro)2 tRNA(Pro)2 NNP 10_1101-2020_12_31_424971 526 22 from from IN 10_1101-2020_12_31_424971 526 23 Salmonella salmonella NN 10_1101-2020_12_31_424971 526 24 1068 1068 CD 10_1101-2020_12_31_424971 526 25 .CC .cc SYM 10_1101-2020_12_31_424971 526 26 - - : 10_1101-2020_12_31_424971 526 27 BY by IN 10_1101-2020_12_31_424971 526 28 - - HYPH 10_1101-2020_12_31_424971 526 29 NC NC NNP 10_1101-2020_12_31_424971 526 30 - - HYPH 10_1101-2020_12_31_424971 526 31 ND ND NNP 10_1101-2020_12_31_424971 526 32 4.0 4.0 CD 10_1101-2020_12_31_424971 526 33 International International NNP 10_1101-2020_12_31_424971 526 34 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 526 35 under under IN 10_1101-2020_12_31_424971 526 36 a a DT 10_1101-2020_12_31_424971 526 37 ( ( -LRB- 10_1101-2020_12_31_424971 526 38 which which WDT 10_1101-2020_12_31_424971 526 39 was be VBD 10_1101-2020_12_31_424971 526 40 not not RB 10_1101-2020_12_31_424971 526 41 certified certify VBN 10_1101-2020_12_31_424971 526 42 by by IN 10_1101-2020_12_31_424971 526 43 peer peer NN 10_1101-2020_12_31_424971 526 44 review review NN 10_1101-2020_12_31_424971 526 45 ) ) -RRB- 10_1101-2020_12_31_424971 526 46 is be VBZ 10_1101-2020_12_31_424971 526 47 the the DT 10_1101-2020_12_31_424971 526 48 author author NN 10_1101-2020_12_31_424971 526 49 / / SYM 10_1101-2020_12_31_424971 526 50 funder funder NN 10_1101-2020_12_31_424971 526 51 , , , 10_1101-2020_12_31_424971 526 52 who who WP 10_1101-2020_12_31_424971 526 53 has have VBZ 10_1101-2020_12_31_424971 526 54 granted grant VBN 10_1101-2020_12_31_424971 526 55 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 526 56 a a DT 10_1101-2020_12_31_424971 526 57 license license NN 10_1101-2020_12_31_424971 526 58 to to TO 10_1101-2020_12_31_424971 526 59 display display VB 10_1101-2020_12_31_424971 526 60 the the DT 10_1101-2020_12_31_424971 526 61 preprint preprint NN 10_1101-2020_12_31_424971 526 62 in in IN 10_1101-2020_12_31_424971 526 63 perpetuity perpetuity NN 10_1101-2020_12_31_424971 526 64 . . . 10_1101-2020_12_31_424971 527 1 It -PRON- PRP 10_1101-2020_12_31_424971 527 2 is be VBZ 10_1101-2020_12_31_424971 527 3 made make VBN 10_1101-2020_12_31_424971 527 4 The the DT 10_1101-2020_12_31_424971 527 5 copyright copyright NN 10_1101-2020_12_31_424971 527 6 holder holder NN 10_1101-2020_12_31_424971 527 7 for for IN 10_1101-2020_12_31_424971 527 8 this this DT 10_1101-2020_12_31_424971 527 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 527 10 version version NN 10_1101-2020_12_31_424971 527 11 posted post VBD 10_1101-2020_12_31_424971 527 12 January January NNP 10_1101-2020_12_31_424971 527 13 2 2 CD 10_1101-2020_12_31_424971 527 14 , , , 10_1101-2020_12_31_424971 527 15 2021 2021 CD 10_1101-2020_12_31_424971 527 16 . . . 10_1101-2020_12_31_424971 527 17 ; ; : 10_1101-2020_12_31_424971 527 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 527 19 : : : 10_1101-2020_12_31_424971 527 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 527 21 preprint preprint NN 10_1101-2020_12_31_424971 527 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 527 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 527 24 43 43 CD 10_1101-2020_12_31_424971 527 25 typhimurium typhimurium NN 10_1101-2020_12_31_424971 527 26 induces induce VBZ 10_1101-2020_12_31_424971 527 27 suppression suppression NN 10_1101-2020_12_31_424971 527 28 of of IN 10_1101-2020_12_31_424971 527 29 frameshift frameshift NN 10_1101-2020_12_31_424971 527 30 mutations mutation NNS 10_1101-2020_12_31_424971 527 31 . . . 10_1101-2020_12_31_424971 528 1 Nucleic Nucleic NNP 10_1101-2020_12_31_424971 528 2 Acids Acids NNPS 10_1101-2020_12_31_424971 528 3 Res Res NNP 10_1101-2020_12_31_424971 528 4 20 20 CD 10_1101-2020_12_31_424971 528 5 , , , 10_1101-2020_12_31_424971 528 6 3463 3463 CD 10_1101-2020_12_31_424971 528 7 - - SYM 10_1101-2020_12_31_424971 528 8 9 9 CD 10_1101-2020_12_31_424971 528 9 1069 1069 CD 10_1101-2020_12_31_424971 528 10 ( ( -LRB- 10_1101-2020_12_31_424971 528 11 1992 1992 CD 10_1101-2020_12_31_424971 528 12 ) ) -RRB- 10_1101-2020_12_31_424971 528 13 . . . 10_1101-2020_12_31_424971 529 1 1070 1070 CD 10_1101-2020_12_31_424971 529 2 21 21 CD 10_1101-2020_12_31_424971 529 3 . . . 10_1101-2020_12_31_424971 530 1 Caliskan Caliskan NNP 10_1101-2020_12_31_424971 530 2 , , , 10_1101-2020_12_31_424971 530 3 N. N. NNP 10_1101-2020_12_31_424971 530 4 , , , 10_1101-2020_12_31_424971 530 5 Katunin Katunin NNP 10_1101-2020_12_31_424971 530 6 , , , 10_1101-2020_12_31_424971 530 7 V.I. V.I. NNP 10_1101-2020_12_31_424971 530 8 , , , 10_1101-2020_12_31_424971 530 9 Belardinelli Belardinelli NNP 10_1101-2020_12_31_424971 530 10 , , , 10_1101-2020_12_31_424971 530 11 R. R. NNP 10_1101-2020_12_31_424971 530 12 , , , 10_1101-2020_12_31_424971 530 13 Peske Peske NNP 10_1101-2020_12_31_424971 530 14 , , , 10_1101-2020_12_31_424971 530 15 F. F. NNP 10_1101-2020_12_31_424971 530 16 & & CC 10_1101-2020_12_31_424971 530 17 Rodnina Rodnina NNP 10_1101-2020_12_31_424971 530 18 , , , 10_1101-2020_12_31_424971 530 19 M.V. M.V. NNP 10_1101-2020_12_31_424971 531 1 Programmed Programmed NNP 10_1101-2020_12_31_424971 531 2 -1 -1 NNP 10_1101-2020_12_31_424971 531 3 1071 1071 CD 10_1101-2020_12_31_424971 531 4 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 531 5 by by IN 10_1101-2020_12_31_424971 531 6 kinetic kinetic NN 10_1101-2020_12_31_424971 531 7 partitioning partitioning NN 10_1101-2020_12_31_424971 531 8 during during IN 10_1101-2020_12_31_424971 531 9 impeded impeded JJ 10_1101-2020_12_31_424971 531 10 translocation translocation NN 10_1101-2020_12_31_424971 531 11 . . . 10_1101-2020_12_31_424971 532 1 Cell Cell NNP 10_1101-2020_12_31_424971 532 2 157 157 CD 10_1101-2020_12_31_424971 532 3 , , , 10_1101-2020_12_31_424971 532 4 1619 1619 CD 10_1101-2020_12_31_424971 532 5 - - SYM 10_1101-2020_12_31_424971 532 6 31 31 CD 10_1101-2020_12_31_424971 532 7 1072 1072 CD 10_1101-2020_12_31_424971 532 8 ( ( -LRB- 10_1101-2020_12_31_424971 532 9 2014 2014 CD 10_1101-2020_12_31_424971 532 10 ) ) -RRB- 10_1101-2020_12_31_424971 532 11 . . . 10_1101-2020_12_31_424971 533 1 1073 1073 CD 10_1101-2020_12_31_424971 533 2 22 22 CD 10_1101-2020_12_31_424971 533 3 . . . 10_1101-2020_12_31_424971 534 1 Taylor Taylor NNP 10_1101-2020_12_31_424971 534 2 , , , 10_1101-2020_12_31_424971 534 3 D.J. D.J. NNP 10_1101-2020_12_31_424971 535 1 et et NNP 10_1101-2020_12_31_424971 535 2 al al NNP 10_1101-2020_12_31_424971 535 3 . . . 10_1101-2020_12_31_424971 536 1 Structures structure NNS 10_1101-2020_12_31_424971 536 2 of of IN 10_1101-2020_12_31_424971 536 3 modified modify VBN 10_1101-2020_12_31_424971 536 4 eEF2 eEF2 NNP 10_1101-2020_12_31_424971 536 5 80S 80S NNP 10_1101-2020_12_31_424971 536 6 ribosome ribosome VBD 10_1101-2020_12_31_424971 536 7 complexes complex NNS 10_1101-2020_12_31_424971 536 8 reveal reveal VBP 10_1101-2020_12_31_424971 536 9 the the DT 10_1101-2020_12_31_424971 536 10 role role NN 10_1101-2020_12_31_424971 536 11 1074 1074 CD 10_1101-2020_12_31_424971 536 12 of of IN 10_1101-2020_12_31_424971 536 13 GTP GTP NNP 10_1101-2020_12_31_424971 536 14 hydrolysis hydrolysis NN 10_1101-2020_12_31_424971 536 15 in in IN 10_1101-2020_12_31_424971 536 16 translocation translocation NN 10_1101-2020_12_31_424971 536 17 . . . 10_1101-2020_12_31_424971 537 1 EMBO EMBO NNP 10_1101-2020_12_31_424971 537 2 J J NNP 10_1101-2020_12_31_424971 537 3 26 26 CD 10_1101-2020_12_31_424971 537 4 , , , 10_1101-2020_12_31_424971 537 5 2421 2421 CD 10_1101-2020_12_31_424971 537 6 - - SYM 10_1101-2020_12_31_424971 537 7 31 31 CD 10_1101-2020_12_31_424971 537 8 ( ( -LRB- 10_1101-2020_12_31_424971 537 9 2007 2007 CD 10_1101-2020_12_31_424971 537 10 ) ) -RRB- 10_1101-2020_12_31_424971 537 11 . . . 10_1101-2020_12_31_424971 538 1 1075 1075 CD 10_1101-2020_12_31_424971 538 2 23 23 CD 10_1101-2020_12_31_424971 538 3 . . . 10_1101-2020_12_31_424971 539 1 Khade Khade NNP 10_1101-2020_12_31_424971 539 2 , , , 10_1101-2020_12_31_424971 539 3 P.K. P.K. NNP 10_1101-2020_12_31_424971 540 1 & & CC 10_1101-2020_12_31_424971 540 2 Joseph Joseph NNP 10_1101-2020_12_31_424971 540 3 , , , 10_1101-2020_12_31_424971 540 4 S. S. NNP 10_1101-2020_12_31_424971 540 5 Messenger Messenger NNP 10_1101-2020_12_31_424971 540 6 RNA RNA NNP 10_1101-2020_12_31_424971 540 7 interactions interaction NNS 10_1101-2020_12_31_424971 540 8 in in IN 10_1101-2020_12_31_424971 540 9 the the DT 10_1101-2020_12_31_424971 540 10 decoding decode VBG 10_1101-2020_12_31_424971 540 11 center center NN 10_1101-2020_12_31_424971 540 12 control control NN 10_1101-2020_12_31_424971 540 13 1076 1076 CD 10_1101-2020_12_31_424971 540 14 the the DT 10_1101-2020_12_31_424971 540 15 rate rate NN 10_1101-2020_12_31_424971 540 16 of of IN 10_1101-2020_12_31_424971 540 17 translocation translocation NN 10_1101-2020_12_31_424971 540 18 . . . 10_1101-2020_12_31_424971 541 1 Nat Nat NNP 10_1101-2020_12_31_424971 541 2 Struct Struct NNP 10_1101-2020_12_31_424971 541 3 Mol Mol NNP 10_1101-2020_12_31_424971 541 4 Biol Biol NNP 10_1101-2020_12_31_424971 541 5 18 18 CD 10_1101-2020_12_31_424971 541 6 , , , 10_1101-2020_12_31_424971 541 7 1300 1300 CD 10_1101-2020_12_31_424971 541 8 - - SYM 10_1101-2020_12_31_424971 541 9 2 2 CD 10_1101-2020_12_31_424971 541 10 ( ( -LRB- 10_1101-2020_12_31_424971 541 11 2011 2011 CD 10_1101-2020_12_31_424971 541 12 ) ) -RRB- 10_1101-2020_12_31_424971 541 13 . . . 10_1101-2020_12_31_424971 542 1 1077 1077 CD 10_1101-2020_12_31_424971 542 2 24 24 CD 10_1101-2020_12_31_424971 542 3 . . . 10_1101-2020_12_31_424971 543 1 Liu Liu NNP 10_1101-2020_12_31_424971 543 2 , , , 10_1101-2020_12_31_424971 543 3 G. G. NNP 10_1101-2020_12_31_424971 543 4 et et NNP 10_1101-2020_12_31_424971 543 5 al al NNP 10_1101-2020_12_31_424971 543 6 . . . 10_1101-2020_12_31_424971 544 1 EF EF NNP 10_1101-2020_12_31_424971 544 2 - - HYPH 10_1101-2020_12_31_424971 544 3 G G NNP 10_1101-2020_12_31_424971 544 4 catalyzes catalyze VBZ 10_1101-2020_12_31_424971 544 5 tRNA trna NN 10_1101-2020_12_31_424971 544 6 translocation translocation NN 10_1101-2020_12_31_424971 544 7 by by IN 10_1101-2020_12_31_424971 544 8 disrupting disrupt VBG 10_1101-2020_12_31_424971 544 9 interactions interaction NNS 10_1101-2020_12_31_424971 544 10 between between IN 10_1101-2020_12_31_424971 544 11 1078 1078 CD 10_1101-2020_12_31_424971 544 12 decoding decode VBG 10_1101-2020_12_31_424971 544 13 center center NN 10_1101-2020_12_31_424971 544 14 and and CC 10_1101-2020_12_31_424971 544 15 codon codon NNP 10_1101-2020_12_31_424971 544 16 - - HYPH 10_1101-2020_12_31_424971 544 17 anticodon anticodon NNP 10_1101-2020_12_31_424971 544 18 duplex duplex NNP 10_1101-2020_12_31_424971 544 19 . . . 10_1101-2020_12_31_424971 545 1 Nat Nat NNP 10_1101-2020_12_31_424971 545 2 Struct Struct NNP 10_1101-2020_12_31_424971 545 3 Mol Mol NNP 10_1101-2020_12_31_424971 545 4 Biol Biol NNP 10_1101-2020_12_31_424971 545 5 21 21 CD 10_1101-2020_12_31_424971 545 6 , , , 10_1101-2020_12_31_424971 545 7 817 817 CD 10_1101-2020_12_31_424971 545 8 - - SYM 10_1101-2020_12_31_424971 545 9 24 24 CD 10_1101-2020_12_31_424971 545 10 ( ( -LRB- 10_1101-2020_12_31_424971 545 11 2014 2014 CD 10_1101-2020_12_31_424971 545 12 ) ) -RRB- 10_1101-2020_12_31_424971 545 13 . . . 10_1101-2020_12_31_424971 546 1 1079 1079 CD 10_1101-2020_12_31_424971 546 2 25 25 CD 10_1101-2020_12_31_424971 546 3 . . . 10_1101-2020_12_31_424971 547 1 Abeyrathne Abeyrathne NNP 10_1101-2020_12_31_424971 547 2 , , , 10_1101-2020_12_31_424971 547 3 P.D. P.D. NNP 10_1101-2020_12_31_424971 547 4 , , , 10_1101-2020_12_31_424971 547 5 Koh Koh NNP 10_1101-2020_12_31_424971 547 6 , , , 10_1101-2020_12_31_424971 547 7 C.S. C.S. NNP 10_1101-2020_12_31_424971 547 8 , , , 10_1101-2020_12_31_424971 547 9 Grant Grant NNP 10_1101-2020_12_31_424971 547 10 , , , 10_1101-2020_12_31_424971 547 11 T. T. NNP 10_1101-2020_12_31_424971 547 12 , , , 10_1101-2020_12_31_424971 547 13 Grigorieff Grigorieff NNP 10_1101-2020_12_31_424971 547 14 , , , 10_1101-2020_12_31_424971 547 15 N. N. NNP 10_1101-2020_12_31_424971 547 16 & & CC 10_1101-2020_12_31_424971 547 17 Korostelev Korostelev NNP 10_1101-2020_12_31_424971 547 18 , , , 10_1101-2020_12_31_424971 547 19 A.A. A.A. NNP 10_1101-2020_12_31_424971 547 20 Ensemble ensemble JJ 10_1101-2020_12_31_424971 547 21 cryo-1080 cryo-1080 VBP 10_1101-2020_12_31_424971 547 22 EM em NN 10_1101-2020_12_31_424971 547 23 uncovers uncover NNS 10_1101-2020_12_31_424971 547 24 inchworm inchworm NN 10_1101-2020_12_31_424971 547 25 - - HYPH 10_1101-2020_12_31_424971 547 26 like like JJ 10_1101-2020_12_31_424971 547 27 translocation translocation NN 10_1101-2020_12_31_424971 547 28 of of IN 10_1101-2020_12_31_424971 547 29 a a DT 10_1101-2020_12_31_424971 547 30 viral viral JJ 10_1101-2020_12_31_424971 547 31 IRES ires NN 10_1101-2020_12_31_424971 547 32 through through IN 10_1101-2020_12_31_424971 547 33 the the DT 10_1101-2020_12_31_424971 547 34 ribosome ribosome NN 10_1101-2020_12_31_424971 547 35 . . . 10_1101-2020_12_31_424971 548 1 Elife elife NN 10_1101-2020_12_31_424971 548 2 5 5 CD 10_1101-2020_12_31_424971 548 3 , , , 10_1101-2020_12_31_424971 548 4 1081 1081 CD 10_1101-2020_12_31_424971 548 5 doi doi XX 10_1101-2020_12_31_424971 548 6 : : : 10_1101-2020_12_31_424971 548 7 10.7554 10.7554 CD 10_1101-2020_12_31_424971 548 8 / / SYM 10_1101-2020_12_31_424971 548 9 eLife.14874 elife.14874 NN 10_1101-2020_12_31_424971 548 10 ( ( -LRB- 10_1101-2020_12_31_424971 548 11 2016 2016 CD 10_1101-2020_12_31_424971 548 12 ) ) -RRB- 10_1101-2020_12_31_424971 548 13 . . . 10_1101-2020_12_31_424971 549 1 1082 1082 CD 10_1101-2020_12_31_424971 549 2 26 26 CD 10_1101-2020_12_31_424971 549 3 . . . 10_1101-2020_12_31_424971 550 1 Schuwirth Schuwirth NNP 10_1101-2020_12_31_424971 550 2 , , , 10_1101-2020_12_31_424971 550 3 B.S. B.S. NNP 10_1101-2020_12_31_424971 551 1 et et NNP 10_1101-2020_12_31_424971 551 2 al al NNP 10_1101-2020_12_31_424971 551 3 . . . 10_1101-2020_12_31_424971 552 1 Structures structure NNS 10_1101-2020_12_31_424971 552 2 of of IN 10_1101-2020_12_31_424971 552 3 the the DT 10_1101-2020_12_31_424971 552 4 bacterial bacterial JJ 10_1101-2020_12_31_424971 552 5 ribosome ribosome NN 10_1101-2020_12_31_424971 552 6 at at IN 10_1101-2020_12_31_424971 552 7 3.5 3.5 CD 10_1101-2020_12_31_424971 552 8 A a DT 10_1101-2020_12_31_424971 552 9 resolution resolution NN 10_1101-2020_12_31_424971 552 10 . . . 10_1101-2020_12_31_424971 553 1 Science science NN 10_1101-2020_12_31_424971 553 2 1083 1083 CD 10_1101-2020_12_31_424971 553 3 310 310 CD 10_1101-2020_12_31_424971 553 4 , , , 10_1101-2020_12_31_424971 553 5 827 827 CD 10_1101-2020_12_31_424971 553 6 - - SYM 10_1101-2020_12_31_424971 553 7 34 34 CD 10_1101-2020_12_31_424971 553 8 ( ( -LRB- 10_1101-2020_12_31_424971 553 9 2005 2005 CD 10_1101-2020_12_31_424971 553 10 ) ) -RRB- 10_1101-2020_12_31_424971 553 11 . . . 10_1101-2020_12_31_424971 554 1 1084 1084 CD 10_1101-2020_12_31_424971 554 2 27 27 CD 10_1101-2020_12_31_424971 554 3 . . . 10_1101-2020_12_31_424971 555 1 Pulk pulk NN 10_1101-2020_12_31_424971 555 2 , , , 10_1101-2020_12_31_424971 555 3 A. a. NN 10_1101-2020_12_31_424971 556 1 & & CC 10_1101-2020_12_31_424971 556 2 Cate Cate NNP 10_1101-2020_12_31_424971 556 3 , , , 10_1101-2020_12_31_424971 556 4 J.H. J.H. NNP 10_1101-2020_12_31_424971 557 1 Control control NN 10_1101-2020_12_31_424971 557 2 of of IN 10_1101-2020_12_31_424971 557 3 ribosomal ribosomal NNP 10_1101-2020_12_31_424971 557 4 subunit subunit NN 10_1101-2020_12_31_424971 557 5 rotation rotation NN 10_1101-2020_12_31_424971 557 6 by by IN 10_1101-2020_12_31_424971 557 7 elongation elongation NN 10_1101-2020_12_31_424971 557 8 factor factor NN 10_1101-2020_12_31_424971 557 9 G. G. NNP 10_1101-2020_12_31_424971 557 10 1085 1085 CD 10_1101-2020_12_31_424971 557 11 Science Science NNP 10_1101-2020_12_31_424971 557 12 340 340 CD 10_1101-2020_12_31_424971 557 13 , , , 10_1101-2020_12_31_424971 557 14 1235970 1235970 CD 10_1101-2020_12_31_424971 557 15 ( ( -LRB- 10_1101-2020_12_31_424971 557 16 2013 2013 CD 10_1101-2020_12_31_424971 557 17 ) ) -RRB- 10_1101-2020_12_31_424971 557 18 . . . 10_1101-2020_12_31_424971 558 1 1086 1086 CD 10_1101-2020_12_31_424971 558 2 28 28 CD 10_1101-2020_12_31_424971 558 3 . . . 10_1101-2020_12_31_424971 559 1 Ratje Ratje NNP 10_1101-2020_12_31_424971 559 2 , , , 10_1101-2020_12_31_424971 559 3 A.H. A.H. NNP 10_1101-2020_12_31_424971 559 4 et et NNP 10_1101-2020_12_31_424971 559 5 al al NNP 10_1101-2020_12_31_424971 559 6 . . . 10_1101-2020_12_31_424971 560 1 Head Head NNP 10_1101-2020_12_31_424971 560 2 swivel swivel NN 10_1101-2020_12_31_424971 560 3 on on IN 10_1101-2020_12_31_424971 560 4 the the DT 10_1101-2020_12_31_424971 560 5 ribosome ribosome JJ 10_1101-2020_12_31_424971 560 6 facilitates facilitate NNS 10_1101-2020_12_31_424971 560 7 translocation translocation NN 10_1101-2020_12_31_424971 560 8 by by IN 10_1101-2020_12_31_424971 560 9 means mean NNS 10_1101-2020_12_31_424971 560 10 of of IN 10_1101-2020_12_31_424971 560 11 intra-1087 intra-1087 NNP 10_1101-2020_12_31_424971 560 12 subunit subunit NN 10_1101-2020_12_31_424971 560 13 tRNA tRNA NNP 10_1101-2020_12_31_424971 560 14 hybrid hybrid JJ 10_1101-2020_12_31_424971 560 15 sites site NNS 10_1101-2020_12_31_424971 560 16 . . . 10_1101-2020_12_31_424971 561 1 Nature nature NN 10_1101-2020_12_31_424971 561 2 468 468 CD 10_1101-2020_12_31_424971 561 3 , , , 10_1101-2020_12_31_424971 561 4 713 713 CD 10_1101-2020_12_31_424971 561 5 - - SYM 10_1101-2020_12_31_424971 561 6 6 6 CD 10_1101-2020_12_31_424971 561 7 ( ( -LRB- 10_1101-2020_12_31_424971 561 8 2010 2010 CD 10_1101-2020_12_31_424971 561 9 ) ) -RRB- 10_1101-2020_12_31_424971 561 10 . . . 10_1101-2020_12_31_424971 562 1 1088 1088 CD 10_1101-2020_12_31_424971 562 2 29 29 CD 10_1101-2020_12_31_424971 562 3 . . . 10_1101-2020_12_31_424971 563 1 Gamper Gamper NNP 10_1101-2020_12_31_424971 563 2 , , , 10_1101-2020_12_31_424971 563 3 H.B. H.B. NNP 10_1101-2020_12_31_424971 563 4 , , , 10_1101-2020_12_31_424971 563 5 Masuda Masuda NNP 10_1101-2020_12_31_424971 563 6 , , , 10_1101-2020_12_31_424971 563 7 I. I. NNP 10_1101-2020_12_31_424971 563 8 , , , 10_1101-2020_12_31_424971 563 9 Frenkel Frenkel NNP 10_1101-2020_12_31_424971 563 10 - - HYPH 10_1101-2020_12_31_424971 563 11 Morgenstern Morgenstern NNP 10_1101-2020_12_31_424971 563 12 , , , 10_1101-2020_12_31_424971 563 13 M. M. NNP 10_1101-2020_12_31_424971 563 14 & & CC 10_1101-2020_12_31_424971 563 15 Hou Hou NNP 10_1101-2020_12_31_424971 563 16 , , , 10_1101-2020_12_31_424971 563 17 Y.M. Y.M. NNP 10_1101-2020_12_31_424971 564 1 Maintenance maintenance NN 10_1101-2020_12_31_424971 564 2 of of IN 10_1101-2020_12_31_424971 564 3 protein protein NN 10_1101-2020_12_31_424971 564 4 1089 1089 CD 10_1101-2020_12_31_424971 564 5 synthesis synthesis NN 10_1101-2020_12_31_424971 564 6 reading reading NN 10_1101-2020_12_31_424971 564 7 frame frame NN 10_1101-2020_12_31_424971 564 8 by by IN 10_1101-2020_12_31_424971 564 9 EF EF NNP 10_1101-2020_12_31_424971 564 10 - - HYPH 10_1101-2020_12_31_424971 564 11 P P NNP 10_1101-2020_12_31_424971 564 12 and and CC 10_1101-2020_12_31_424971 564 13 m(1)G37-tRNA m(1)G37-tRNA NNP 10_1101-2020_12_31_424971 564 14 . . . 10_1101-2020_12_31_424971 565 1 Nat Nat NNP 10_1101-2020_12_31_424971 565 2 Commun Commun NNP 10_1101-2020_12_31_424971 565 3 6 6 CD 10_1101-2020_12_31_424971 565 4 , , , 10_1101-2020_12_31_424971 565 5 7226 7226 CD 10_1101-2020_12_31_424971 565 6 ( ( -LRB- 10_1101-2020_12_31_424971 565 7 2015 2015 CD 10_1101-2020_12_31_424971 565 8 ) ) -RRB- 10_1101-2020_12_31_424971 565 9 . . . 10_1101-2020_12_31_424971 566 1 1090 1090 CD 10_1101-2020_12_31_424971 566 2 30 30 CD 10_1101-2020_12_31_424971 566 3 . . . 10_1101-2020_12_31_424971 567 1 Masuda Masuda NNP 10_1101-2020_12_31_424971 567 2 , , , 10_1101-2020_12_31_424971 567 3 I. I. NNP 10_1101-2020_12_31_424971 567 4 et et NNP 10_1101-2020_12_31_424971 567 5 al al NNP 10_1101-2020_12_31_424971 567 6 . . . 10_1101-2020_12_31_424971 568 1 tRNA tRNA NNP 10_1101-2020_12_31_424971 568 2 Methylation methylation NN 10_1101-2020_12_31_424971 568 3 Is be VBZ 10_1101-2020_12_31_424971 568 4 a a DT 10_1101-2020_12_31_424971 568 5 Global Global NNP 10_1101-2020_12_31_424971 568 6 Determinant Determinant NNP 10_1101-2020_12_31_424971 568 7 of of IN 10_1101-2020_12_31_424971 568 8 Bacterial Bacterial NNP 10_1101-2020_12_31_424971 568 9 Multi Multi NNP 10_1101-2020_12_31_424971 568 10 - - JJ 10_1101-2020_12_31_424971 568 11 drug drug NNP 10_1101-2020_12_31_424971 568 12 1091 1091 CD 10_1101-2020_12_31_424971 568 13 Resistance Resistance NNP 10_1101-2020_12_31_424971 568 14 . . . 10_1101-2020_12_31_424971 569 1 Cell Cell NNP 10_1101-2020_12_31_424971 569 2 Syst Syst NNP 10_1101-2020_12_31_424971 569 3 8 8 CD 10_1101-2020_12_31_424971 569 4 , , , 10_1101-2020_12_31_424971 569 5 302 302 CD 10_1101-2020_12_31_424971 569 6 - - HYPH 10_1101-2020_12_31_424971 569 7 314 314 CD 10_1101-2020_12_31_424971 569 8 e8 e8 NNS 10_1101-2020_12_31_424971 569 9 ( ( -LRB- 10_1101-2020_12_31_424971 569 10 2019 2019 CD 10_1101-2020_12_31_424971 569 11 ) ) -RRB- 10_1101-2020_12_31_424971 569 12 . . . 10_1101-2020_12_31_424971 570 1 1092 1092 CD 10_1101-2020_12_31_424971 570 2 31 31 CD 10_1101-2020_12_31_424971 570 3 . . . 10_1101-2020_12_31_424971 571 1 Christian Christian NNP 10_1101-2020_12_31_424971 571 2 , , , 10_1101-2020_12_31_424971 571 3 T. T. NNP 10_1101-2020_12_31_424971 571 4 & & CC 10_1101-2020_12_31_424971 571 5 Hou Hou NNP 10_1101-2020_12_31_424971 571 6 , , , 10_1101-2020_12_31_424971 571 7 Y.M. Y.M. NNP 10_1101-2020_12_31_424971 572 1 Distinct distinct JJ 10_1101-2020_12_31_424971 572 2 determinants determinant NNS 10_1101-2020_12_31_424971 572 3 of of IN 10_1101-2020_12_31_424971 572 4 tRNA trna NN 10_1101-2020_12_31_424971 572 5 recognition recognition NN 10_1101-2020_12_31_424971 572 6 by by IN 10_1101-2020_12_31_424971 572 7 the the DT 10_1101-2020_12_31_424971 572 8 TrmD TrmD NNP 10_1101-2020_12_31_424971 572 9 and and CC 10_1101-2020_12_31_424971 572 10 1093 1093 CD 10_1101-2020_12_31_424971 572 11 Trm5 Trm5 NNP 10_1101-2020_12_31_424971 572 12 methyl methyl NN 10_1101-2020_12_31_424971 572 13 transferases transferase VBZ 10_1101-2020_12_31_424971 572 14 . . . 10_1101-2020_12_31_424971 573 1 J J NNP 10_1101-2020_12_31_424971 573 2 Mol Mol NNP 10_1101-2020_12_31_424971 573 3 Biol Biol NNP 10_1101-2020_12_31_424971 573 4 373 373 CD 10_1101-2020_12_31_424971 573 5 , , , 10_1101-2020_12_31_424971 573 6 623 623 CD 10_1101-2020_12_31_424971 573 7 - - SYM 10_1101-2020_12_31_424971 573 8 32 32 CD 10_1101-2020_12_31_424971 573 9 ( ( -LRB- 10_1101-2020_12_31_424971 573 10 2007 2007 CD 10_1101-2020_12_31_424971 573 11 ) ) -RRB- 10_1101-2020_12_31_424971 573 12 . . . 10_1101-2020_12_31_424971 574 1 1094 1094 CD 10_1101-2020_12_31_424971 574 2 32 32 CD 10_1101-2020_12_31_424971 574 3 . . . 10_1101-2020_12_31_424971 575 1 Murakami Murakami NNP 10_1101-2020_12_31_424971 575 2 , , , 10_1101-2020_12_31_424971 575 3 H. H. NNP 10_1101-2020_12_31_424971 575 4 , , , 10_1101-2020_12_31_424971 575 5 Ohta Ohta NNP 10_1101-2020_12_31_424971 575 6 , , , 10_1101-2020_12_31_424971 575 7 A. A. NNP 10_1101-2020_12_31_424971 575 8 , , , 10_1101-2020_12_31_424971 575 9 Ashigai Ashigai NNP 10_1101-2020_12_31_424971 575 10 , , , 10_1101-2020_12_31_424971 575 11 H. H. NNP 10_1101-2020_12_31_424971 575 12 & & CC 10_1101-2020_12_31_424971 575 13 Suga Suga NNP 10_1101-2020_12_31_424971 575 14 , , , 10_1101-2020_12_31_424971 575 15 H. H. NNP 10_1101-2020_12_31_424971 575 16 A a DT 10_1101-2020_12_31_424971 575 17 highly highly RB 10_1101-2020_12_31_424971 575 18 flexible flexible JJ 10_1101-2020_12_31_424971 575 19 tRNA trna NN 10_1101-2020_12_31_424971 575 20 acylation acylation NN 10_1101-2020_12_31_424971 575 21 method method NN 10_1101-2020_12_31_424971 575 22 1095 1095 CD 10_1101-2020_12_31_424971 575 23 for for IN 10_1101-2020_12_31_424971 575 24 non non JJ 10_1101-2020_12_31_424971 575 25 - - JJ 10_1101-2020_12_31_424971 575 26 natural natural JJ 10_1101-2020_12_31_424971 575 27 polypeptide polypeptide NN 10_1101-2020_12_31_424971 575 28 synthesis synthesis NN 10_1101-2020_12_31_424971 575 29 . . . 10_1101-2020_12_31_424971 576 1 Nat Nat NNP 10_1101-2020_12_31_424971 576 2 Methods Methods NNP 10_1101-2020_12_31_424971 576 3 3 3 CD 10_1101-2020_12_31_424971 576 4 , , , 10_1101-2020_12_31_424971 576 5 357 357 CD 10_1101-2020_12_31_424971 576 6 - - SYM 10_1101-2020_12_31_424971 576 7 9 9 CD 10_1101-2020_12_31_424971 576 8 ( ( -LRB- 10_1101-2020_12_31_424971 576 9 2006 2006 CD 10_1101-2020_12_31_424971 576 10 ) ) -RRB- 10_1101-2020_12_31_424971 576 11 . . . 10_1101-2020_12_31_424971 577 1 1096 1096 CD 10_1101-2020_12_31_424971 577 2 33 33 CD 10_1101-2020_12_31_424971 577 3 . . . 10_1101-2020_12_31_424971 578 1 Walker Walker NNP 10_1101-2020_12_31_424971 578 2 , , , 10_1101-2020_12_31_424971 578 3 S.E. S.E. NNP 10_1101-2020_12_31_424971 579 1 & & CC 10_1101-2020_12_31_424971 579 2 Fredrick Fredrick NNP 10_1101-2020_12_31_424971 579 3 , , , 10_1101-2020_12_31_424971 579 4 K. K. NNP 10_1101-2020_12_31_424971 579 5 Recognition Recognition NNP 10_1101-2020_12_31_424971 579 6 and and CC 10_1101-2020_12_31_424971 579 7 positioning positioning NN 10_1101-2020_12_31_424971 579 8 of of IN 10_1101-2020_12_31_424971 579 9 mRNA mRNA NNS 10_1101-2020_12_31_424971 579 10 in in IN 10_1101-2020_12_31_424971 579 11 the the DT 10_1101-2020_12_31_424971 579 12 ribosome ribosome NN 10_1101-2020_12_31_424971 579 13 by by IN 10_1101-2020_12_31_424971 579 14 1097 1097 CD 10_1101-2020_12_31_424971 579 15 tRNAs trnas CD 10_1101-2020_12_31_424971 579 16 with with IN 10_1101-2020_12_31_424971 579 17 expanded expand VBN 10_1101-2020_12_31_424971 579 18 anticodons anticodon NNS 10_1101-2020_12_31_424971 579 19 . . . 10_1101-2020_12_31_424971 580 1 J J NNP 10_1101-2020_12_31_424971 580 2 Mol Mol NNP 10_1101-2020_12_31_424971 580 3 Biol Biol NNP 10_1101-2020_12_31_424971 580 4 360 360 CD 10_1101-2020_12_31_424971 580 5 , , , 10_1101-2020_12_31_424971 580 6 599 599 CD 10_1101-2020_12_31_424971 580 7 - - SYM 10_1101-2020_12_31_424971 580 8 609 609 CD 10_1101-2020_12_31_424971 580 9 ( ( -LRB- 10_1101-2020_12_31_424971 580 10 2006 2006 CD 10_1101-2020_12_31_424971 580 11 ) ) -RRB- 10_1101-2020_12_31_424971 580 12 . . . 10_1101-2020_12_31_424971 581 1 1098 1098 CD 10_1101-2020_12_31_424971 581 2 34 34 CD 10_1101-2020_12_31_424971 581 3 . . . 10_1101-2020_12_31_424971 582 1 Gamper Gamper NNP 10_1101-2020_12_31_424971 582 2 , , , 10_1101-2020_12_31_424971 582 3 H.B. H.B. NNP 10_1101-2020_12_31_424971 582 4 , , , 10_1101-2020_12_31_424971 582 5 Masuda Masuda NNP 10_1101-2020_12_31_424971 582 6 , , , 10_1101-2020_12_31_424971 582 7 I. I. NNP 10_1101-2020_12_31_424971 582 8 , , , 10_1101-2020_12_31_424971 582 9 Frenkel Frenkel NNP 10_1101-2020_12_31_424971 582 10 - - HYPH 10_1101-2020_12_31_424971 582 11 Morgenstern Morgenstern NNP 10_1101-2020_12_31_424971 582 12 , , , 10_1101-2020_12_31_424971 582 13 M. M. NNP 10_1101-2020_12_31_424971 582 14 & & CC 10_1101-2020_12_31_424971 582 15 Hou Hou NNP 10_1101-2020_12_31_424971 582 16 , , , 10_1101-2020_12_31_424971 582 17 Y.M. Y.M. NNP 10_1101-2020_12_31_424971 583 1 The the DT 10_1101-2020_12_31_424971 583 2 UGG UGG NNP 10_1101-2020_12_31_424971 583 3 Isoacceptor Isoacceptor NNP 10_1101-2020_12_31_424971 583 4 1099 1099 CD 10_1101-2020_12_31_424971 583 5 of of IN 10_1101-2020_12_31_424971 583 6 tRNAPro tRNAPro NNP 10_1101-2020_12_31_424971 583 7 Is be VBZ 10_1101-2020_12_31_424971 583 8 Naturally naturally RB 10_1101-2020_12_31_424971 583 9 Prone prone JJ 10_1101-2020_12_31_424971 583 10 to to IN 10_1101-2020_12_31_424971 583 11 Frameshifts Frameshifts NNP 10_1101-2020_12_31_424971 583 12 . . . 10_1101-2020_12_31_424971 584 1 Int Int NNP 10_1101-2020_12_31_424971 584 2 J J NNP 10_1101-2020_12_31_424971 584 3 Mol Mol NNP 10_1101-2020_12_31_424971 584 4 Sci Sci NNP 10_1101-2020_12_31_424971 584 5 16 16 CD 10_1101-2020_12_31_424971 584 6 , , , 10_1101-2020_12_31_424971 584 7 14866 14866 CD 10_1101-2020_12_31_424971 584 8 - - SYM 10_1101-2020_12_31_424971 584 9 83 83 CD 10_1101-2020_12_31_424971 584 10 ( ( -LRB- 10_1101-2020_12_31_424971 584 11 2015 2015 CD 10_1101-2020_12_31_424971 584 12 ) ) -RRB- 10_1101-2020_12_31_424971 584 13 . . . 10_1101-2020_12_31_424971 585 1 1100 1100 CD 10_1101-2020_12_31_424971 585 2 35 35 CD 10_1101-2020_12_31_424971 585 3 . . . 10_1101-2020_12_31_424971 586 1 Fei Fei NNP 10_1101-2020_12_31_424971 586 2 , , , 10_1101-2020_12_31_424971 586 3 J. J. NNP 10_1101-2020_12_31_424971 586 4 et et NNP 10_1101-2020_12_31_424971 586 5 al al NNP 10_1101-2020_12_31_424971 586 6 . . . 10_1101-2020_12_31_424971 587 1 Allosteric allosteric JJ 10_1101-2020_12_31_424971 587 2 collaboration collaboration NN 10_1101-2020_12_31_424971 587 3 between between IN 10_1101-2020_12_31_424971 587 4 elongation elongation NN 10_1101-2020_12_31_424971 587 5 factor factor NN 10_1101-2020_12_31_424971 587 6 G g NN 10_1101-2020_12_31_424971 587 7 and and CC 10_1101-2020_12_31_424971 587 8 the the DT 10_1101-2020_12_31_424971 587 9 ribosomal ribosomal NN 10_1101-2020_12_31_424971 587 10 L1 L1 NNP 10_1101-2020_12_31_424971 587 11 1101 1101 CD 10_1101-2020_12_31_424971 587 12 stalk stalk NN 10_1101-2020_12_31_424971 587 13 directs direct VBZ 10_1101-2020_12_31_424971 587 14 tRNA trna JJ 10_1101-2020_12_31_424971 587 15 movements movement NNS 10_1101-2020_12_31_424971 587 16 during during IN 10_1101-2020_12_31_424971 587 17 translation translation NN 10_1101-2020_12_31_424971 587 18 . . . 10_1101-2020_12_31_424971 588 1 Proc Proc NNP 10_1101-2020_12_31_424971 588 2 Natl Natl NNP 10_1101-2020_12_31_424971 588 3 Acad Acad NNP 10_1101-2020_12_31_424971 588 4 Sci Sci NNP 10_1101-2020_12_31_424971 588 5 U U NNP 10_1101-2020_12_31_424971 588 6 S S NNP 10_1101-2020_12_31_424971 588 7 A A NNP 10_1101-2020_12_31_424971 588 8 106 106 CD 10_1101-2020_12_31_424971 588 9 , , , 10_1101-2020_12_31_424971 588 10 15702 15702 CD 10_1101-2020_12_31_424971 588 11 - - SYM 10_1101-2020_12_31_424971 588 12 1102 1102 CD 10_1101-2020_12_31_424971 588 13 7 7 CD 10_1101-2020_12_31_424971 588 14 ( ( -LRB- 10_1101-2020_12_31_424971 588 15 2009 2009 CD 10_1101-2020_12_31_424971 588 16 ) ) -RRB- 10_1101-2020_12_31_424971 588 17 . . . 10_1101-2020_12_31_424971 589 1 1103 1103 CD 10_1101-2020_12_31_424971 589 2 36 36 CD 10_1101-2020_12_31_424971 589 3 . . . 10_1101-2020_12_31_424971 590 1 Ning Ning NNP 10_1101-2020_12_31_424971 590 2 , , , 10_1101-2020_12_31_424971 590 3 W. W. NNP 10_1101-2020_12_31_424971 590 4 , , , 10_1101-2020_12_31_424971 590 5 Fei Fei NNP 10_1101-2020_12_31_424971 590 6 , , , 10_1101-2020_12_31_424971 590 7 J. J. NNP 10_1101-2020_12_31_424971 591 1 & & CC 10_1101-2020_12_31_424971 591 2 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 591 3 , , , 10_1101-2020_12_31_424971 591 4 R.L. R.L. NNP 10_1101-2020_12_31_424971 591 5 , , , 10_1101-2020_12_31_424971 591 6 Jr. Jr. NNP 10_1101-2020_12_31_424971 592 1 The the DT 10_1101-2020_12_31_424971 592 2 ribosome ribosome NN 10_1101-2020_12_31_424971 592 3 uses use VBZ 10_1101-2020_12_31_424971 592 4 cooperative cooperative JJ 10_1101-2020_12_31_424971 592 5 conformational conformational JJ 10_1101-2020_12_31_424971 592 6 1104 1104 CD 10_1101-2020_12_31_424971 592 7 changes change NNS 10_1101-2020_12_31_424971 592 8 to to TO 10_1101-2020_12_31_424971 592 9 maximize maximize VB 10_1101-2020_12_31_424971 592 10 and and CC 10_1101-2020_12_31_424971 592 11 regulate regulate VB 10_1101-2020_12_31_424971 592 12 the the DT 10_1101-2020_12_31_424971 592 13 efficiency efficiency NN 10_1101-2020_12_31_424971 592 14 of of IN 10_1101-2020_12_31_424971 592 15 translation translation NN 10_1101-2020_12_31_424971 592 16 . . . 10_1101-2020_12_31_424971 593 1 Proc Proc NNP 10_1101-2020_12_31_424971 593 2 Natl Natl NNP 10_1101-2020_12_31_424971 593 3 Acad Acad NNP 10_1101-2020_12_31_424971 593 4 Sci Sci NNP 10_1101-2020_12_31_424971 593 5 U U NNP 10_1101-2020_12_31_424971 593 6 S S NNP 10_1101-2020_12_31_424971 593 7 A A NNP 10_1101-2020_12_31_424971 593 8 1105 1105 CD 10_1101-2020_12_31_424971 593 9 111 111 CD 10_1101-2020_12_31_424971 593 10 , , , 10_1101-2020_12_31_424971 593 11 12073 12073 CD 10_1101-2020_12_31_424971 593 12 - - SYM 10_1101-2020_12_31_424971 593 13 8 8 CD 10_1101-2020_12_31_424971 593 14 ( ( -LRB- 10_1101-2020_12_31_424971 593 15 2014 2014 CD 10_1101-2020_12_31_424971 593 16 ) ) -RRB- 10_1101-2020_12_31_424971 593 17 . . . 10_1101-2020_12_31_424971 594 1 1106 1106 CD 10_1101-2020_12_31_424971 594 2 37 37 CD 10_1101-2020_12_31_424971 594 3 . . . 10_1101-2020_12_31_424971 595 1 Fei Fei NNP 10_1101-2020_12_31_424971 595 2 , , , 10_1101-2020_12_31_424971 595 3 J. J. NNP 10_1101-2020_12_31_424971 595 4 , , , 10_1101-2020_12_31_424971 595 5 Kosuri Kosuri NNP 10_1101-2020_12_31_424971 595 6 , , , 10_1101-2020_12_31_424971 595 7 P. P. NNP 10_1101-2020_12_31_424971 595 8 , , , 10_1101-2020_12_31_424971 595 9 MacDougall MacDougall NNP 10_1101-2020_12_31_424971 595 10 , , , 10_1101-2020_12_31_424971 595 11 D.D. D.D. NNP 10_1101-2020_12_31_424971 596 1 & & CC 10_1101-2020_12_31_424971 596 2 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 596 3 , , , 10_1101-2020_12_31_424971 596 4 R.L. R.L. NNP 10_1101-2020_12_31_424971 596 5 , , , 10_1101-2020_12_31_424971 596 6 Jr. Jr. NNP 10_1101-2020_12_31_424971 596 7 Coupling Coupling NNP 10_1101-2020_12_31_424971 596 8 of of IN 10_1101-2020_12_31_424971 596 9 ribosomal ribosomal NNP 10_1101-2020_12_31_424971 596 10 L1 L1 NNP 10_1101-2020_12_31_424971 596 11 stalk stalk NNP 10_1101-2020_12_31_424971 596 12 1107 1107 CD 10_1101-2020_12_31_424971 596 13 and and CC 10_1101-2020_12_31_424971 596 14 tRNA trna JJ 10_1101-2020_12_31_424971 596 15 dynamics dynamic NNS 10_1101-2020_12_31_424971 596 16 during during IN 10_1101-2020_12_31_424971 596 17 translation translation NN 10_1101-2020_12_31_424971 596 18 elongation elongation NN 10_1101-2020_12_31_424971 596 19 . . . 10_1101-2020_12_31_424971 597 1 Mol Mol NNP 10_1101-2020_12_31_424971 597 2 Cell Cell NNP 10_1101-2020_12_31_424971 597 3 30 30 CD 10_1101-2020_12_31_424971 597 4 , , , 10_1101-2020_12_31_424971 597 5 348 348 CD 10_1101-2020_12_31_424971 597 6 - - HYPH 10_1101-2020_12_31_424971 597 7 59 59 CD 10_1101-2020_12_31_424971 597 8 ( ( -LRB- 10_1101-2020_12_31_424971 597 9 2008 2008 CD 10_1101-2020_12_31_424971 597 10 ) ) -RRB- 10_1101-2020_12_31_424971 597 11 . . . 10_1101-2020_12_31_424971 598 1 1108 1108 CD 10_1101-2020_12_31_424971 598 2 38 38 CD 10_1101-2020_12_31_424971 598 3 . . . 10_1101-2020_12_31_424971 599 1 Fei Fei NNP 10_1101-2020_12_31_424971 599 2 , , , 10_1101-2020_12_31_424971 599 3 J. J. NNP 10_1101-2020_12_31_424971 599 4 , , , 10_1101-2020_12_31_424971 599 5 Richard Richard NNP 10_1101-2020_12_31_424971 599 6 , , , 10_1101-2020_12_31_424971 599 7 A.C. A.C. NNP 10_1101-2020_12_31_424971 599 8 , , , 10_1101-2020_12_31_424971 599 9 Bronson Bronson NNP 10_1101-2020_12_31_424971 599 10 , , , 10_1101-2020_12_31_424971 599 11 J.E. J.E. NNP 10_1101-2020_12_31_424971 600 1 & & CC 10_1101-2020_12_31_424971 600 2 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 600 3 , , , 10_1101-2020_12_31_424971 600 4 R.L. R.L. NNP 10_1101-2020_12_31_424971 600 5 , , , 10_1101-2020_12_31_424971 600 6 Jr. Jr. NNP 10_1101-2020_12_31_424971 601 1 Transfer Transfer NNP 10_1101-2020_12_31_424971 601 2 RNA RNA NNP 10_1101-2020_12_31_424971 601 3 - - HYPH 10_1101-2020_12_31_424971 601 4 mediated mediate VBN 10_1101-2020_12_31_424971 601 5 1109 1109 CD 10_1101-2020_12_31_424971 601 6 regulation regulation NN 10_1101-2020_12_31_424971 601 7 of of IN 10_1101-2020_12_31_424971 601 8 ribosome ribosome JJ 10_1101-2020_12_31_424971 601 9 dynamics dynamic NNS 10_1101-2020_12_31_424971 601 10 during during IN 10_1101-2020_12_31_424971 601 11 protein protein NN 10_1101-2020_12_31_424971 601 12 synthesis synthesis NN 10_1101-2020_12_31_424971 601 13 . . . 10_1101-2020_12_31_424971 602 1 Nat Nat NNP 10_1101-2020_12_31_424971 602 2 Struct Struct NNP 10_1101-2020_12_31_424971 602 3 Mol Mol NNP 10_1101-2020_12_31_424971 602 4 Biol Biol NNP 10_1101-2020_12_31_424971 602 5 18 18 CD 10_1101-2020_12_31_424971 602 6 , , , 10_1101-2020_12_31_424971 602 7 1043 1043 CD 10_1101-2020_12_31_424971 602 8 - - SYM 10_1101-2020_12_31_424971 602 9 1110 1110 CD 10_1101-2020_12_31_424971 602 10 51 51 CD 10_1101-2020_12_31_424971 602 11 ( ( -LRB- 10_1101-2020_12_31_424971 602 12 2011 2011 CD 10_1101-2020_12_31_424971 602 13 ) ) -RRB- 10_1101-2020_12_31_424971 602 14 . . . 10_1101-2020_12_31_424971 603 1 1111 1111 CD 10_1101-2020_12_31_424971 603 2 39 39 CD 10_1101-2020_12_31_424971 603 3 . . . 10_1101-2020_12_31_424971 604 1 Boel Boel NNP 10_1101-2020_12_31_424971 604 2 , , , 10_1101-2020_12_31_424971 604 3 G. G. NNP 10_1101-2020_12_31_424971 604 4 et et NNP 10_1101-2020_12_31_424971 604 5 al al NNP 10_1101-2020_12_31_424971 604 6 . . . 10_1101-2020_12_31_424971 605 1 The the DT 10_1101-2020_12_31_424971 605 2 ABC ABC NNP 10_1101-2020_12_31_424971 605 3 - - HYPH 10_1101-2020_12_31_424971 605 4 F F NNP 10_1101-2020_12_31_424971 605 5 protein protein NN 10_1101-2020_12_31_424971 605 6 EttA EttA NNP 10_1101-2020_12_31_424971 605 7 gates gate NNS 10_1101-2020_12_31_424971 605 8 ribosome ribosome VBD 10_1101-2020_12_31_424971 605 9 entry entry NN 10_1101-2020_12_31_424971 605 10 into into IN 10_1101-2020_12_31_424971 605 11 the the DT 10_1101-2020_12_31_424971 605 12 translation translation NN 10_1101-2020_12_31_424971 605 13 1112 1112 CD 10_1101-2020_12_31_424971 605 14 elongation elongation NN 10_1101-2020_12_31_424971 605 15 cycle cycle NN 10_1101-2020_12_31_424971 605 16 . . . 10_1101-2020_12_31_424971 606 1 Nat Nat NNP 10_1101-2020_12_31_424971 606 2 Struct Struct NNP 10_1101-2020_12_31_424971 606 3 Mol Mol NNP 10_1101-2020_12_31_424971 606 4 Biol Biol NNP 10_1101-2020_12_31_424971 606 5 21 21 CD 10_1101-2020_12_31_424971 606 6 , , , 10_1101-2020_12_31_424971 606 7 143 143 CD 10_1101-2020_12_31_424971 606 8 - - SYM 10_1101-2020_12_31_424971 606 9 51 51 CD 10_1101-2020_12_31_424971 606 10 ( ( -LRB- 10_1101-2020_12_31_424971 606 11 2014 2014 CD 10_1101-2020_12_31_424971 606 12 ) ) -RRB- 10_1101-2020_12_31_424971 606 13 . . . 10_1101-2020_12_31_424971 607 1 1113 1113 CD 10_1101-2020_12_31_424971 607 2 40 40 CD 10_1101-2020_12_31_424971 607 3 . . . 10_1101-2020_12_31_424971 608 1 Chen Chen NNP 10_1101-2020_12_31_424971 608 2 , , , 10_1101-2020_12_31_424971 608 3 B. B. NNP 10_1101-2020_12_31_424971 608 4 et et NNP 10_1101-2020_12_31_424971 608 5 al al NNP 10_1101-2020_12_31_424971 608 6 . . . 10_1101-2020_12_31_424971 609 1 EttA EttA NNP 10_1101-2020_12_31_424971 609 2 regulates regulate VBZ 10_1101-2020_12_31_424971 609 3 translation translation NN 10_1101-2020_12_31_424971 609 4 by by IN 10_1101-2020_12_31_424971 609 5 binding bind VBG 10_1101-2020_12_31_424971 609 6 the the DT 10_1101-2020_12_31_424971 609 7 ribosomal ribosomal NN 10_1101-2020_12_31_424971 609 8 E e NN 10_1101-2020_12_31_424971 609 9 site site NN 10_1101-2020_12_31_424971 609 10 and and CC 10_1101-2020_12_31_424971 609 11 restricting restrict VBG 10_1101-2020_12_31_424971 609 12 1114 1114 CD 10_1101-2020_12_31_424971 609 13 ribosome ribosome NN 10_1101-2020_12_31_424971 609 14 - - HYPH 10_1101-2020_12_31_424971 609 15 tRNA trna NN 10_1101-2020_12_31_424971 609 16 dynamics dynamic NNS 10_1101-2020_12_31_424971 609 17 . . . 10_1101-2020_12_31_424971 610 1 Nat Nat NNP 10_1101-2020_12_31_424971 610 2 Struct Struct NNP 10_1101-2020_12_31_424971 610 3 Mol Mol NNP 10_1101-2020_12_31_424971 610 4 Biol Biol NNP 10_1101-2020_12_31_424971 610 5 21 21 CD 10_1101-2020_12_31_424971 610 6 , , , 10_1101-2020_12_31_424971 610 7 152 152 CD 10_1101-2020_12_31_424971 610 8 - - SYM 10_1101-2020_12_31_424971 610 9 9 9 CD 10_1101-2020_12_31_424971 610 10 ( ( -LRB- 10_1101-2020_12_31_424971 610 11 2014 2014 CD 10_1101-2020_12_31_424971 610 12 ) ) -RRB- 10_1101-2020_12_31_424971 610 13 . . . 10_1101-2020_12_31_424971 611 1 1115 1115 CD 10_1101-2020_12_31_424971 611 2 41 41 CD 10_1101-2020_12_31_424971 611 3 . . . 10_1101-2020_12_31_424971 612 1 Kim Kim NNP 10_1101-2020_12_31_424971 612 2 , , , 10_1101-2020_12_31_424971 612 3 H.K. H.K. NNP 10_1101-2020_12_31_424971 613 1 et et NNP 10_1101-2020_12_31_424971 613 2 al al NNP 10_1101-2020_12_31_424971 613 3 . . . 10_1101-2020_12_31_424971 614 1 A a DT 10_1101-2020_12_31_424971 614 2 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 614 3 stimulatory stimulatory JJ 10_1101-2020_12_31_424971 614 4 stem stem NN 10_1101-2020_12_31_424971 614 5 loop loop NN 10_1101-2020_12_31_424971 614 6 destabilizes destabilize VBZ 10_1101-2020_12_31_424971 614 7 the the DT 10_1101-2020_12_31_424971 614 8 hybrid hybrid JJ 10_1101-2020_12_31_424971 614 9 state state NN 10_1101-2020_12_31_424971 614 10 and and CC 10_1101-2020_12_31_424971 614 11 1116 1116 CD 10_1101-2020_12_31_424971 614 12 impedes impede NNS 10_1101-2020_12_31_424971 614 13 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 614 14 translocation translocation NN 10_1101-2020_12_31_424971 614 15 . . . 10_1101-2020_12_31_424971 615 1 Proc Proc NNP 10_1101-2020_12_31_424971 615 2 Natl Natl NNP 10_1101-2020_12_31_424971 615 3 Acad Acad NNP 10_1101-2020_12_31_424971 615 4 Sci Sci NNP 10_1101-2020_12_31_424971 615 5 U U NNP 10_1101-2020_12_31_424971 615 6 S S NNP 10_1101-2020_12_31_424971 615 7 A A NNP 10_1101-2020_12_31_424971 615 8 111 111 CD 10_1101-2020_12_31_424971 615 9 , , , 10_1101-2020_12_31_424971 615 10 5538 5538 CD 10_1101-2020_12_31_424971 615 11 - - SYM 10_1101-2020_12_31_424971 615 12 43 43 CD 10_1101-2020_12_31_424971 615 13 ( ( -LRB- 10_1101-2020_12_31_424971 615 14 2014 2014 CD 10_1101-2020_12_31_424971 615 15 ) ) -RRB- 10_1101-2020_12_31_424971 615 16 . . . 10_1101-2020_12_31_424971 616 1 1117 1117 CD 10_1101-2020_12_31_424971 616 2 .CC .CC : 10_1101-2020_12_31_424971 616 3 - - : 10_1101-2020_12_31_424971 616 4 BY by IN 10_1101-2020_12_31_424971 616 5 - - HYPH 10_1101-2020_12_31_424971 616 6 NC NC NNP 10_1101-2020_12_31_424971 616 7 - - HYPH 10_1101-2020_12_31_424971 616 8 ND ND NNP 10_1101-2020_12_31_424971 616 9 4.0 4.0 CD 10_1101-2020_12_31_424971 616 10 International International NNP 10_1101-2020_12_31_424971 616 11 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 616 12 under under IN 10_1101-2020_12_31_424971 616 13 a a DT 10_1101-2020_12_31_424971 616 14 ( ( -LRB- 10_1101-2020_12_31_424971 616 15 which which WDT 10_1101-2020_12_31_424971 616 16 was be VBD 10_1101-2020_12_31_424971 616 17 not not RB 10_1101-2020_12_31_424971 616 18 certified certify VBN 10_1101-2020_12_31_424971 616 19 by by IN 10_1101-2020_12_31_424971 616 20 peer peer NN 10_1101-2020_12_31_424971 616 21 review review NN 10_1101-2020_12_31_424971 616 22 ) ) -RRB- 10_1101-2020_12_31_424971 616 23 is be VBZ 10_1101-2020_12_31_424971 616 24 the the DT 10_1101-2020_12_31_424971 616 25 author author NN 10_1101-2020_12_31_424971 616 26 / / SYM 10_1101-2020_12_31_424971 616 27 funder funder NN 10_1101-2020_12_31_424971 616 28 , , , 10_1101-2020_12_31_424971 616 29 who who WP 10_1101-2020_12_31_424971 616 30 has have VBZ 10_1101-2020_12_31_424971 616 31 granted grant VBN 10_1101-2020_12_31_424971 616 32 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 616 33 a a DT 10_1101-2020_12_31_424971 616 34 license license NN 10_1101-2020_12_31_424971 616 35 to to TO 10_1101-2020_12_31_424971 616 36 display display VB 10_1101-2020_12_31_424971 616 37 the the DT 10_1101-2020_12_31_424971 616 38 preprint preprint NN 10_1101-2020_12_31_424971 616 39 in in IN 10_1101-2020_12_31_424971 616 40 perpetuity perpetuity NN 10_1101-2020_12_31_424971 616 41 . . . 10_1101-2020_12_31_424971 617 1 It -PRON- PRP 10_1101-2020_12_31_424971 617 2 is be VBZ 10_1101-2020_12_31_424971 617 3 made make VBN 10_1101-2020_12_31_424971 617 4 The the DT 10_1101-2020_12_31_424971 617 5 copyright copyright NN 10_1101-2020_12_31_424971 617 6 holder holder NN 10_1101-2020_12_31_424971 617 7 for for IN 10_1101-2020_12_31_424971 617 8 this this DT 10_1101-2020_12_31_424971 617 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 617 10 version version NN 10_1101-2020_12_31_424971 617 11 posted post VBD 10_1101-2020_12_31_424971 617 12 January January NNP 10_1101-2020_12_31_424971 617 13 2 2 CD 10_1101-2020_12_31_424971 617 14 , , , 10_1101-2020_12_31_424971 617 15 2021 2021 CD 10_1101-2020_12_31_424971 617 16 . . . 10_1101-2020_12_31_424971 617 17 ; ; : 10_1101-2020_12_31_424971 617 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 617 19 : : : 10_1101-2020_12_31_424971 617 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 617 21 preprint preprint NN 10_1101-2020_12_31_424971 617 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NNP 10_1101-2020_12_31_424971 617 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 617 24 44 44 CD 10_1101-2020_12_31_424971 617 25 42 42 CD 10_1101-2020_12_31_424971 617 26 . . . 10_1101-2020_12_31_424971 618 1 Munro Munro NNP 10_1101-2020_12_31_424971 618 2 , , , 10_1101-2020_12_31_424971 618 3 J.B. J.B. NNP 10_1101-2020_12_31_424971 618 4 , , , 10_1101-2020_12_31_424971 618 5 Wasserman Wasserman NNP 10_1101-2020_12_31_424971 618 6 , , , 10_1101-2020_12_31_424971 618 7 M.R. M.R. NNP 10_1101-2020_12_31_424971 618 8 , , , 10_1101-2020_12_31_424971 618 9 Altman Altman NNP 10_1101-2020_12_31_424971 618 10 , , , 10_1101-2020_12_31_424971 618 11 R.B. R.B. NNP 10_1101-2020_12_31_424971 618 12 , , , 10_1101-2020_12_31_424971 618 13 Wang Wang NNP 10_1101-2020_12_31_424971 618 14 , , , 10_1101-2020_12_31_424971 618 15 L. L. NNP 10_1101-2020_12_31_424971 618 16 & & CC 10_1101-2020_12_31_424971 618 17 Blanchard Blanchard NNP 10_1101-2020_12_31_424971 618 18 , , , 10_1101-2020_12_31_424971 618 19 S.C. South Carolina NNP 10_1101-2020_12_31_424971 618 20 Correlated correlate VBD 10_1101-2020_12_31_424971 618 21 1118 1118 CD 10_1101-2020_12_31_424971 618 22 conformational conformational JJ 10_1101-2020_12_31_424971 618 23 events event NNS 10_1101-2020_12_31_424971 618 24 in in IN 10_1101-2020_12_31_424971 618 25 EF EF NNP 10_1101-2020_12_31_424971 618 26 - - HYPH 10_1101-2020_12_31_424971 618 27 G G NNP 10_1101-2020_12_31_424971 618 28 and and CC 10_1101-2020_12_31_424971 618 29 the the DT 10_1101-2020_12_31_424971 618 30 ribosome ribosome JJ 10_1101-2020_12_31_424971 618 31 regulate regulate NN 10_1101-2020_12_31_424971 618 32 translocation translocation NN 10_1101-2020_12_31_424971 618 33 . . . 10_1101-2020_12_31_424971 619 1 Nat Nat NNP 10_1101-2020_12_31_424971 619 2 Struct Struct NNP 10_1101-2020_12_31_424971 619 3 Mol Mol NNP 10_1101-2020_12_31_424971 619 4 1119 1119 CD 10_1101-2020_12_31_424971 619 5 Biol Biol NNP 10_1101-2020_12_31_424971 619 6 17 17 CD 10_1101-2020_12_31_424971 619 7 , , , 10_1101-2020_12_31_424971 619 8 1470 1470 CD 10_1101-2020_12_31_424971 619 9 - - SYM 10_1101-2020_12_31_424971 619 10 7 7 CD 10_1101-2020_12_31_424971 619 11 ( ( -LRB- 10_1101-2020_12_31_424971 619 12 2010 2010 CD 10_1101-2020_12_31_424971 619 13 ) ) -RRB- 10_1101-2020_12_31_424971 619 14 . . . 10_1101-2020_12_31_424971 620 1 1120 1120 CD 10_1101-2020_12_31_424971 620 2 43 43 CD 10_1101-2020_12_31_424971 620 3 . . . 10_1101-2020_12_31_424971 621 1 Blanchard Blanchard NNP 10_1101-2020_12_31_424971 621 2 , , , 10_1101-2020_12_31_424971 621 3 S.C. South Carolina NNP 10_1101-2020_12_31_424971 621 4 , , , 10_1101-2020_12_31_424971 621 5 Kim Kim NNP 10_1101-2020_12_31_424971 621 6 , , , 10_1101-2020_12_31_424971 621 7 H.D. H.D. NNP 10_1101-2020_12_31_424971 621 8 , , , 10_1101-2020_12_31_424971 621 9 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 621 10 , , , 10_1101-2020_12_31_424971 621 11 R.L. R.L. NNP 10_1101-2020_12_31_424971 621 12 , , , 10_1101-2020_12_31_424971 621 13 Jr. Jr. NNP 10_1101-2020_12_31_424971 621 14 , , , 10_1101-2020_12_31_424971 621 15 Puglisi Puglisi NNP 10_1101-2020_12_31_424971 621 16 , , , 10_1101-2020_12_31_424971 621 17 J.D. J.D. NNP 10_1101-2020_12_31_424971 622 1 & & CC 10_1101-2020_12_31_424971 622 2 Chu Chu NNP 10_1101-2020_12_31_424971 622 3 , , , 10_1101-2020_12_31_424971 622 4 S. S. NNP 10_1101-2020_12_31_424971 622 5 tRNA trna NN 10_1101-2020_12_31_424971 622 6 dynamics dynamic NNS 10_1101-2020_12_31_424971 622 7 1121 1121 CD 10_1101-2020_12_31_424971 622 8 on on IN 10_1101-2020_12_31_424971 622 9 the the DT 10_1101-2020_12_31_424971 622 10 ribosome ribosome NN 10_1101-2020_12_31_424971 622 11 during during IN 10_1101-2020_12_31_424971 622 12 translation translation NN 10_1101-2020_12_31_424971 622 13 . . . 10_1101-2020_12_31_424971 623 1 Proc Proc NNP 10_1101-2020_12_31_424971 623 2 Natl Natl NNP 10_1101-2020_12_31_424971 623 3 Acad Acad NNP 10_1101-2020_12_31_424971 623 4 Sci Sci NNP 10_1101-2020_12_31_424971 623 5 U U NNP 10_1101-2020_12_31_424971 623 6 S S NNP 10_1101-2020_12_31_424971 623 7 A A NNP 10_1101-2020_12_31_424971 623 8 101 101 CD 10_1101-2020_12_31_424971 623 9 , , , 10_1101-2020_12_31_424971 623 10 12893 12893 CD 10_1101-2020_12_31_424971 623 11 - - SYM 10_1101-2020_12_31_424971 623 12 8 8 CD 10_1101-2020_12_31_424971 623 13 ( ( -LRB- 10_1101-2020_12_31_424971 623 14 2004 2004 CD 10_1101-2020_12_31_424971 623 15 ) ) -RRB- 10_1101-2020_12_31_424971 623 16 . . . 10_1101-2020_12_31_424971 624 1 1122 1122 CD 10_1101-2020_12_31_424971 624 2 44 44 CD 10_1101-2020_12_31_424971 624 3 . . . 10_1101-2020_12_31_424971 625 1 Studer Studer NNP 10_1101-2020_12_31_424971 625 2 , , , 10_1101-2020_12_31_424971 625 3 S.M. S.M. NNP 10_1101-2020_12_31_424971 625 4 , , , 10_1101-2020_12_31_424971 625 5 Feinberg Feinberg NNP 10_1101-2020_12_31_424971 625 6 , , , 10_1101-2020_12_31_424971 625 7 J.S. J.S. NNP 10_1101-2020_12_31_424971 626 1 & & CC 10_1101-2020_12_31_424971 626 2 Joseph Joseph NNP 10_1101-2020_12_31_424971 626 3 , , , 10_1101-2020_12_31_424971 626 4 S. S. NNP 10_1101-2020_12_31_424971 626 5 Rapid Rapid NNP 10_1101-2020_12_31_424971 626 6 kinetic kinetic JJ 10_1101-2020_12_31_424971 626 7 analysis analysis NN 10_1101-2020_12_31_424971 626 8 of of IN 10_1101-2020_12_31_424971 626 9 EF EF NNP 10_1101-2020_12_31_424971 626 10 - - HYPH 10_1101-2020_12_31_424971 626 11 G G NNP 10_1101-2020_12_31_424971 626 12 - - HYPH 10_1101-2020_12_31_424971 626 13 dependent dependent JJ 10_1101-2020_12_31_424971 626 14 1123 1123 CD 10_1101-2020_12_31_424971 626 15 mRNA mrna NN 10_1101-2020_12_31_424971 626 16 translocation translocation NN 10_1101-2020_12_31_424971 626 17 in in IN 10_1101-2020_12_31_424971 626 18 the the DT 10_1101-2020_12_31_424971 626 19 ribosome ribosome NN 10_1101-2020_12_31_424971 626 20 . . . 10_1101-2020_12_31_424971 627 1 J J NNP 10_1101-2020_12_31_424971 627 2 Mol Mol NNP 10_1101-2020_12_31_424971 627 3 Biol Biol NNP 10_1101-2020_12_31_424971 627 4 327 327 CD 10_1101-2020_12_31_424971 627 5 , , , 10_1101-2020_12_31_424971 627 6 369 369 CD 10_1101-2020_12_31_424971 627 7 - - SYM 10_1101-2020_12_31_424971 627 8 81 81 CD 10_1101-2020_12_31_424971 627 9 ( ( -LRB- 10_1101-2020_12_31_424971 627 10 2003 2003 CD 10_1101-2020_12_31_424971 627 11 ) ) -RRB- 10_1101-2020_12_31_424971 627 12 . . . 10_1101-2020_12_31_424971 628 1 1124 1124 CD 10_1101-2020_12_31_424971 628 2 45 45 CD 10_1101-2020_12_31_424971 628 3 . . . 10_1101-2020_12_31_424971 629 1 Wintermeyer Wintermeyer NNP 10_1101-2020_12_31_424971 629 2 , , , 10_1101-2020_12_31_424971 629 3 W. W. NNP 10_1101-2020_12_31_424971 629 4 & & CC 10_1101-2020_12_31_424971 629 5 Rodnina Rodnina NNP 10_1101-2020_12_31_424971 629 6 , , , 10_1101-2020_12_31_424971 629 7 M.V. M.V. NNP 10_1101-2020_12_31_424971 630 1 Translational translational JJ 10_1101-2020_12_31_424971 630 2 elongation elongation NN 10_1101-2020_12_31_424971 630 3 factor factor NN 10_1101-2020_12_31_424971 630 4 G g NN 10_1101-2020_12_31_424971 630 5 : : : 10_1101-2020_12_31_424971 630 6 a a DT 10_1101-2020_12_31_424971 630 7 GTP gtp NN 10_1101-2020_12_31_424971 630 8 - - HYPH 10_1101-2020_12_31_424971 630 9 driven drive VBN 10_1101-2020_12_31_424971 630 10 motor motor NN 10_1101-2020_12_31_424971 630 11 1125 1125 CD 10_1101-2020_12_31_424971 630 12 of of IN 10_1101-2020_12_31_424971 630 13 the the DT 10_1101-2020_12_31_424971 630 14 ribosome ribosome NN 10_1101-2020_12_31_424971 630 15 . . . 10_1101-2020_12_31_424971 631 1 Essays Essays NNPS 10_1101-2020_12_31_424971 631 2 Biochem Biochem NNP 10_1101-2020_12_31_424971 631 3 35 35 CD 10_1101-2020_12_31_424971 631 4 , , , 10_1101-2020_12_31_424971 631 5 117 117 CD 10_1101-2020_12_31_424971 631 6 - - SYM 10_1101-2020_12_31_424971 631 7 29 29 CD 10_1101-2020_12_31_424971 631 8 ( ( -LRB- 10_1101-2020_12_31_424971 631 9 2000 2000 CD 10_1101-2020_12_31_424971 631 10 ) ) -RRB- 10_1101-2020_12_31_424971 631 11 . . . 10_1101-2020_12_31_424971 632 1 1126 1126 CD 10_1101-2020_12_31_424971 632 2 46 46 CD 10_1101-2020_12_31_424971 632 3 . . . 10_1101-2020_12_31_424971 633 1 Ermolenko Ermolenko NNP 10_1101-2020_12_31_424971 633 2 , , , 10_1101-2020_12_31_424971 633 3 D.N. D.N. NNP 10_1101-2020_12_31_424971 634 1 et et NNP 10_1101-2020_12_31_424971 634 2 al al NNP 10_1101-2020_12_31_424971 634 3 . . . 10_1101-2020_12_31_424971 635 1 Observation observation NN 10_1101-2020_12_31_424971 635 2 of of IN 10_1101-2020_12_31_424971 635 3 intersubunit intersubunit JJ 10_1101-2020_12_31_424971 635 4 movement movement NN 10_1101-2020_12_31_424971 635 5 of of IN 10_1101-2020_12_31_424971 635 6 the the DT 10_1101-2020_12_31_424971 635 7 ribosome ribosome NN 10_1101-2020_12_31_424971 635 8 in in IN 10_1101-2020_12_31_424971 635 9 solution solution NN 10_1101-2020_12_31_424971 635 10 1127 1127 CD 10_1101-2020_12_31_424971 635 11 using use VBG 10_1101-2020_12_31_424971 635 12 FRET FRET NNP 10_1101-2020_12_31_424971 635 13 . . . 10_1101-2020_12_31_424971 636 1 J J NNP 10_1101-2020_12_31_424971 636 2 Mol Mol NNP 10_1101-2020_12_31_424971 636 3 Biol Biol NNP 10_1101-2020_12_31_424971 636 4 370 370 CD 10_1101-2020_12_31_424971 636 5 , , , 10_1101-2020_12_31_424971 636 6 530 530 CD 10_1101-2020_12_31_424971 636 7 - - SYM 10_1101-2020_12_31_424971 636 8 40 40 CD 10_1101-2020_12_31_424971 636 9 ( ( -LRB- 10_1101-2020_12_31_424971 636 10 2007 2007 CD 10_1101-2020_12_31_424971 636 11 ) ) -RRB- 10_1101-2020_12_31_424971 636 12 . . . 10_1101-2020_12_31_424971 637 1 1128 1128 CD 10_1101-2020_12_31_424971 637 2 47 47 CD 10_1101-2020_12_31_424971 637 3 . . . 10_1101-2020_12_31_424971 638 1 Ermolenko Ermolenko NNP 10_1101-2020_12_31_424971 638 2 , , , 10_1101-2020_12_31_424971 638 3 D.N. D.N. NNP 10_1101-2020_12_31_424971 639 1 & & CC 10_1101-2020_12_31_424971 639 2 Noller Noller NNP 10_1101-2020_12_31_424971 639 3 , , , 10_1101-2020_12_31_424971 639 4 H.F. H.F. NNP 10_1101-2020_12_31_424971 639 5 mRNA mrna NN 10_1101-2020_12_31_424971 639 6 translocation translocation NN 10_1101-2020_12_31_424971 639 7 occurs occur VBZ 10_1101-2020_12_31_424971 639 8 during during IN 10_1101-2020_12_31_424971 639 9 the the DT 10_1101-2020_12_31_424971 639 10 second second JJ 10_1101-2020_12_31_424971 639 11 step step NN 10_1101-2020_12_31_424971 639 12 of of IN 10_1101-2020_12_31_424971 639 13 1129 1129 CD 10_1101-2020_12_31_424971 639 14 ribosomal ribosomal JJ 10_1101-2020_12_31_424971 639 15 intersubunit intersubunit JJ 10_1101-2020_12_31_424971 639 16 rotation rotation NN 10_1101-2020_12_31_424971 639 17 . . . 10_1101-2020_12_31_424971 640 1 Nat Nat NNP 10_1101-2020_12_31_424971 640 2 Struct Struct NNP 10_1101-2020_12_31_424971 640 3 Mol Mol NNP 10_1101-2020_12_31_424971 640 4 Biol Biol NNP 10_1101-2020_12_31_424971 640 5 18 18 CD 10_1101-2020_12_31_424971 640 6 , , , 10_1101-2020_12_31_424971 640 7 457 457 CD 10_1101-2020_12_31_424971 640 8 - - SYM 10_1101-2020_12_31_424971 640 9 62 62 CD 10_1101-2020_12_31_424971 640 10 ( ( -LRB- 10_1101-2020_12_31_424971 640 11 2011 2011 CD 10_1101-2020_12_31_424971 640 12 ) ) -RRB- 10_1101-2020_12_31_424971 640 13 . . . 10_1101-2020_12_31_424971 641 1 1130 1130 CD 10_1101-2020_12_31_424971 641 2 48 48 CD 10_1101-2020_12_31_424971 641 3 . . . 10_1101-2020_12_31_424971 642 1 Cornish Cornish NNP 10_1101-2020_12_31_424971 642 2 , , , 10_1101-2020_12_31_424971 642 3 P.V. P.V. NNP 10_1101-2020_12_31_424971 643 1 et et NNP 10_1101-2020_12_31_424971 643 2 al al NNP 10_1101-2020_12_31_424971 643 3 . . . 10_1101-2020_12_31_424971 644 1 Following follow VBG 10_1101-2020_12_31_424971 644 2 movement movement NN 10_1101-2020_12_31_424971 644 3 of of IN 10_1101-2020_12_31_424971 644 4 the the DT 10_1101-2020_12_31_424971 644 5 L1 L1 NNP 10_1101-2020_12_31_424971 644 6 stalk stalk NN 10_1101-2020_12_31_424971 644 7 between between IN 10_1101-2020_12_31_424971 644 8 three three CD 10_1101-2020_12_31_424971 644 9 functional functional JJ 10_1101-2020_12_31_424971 644 10 states state NNS 10_1101-2020_12_31_424971 644 11 1131 1131 CD 10_1101-2020_12_31_424971 644 12 in in IN 10_1101-2020_12_31_424971 644 13 single single JJ 10_1101-2020_12_31_424971 644 14 ribosomes ribosome NNS 10_1101-2020_12_31_424971 644 15 . . . 10_1101-2020_12_31_424971 645 1 Proc Proc NNP 10_1101-2020_12_31_424971 645 2 Natl Natl NNP 10_1101-2020_12_31_424971 645 3 Acad Acad NNP 10_1101-2020_12_31_424971 645 4 Sci Sci NNP 10_1101-2020_12_31_424971 645 5 U U NNP 10_1101-2020_12_31_424971 645 6 S S NNP 10_1101-2020_12_31_424971 645 7 A A NNP 10_1101-2020_12_31_424971 645 8 106 106 CD 10_1101-2020_12_31_424971 645 9 , , , 10_1101-2020_12_31_424971 645 10 2571 2571 CD 10_1101-2020_12_31_424971 645 11 - - SYM 10_1101-2020_12_31_424971 645 12 6 6 CD 10_1101-2020_12_31_424971 645 13 ( ( -LRB- 10_1101-2020_12_31_424971 645 14 2009 2009 CD 10_1101-2020_12_31_424971 645 15 ) ) -RRB- 10_1101-2020_12_31_424971 645 16 . . . 10_1101-2020_12_31_424971 646 1 1132 1132 CD 10_1101-2020_12_31_424971 646 2 49 49 CD 10_1101-2020_12_31_424971 646 3 . . . 10_1101-2020_12_31_424971 647 1 Nguyen Nguyen NNP 10_1101-2020_12_31_424971 647 2 , , , 10_1101-2020_12_31_424971 647 3 H.A. H.A. NNP 10_1101-2020_12_31_424971 647 4 , , , 10_1101-2020_12_31_424971 647 5 Hoffer Hoffer NNP 10_1101-2020_12_31_424971 647 6 , , , 10_1101-2020_12_31_424971 647 7 E.D. E.D. NNP 10_1101-2020_12_31_424971 648 1 & & CC 10_1101-2020_12_31_424971 648 2 Dunham Dunham NNP 10_1101-2020_12_31_424971 648 3 , , , 10_1101-2020_12_31_424971 648 4 C.M. C.M. NNP 10_1101-2020_12_31_424971 649 1 Importance importance NN 10_1101-2020_12_31_424971 649 2 of of IN 10_1101-2020_12_31_424971 649 3 a a DT 10_1101-2020_12_31_424971 649 4 tRNA trna NN 10_1101-2020_12_31_424971 649 5 anticodon anticodon NNP 10_1101-2020_12_31_424971 649 6 loop loop NNP 10_1101-2020_12_31_424971 649 7 1133 1133 CD 10_1101-2020_12_31_424971 649 8 modification modification NN 10_1101-2020_12_31_424971 649 9 and and CC 10_1101-2020_12_31_424971 649 10 a a DT 10_1101-2020_12_31_424971 649 11 conserved conserve VBN 10_1101-2020_12_31_424971 649 12 , , , 10_1101-2020_12_31_424971 649 13 noncanonical noncanonical JJ 10_1101-2020_12_31_424971 649 14 anticodon anticodon NN 10_1101-2020_12_31_424971 649 15 stem stem NN 10_1101-2020_12_31_424971 649 16 pairing pair VBG 10_1101-2020_12_31_424971 649 17 in in IN 10_1101-2020_12_31_424971 649 18 tRNACGGProfor trnacggprofor CD 10_1101-2020_12_31_424971 649 19 1134 1134 CD 10_1101-2020_12_31_424971 649 20 decoding decoding NN 10_1101-2020_12_31_424971 649 21 . . . 10_1101-2020_12_31_424971 650 1 J J NNP 10_1101-2020_12_31_424971 650 2 Biol Biol NNP 10_1101-2020_12_31_424971 650 3 Chem Chem NNP 10_1101-2020_12_31_424971 650 4 294 294 CD 10_1101-2020_12_31_424971 650 5 , , , 10_1101-2020_12_31_424971 650 6 5281 5281 CD 10_1101-2020_12_31_424971 650 7 - - SYM 10_1101-2020_12_31_424971 650 8 5291 5291 CD 10_1101-2020_12_31_424971 650 9 ( ( -LRB- 10_1101-2020_12_31_424971 650 10 2019 2019 CD 10_1101-2020_12_31_424971 650 11 ) ) -RRB- 10_1101-2020_12_31_424971 650 12 . . . 10_1101-2020_12_31_424971 651 1 1135 1135 CD 10_1101-2020_12_31_424971 651 2 50 50 CD 10_1101-2020_12_31_424971 651 3 . . . 10_1101-2020_12_31_424971 652 1 Guo Guo NNP 10_1101-2020_12_31_424971 652 2 , , , 10_1101-2020_12_31_424971 652 3 Z. Z. NNP 10_1101-2020_12_31_424971 653 1 & & CC 10_1101-2020_12_31_424971 653 2 Noller Noller NNP 10_1101-2020_12_31_424971 653 3 , , , 10_1101-2020_12_31_424971 653 4 H.F. H.F. NNP 10_1101-2020_12_31_424971 653 5 Rotation Rotation NNP 10_1101-2020_12_31_424971 653 6 of of IN 10_1101-2020_12_31_424971 653 7 the the DT 10_1101-2020_12_31_424971 653 8 head head NN 10_1101-2020_12_31_424971 653 9 of of IN 10_1101-2020_12_31_424971 653 10 the the DT 10_1101-2020_12_31_424971 653 11 30S 30s CD 10_1101-2020_12_31_424971 653 12 ribosomal ribosomal NN 10_1101-2020_12_31_424971 653 13 subunit subunit NN 10_1101-2020_12_31_424971 653 14 during during IN 10_1101-2020_12_31_424971 653 15 mRNA mRNA NNP 10_1101-2020_12_31_424971 653 16 1136 1136 CD 10_1101-2020_12_31_424971 653 17 translocation translocation NN 10_1101-2020_12_31_424971 653 18 . . . 10_1101-2020_12_31_424971 654 1 Proc Proc NNP 10_1101-2020_12_31_424971 654 2 Natl Natl NNP 10_1101-2020_12_31_424971 654 3 Acad Acad NNP 10_1101-2020_12_31_424971 654 4 Sci Sci NNP 10_1101-2020_12_31_424971 654 5 U U NNP 10_1101-2020_12_31_424971 654 6 S S NNP 10_1101-2020_12_31_424971 654 7 A A NNP 10_1101-2020_12_31_424971 654 8 109 109 CD 10_1101-2020_12_31_424971 654 9 , , , 10_1101-2020_12_31_424971 654 10 20391 20391 CD 10_1101-2020_12_31_424971 654 11 - - SYM 10_1101-2020_12_31_424971 654 12 4 4 CD 10_1101-2020_12_31_424971 654 13 ( ( -LRB- 10_1101-2020_12_31_424971 654 14 2012 2012 CD 10_1101-2020_12_31_424971 654 15 ) ) -RRB- 10_1101-2020_12_31_424971 654 16 . . . 10_1101-2020_12_31_424971 655 1 1137 1137 CD 10_1101-2020_12_31_424971 655 2 51 51 CD 10_1101-2020_12_31_424971 655 3 . . . 10_1101-2020_12_31_424971 656 1 Zhou Zhou NNP 10_1101-2020_12_31_424971 656 2 , , , 10_1101-2020_12_31_424971 656 3 J. J. NNP 10_1101-2020_12_31_424971 656 4 , , , 10_1101-2020_12_31_424971 656 5 Lancaster Lancaster NNP 10_1101-2020_12_31_424971 656 6 , , , 10_1101-2020_12_31_424971 656 7 L. L. NNP 10_1101-2020_12_31_424971 656 8 , , , 10_1101-2020_12_31_424971 656 9 Donohue Donohue NNP 10_1101-2020_12_31_424971 656 10 , , , 10_1101-2020_12_31_424971 656 11 J.P. J.P. NNP 10_1101-2020_12_31_424971 656 12 & & CC 10_1101-2020_12_31_424971 656 13 Noller Noller NNP 10_1101-2020_12_31_424971 656 14 , , , 10_1101-2020_12_31_424971 656 15 H.F. H.F. NNP 10_1101-2020_12_31_424971 656 16 Spontaneous spontaneous JJ 10_1101-2020_12_31_424971 656 17 ribosomal ribosomal NN 10_1101-2020_12_31_424971 656 18 1138 1138 CD 10_1101-2020_12_31_424971 656 19 translocation translocation NN 10_1101-2020_12_31_424971 656 20 of of IN 10_1101-2020_12_31_424971 656 21 mRNA mRNA NNS 10_1101-2020_12_31_424971 656 22 and and CC 10_1101-2020_12_31_424971 656 23 tRNAs trnas CD 10_1101-2020_12_31_424971 656 24 into into IN 10_1101-2020_12_31_424971 656 25 a a DT 10_1101-2020_12_31_424971 656 26 chimeric chimeric JJ 10_1101-2020_12_31_424971 656 27 hybrid hybrid JJ 10_1101-2020_12_31_424971 656 28 state state NN 10_1101-2020_12_31_424971 656 29 . . . 10_1101-2020_12_31_424971 657 1 Proc Proc NNP 10_1101-2020_12_31_424971 657 2 Natl Natl NNP 10_1101-2020_12_31_424971 657 3 Acad Acad NNP 10_1101-2020_12_31_424971 657 4 Sci Sci NNP 10_1101-2020_12_31_424971 657 5 U U NNP 10_1101-2020_12_31_424971 657 6 S S NNP 10_1101-2020_12_31_424971 657 7 A A NNP 10_1101-2020_12_31_424971 657 8 1139 1139 CD 10_1101-2020_12_31_424971 657 9 116 116 CD 10_1101-2020_12_31_424971 657 10 , , , 10_1101-2020_12_31_424971 657 11 7813 7813 CD 10_1101-2020_12_31_424971 657 12 - - SYM 10_1101-2020_12_31_424971 657 13 7818 7818 CD 10_1101-2020_12_31_424971 657 14 ( ( -LRB- 10_1101-2020_12_31_424971 657 15 2019 2019 CD 10_1101-2020_12_31_424971 657 16 ) ) -RRB- 10_1101-2020_12_31_424971 657 17 . . . 10_1101-2020_12_31_424971 658 1 1140 1140 CD 10_1101-2020_12_31_424971 658 2 52 52 CD 10_1101-2020_12_31_424971 658 3 . . . 10_1101-2020_12_31_424971 659 1 Korniy Korniy NNP 10_1101-2020_12_31_424971 659 2 , , , 10_1101-2020_12_31_424971 659 3 N. N. NNP 10_1101-2020_12_31_424971 659 4 , , , 10_1101-2020_12_31_424971 659 5 Samatova Samatova NNP 10_1101-2020_12_31_424971 659 6 , , , 10_1101-2020_12_31_424971 659 7 E. E. NNP 10_1101-2020_12_31_424971 659 8 , , , 10_1101-2020_12_31_424971 659 9 Anokhina Anokhina NNP 10_1101-2020_12_31_424971 659 10 , , , 10_1101-2020_12_31_424971 659 11 M.M. M.M. NNP 10_1101-2020_12_31_424971 659 12 , , , 10_1101-2020_12_31_424971 659 13 Peske Peske NNP 10_1101-2020_12_31_424971 659 14 , , , 10_1101-2020_12_31_424971 659 15 F. F. NNP 10_1101-2020_12_31_424971 659 16 & & CC 10_1101-2020_12_31_424971 659 17 Rodnina Rodnina NNP 10_1101-2020_12_31_424971 659 18 , , , 10_1101-2020_12_31_424971 659 19 M.V. M.V. NNP 10_1101-2020_12_31_424971 660 1 Mechanisms mechanism NNS 10_1101-2020_12_31_424971 660 2 and and CC 10_1101-2020_12_31_424971 660 3 1141 1141 CD 10_1101-2020_12_31_424971 660 4 biomedical biomedical JJ 10_1101-2020_12_31_424971 660 5 implications implication NNS 10_1101-2020_12_31_424971 660 6 of of IN 10_1101-2020_12_31_424971 660 7 -1 -1 . 10_1101-2020_12_31_424971 660 8 programmed program VBN 10_1101-2020_12_31_424971 660 9 ribosome ribosome NNP 10_1101-2020_12_31_424971 660 10 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 660 11 on on IN 10_1101-2020_12_31_424971 660 12 viral viral JJ 10_1101-2020_12_31_424971 660 13 and and CC 10_1101-2020_12_31_424971 660 14 bacterial bacterial JJ 10_1101-2020_12_31_424971 660 15 1142 1142 CD 10_1101-2020_12_31_424971 660 16 mRNAs mrnas ADD 10_1101-2020_12_31_424971 660 17 . . . 10_1101-2020_12_31_424971 661 1 FEBS FEBS NNP 10_1101-2020_12_31_424971 661 2 Lett Lett NNP 10_1101-2020_12_31_424971 661 3 593 593 CD 10_1101-2020_12_31_424971 661 4 , , , 10_1101-2020_12_31_424971 661 5 1468 1468 CD 10_1101-2020_12_31_424971 661 6 - - SYM 10_1101-2020_12_31_424971 661 7 1482 1482 CD 10_1101-2020_12_31_424971 661 8 ( ( -LRB- 10_1101-2020_12_31_424971 661 9 2019 2019 CD 10_1101-2020_12_31_424971 661 10 ) ) -RRB- 10_1101-2020_12_31_424971 661 11 . . . 10_1101-2020_12_31_424971 662 1 1143 1143 CD 10_1101-2020_12_31_424971 662 2 53 53 CD 10_1101-2020_12_31_424971 662 3 . . . 10_1101-2020_12_31_424971 663 1 Lajoie Lajoie NNP 10_1101-2020_12_31_424971 663 2 , , , 10_1101-2020_12_31_424971 663 3 M.J. M.J. NNP 10_1101-2020_12_31_424971 663 4 et et NNP 10_1101-2020_12_31_424971 663 5 al al NNP 10_1101-2020_12_31_424971 663 6 . . . 10_1101-2020_12_31_424971 664 1 Genomically genomically RB 10_1101-2020_12_31_424971 664 2 recoded recode VBD 10_1101-2020_12_31_424971 664 3 organisms organism NNS 10_1101-2020_12_31_424971 664 4 expand expand VBP 10_1101-2020_12_31_424971 664 5 biological biological JJ 10_1101-2020_12_31_424971 664 6 functions function NNS 10_1101-2020_12_31_424971 664 7 . . . 10_1101-2020_12_31_424971 665 1 Science science NN 10_1101-2020_12_31_424971 665 2 1144 1144 CD 10_1101-2020_12_31_424971 665 3 342 342 CD 10_1101-2020_12_31_424971 665 4 , , , 10_1101-2020_12_31_424971 665 5 357 357 CD 10_1101-2020_12_31_424971 665 6 - - HYPH 10_1101-2020_12_31_424971 665 7 60 60 CD 10_1101-2020_12_31_424971 665 8 ( ( -LRB- 10_1101-2020_12_31_424971 665 9 2013 2013 CD 10_1101-2020_12_31_424971 665 10 ) ) -RRB- 10_1101-2020_12_31_424971 665 11 . . . 10_1101-2020_12_31_424971 666 1 1145 1145 CD 10_1101-2020_12_31_424971 666 2 54 54 CD 10_1101-2020_12_31_424971 666 3 . . . 10_1101-2020_12_31_424971 667 1 Wang Wang NNP 10_1101-2020_12_31_424971 667 2 , , , 10_1101-2020_12_31_424971 667 3 K. K. NNP 10_1101-2020_12_31_424971 667 4 , , , 10_1101-2020_12_31_424971 667 5 de de NNP 10_1101-2020_12_31_424971 667 6 la la NNP 10_1101-2020_12_31_424971 667 7 Torre Torre NNP 10_1101-2020_12_31_424971 667 8 , , , 10_1101-2020_12_31_424971 667 9 D. D. NNP 10_1101-2020_12_31_424971 667 10 , , , 10_1101-2020_12_31_424971 667 11 Robertson Robertson NNP 10_1101-2020_12_31_424971 667 12 , , , 10_1101-2020_12_31_424971 667 13 W.E. W.E. NNP 10_1101-2020_12_31_424971 668 1 & & CC 10_1101-2020_12_31_424971 668 2 Chin Chin NNP 10_1101-2020_12_31_424971 668 3 , , , 10_1101-2020_12_31_424971 668 4 J.W. J.W. NNP 10_1101-2020_12_31_424971 669 1 Programmed program VBN 10_1101-2020_12_31_424971 669 2 chromosome chromosome NN 10_1101-2020_12_31_424971 669 3 1146 1146 CD 10_1101-2020_12_31_424971 669 4 fission fission NN 10_1101-2020_12_31_424971 669 5 and and CC 10_1101-2020_12_31_424971 669 6 fusion fusion NN 10_1101-2020_12_31_424971 669 7 enable enable VBP 10_1101-2020_12_31_424971 669 8 precise precise JJ 10_1101-2020_12_31_424971 669 9 large large JJ 10_1101-2020_12_31_424971 669 10 - - HYPH 10_1101-2020_12_31_424971 669 11 scale scale NN 10_1101-2020_12_31_424971 669 12 genome genome JJ 10_1101-2020_12_31_424971 669 13 rearrangement rearrangement NN 10_1101-2020_12_31_424971 669 14 and and CC 10_1101-2020_12_31_424971 669 15 assembly assembly NN 10_1101-2020_12_31_424971 669 16 . . . 10_1101-2020_12_31_424971 670 1 1147 1147 CD 10_1101-2020_12_31_424971 670 2 Science Science NNP 10_1101-2020_12_31_424971 670 3 365 365 CD 10_1101-2020_12_31_424971 670 4 , , , 10_1101-2020_12_31_424971 670 5 922 922 CD 10_1101-2020_12_31_424971 670 6 - - SYM 10_1101-2020_12_31_424971 670 7 926 926 CD 10_1101-2020_12_31_424971 670 8 ( ( -LRB- 10_1101-2020_12_31_424971 670 9 2019 2019 CD 10_1101-2020_12_31_424971 670 10 ) ) -RRB- 10_1101-2020_12_31_424971 670 11 . . . 10_1101-2020_12_31_424971 671 1 1148 1148 CD 10_1101-2020_12_31_424971 671 2 55 55 CD 10_1101-2020_12_31_424971 671 3 . . . 10_1101-2020_12_31_424971 672 1 Mohan Mohan NNP 10_1101-2020_12_31_424971 672 2 , , , 10_1101-2020_12_31_424971 672 3 S. S. NNP 10_1101-2020_12_31_424971 672 4 , , , 10_1101-2020_12_31_424971 672 5 Donohue Donohue NNP 10_1101-2020_12_31_424971 672 6 , , , 10_1101-2020_12_31_424971 672 7 J.P. J.P. NNP 10_1101-2020_12_31_424971 672 8 & & CC 10_1101-2020_12_31_424971 672 9 Noller Noller NNP 10_1101-2020_12_31_424971 672 10 , , , 10_1101-2020_12_31_424971 672 11 H.F. H.F. NNP 10_1101-2020_12_31_424971 672 12 Molecular Molecular NNP 10_1101-2020_12_31_424971 672 13 mechanics mechanic NNS 10_1101-2020_12_31_424971 672 14 of of IN 10_1101-2020_12_31_424971 672 15 30S 30s CD 10_1101-2020_12_31_424971 672 16 subunit subunit NN 10_1101-2020_12_31_424971 672 17 head head VB 10_1101-2020_12_31_424971 672 18 1149 1149 CD 10_1101-2020_12_31_424971 672 19 rotation rotation NN 10_1101-2020_12_31_424971 672 20 . . . 10_1101-2020_12_31_424971 673 1 Proc Proc NNP 10_1101-2020_12_31_424971 673 2 Natl Natl NNP 10_1101-2020_12_31_424971 673 3 Acad Acad NNP 10_1101-2020_12_31_424971 673 4 Sci Sci NNP 10_1101-2020_12_31_424971 673 5 U U NNP 10_1101-2020_12_31_424971 673 6 S S NNP 10_1101-2020_12_31_424971 673 7 A A NNP 10_1101-2020_12_31_424971 673 8 111 111 CD 10_1101-2020_12_31_424971 673 9 , , , 10_1101-2020_12_31_424971 673 10 13325 13325 CD 10_1101-2020_12_31_424971 673 11 - - SYM 10_1101-2020_12_31_424971 673 12 30 30 CD 10_1101-2020_12_31_424971 673 13 ( ( -LRB- 10_1101-2020_12_31_424971 673 14 2014 2014 CD 10_1101-2020_12_31_424971 673 15 ) ) -RRB- 10_1101-2020_12_31_424971 673 16 . . . 10_1101-2020_12_31_424971 674 1 1150 1150 CD 10_1101-2020_12_31_424971 674 2 56 56 CD 10_1101-2020_12_31_424971 674 3 . . . 10_1101-2020_12_31_424971 675 1 Kaledhonkar Kaledhonkar NNP 10_1101-2020_12_31_424971 675 2 , , , 10_1101-2020_12_31_424971 675 3 S. S. NNP 10_1101-2020_12_31_424971 675 4 et et NNP 10_1101-2020_12_31_424971 675 5 al al NNP 10_1101-2020_12_31_424971 675 6 . . . 10_1101-2020_12_31_424971 676 1 Late late JJ 10_1101-2020_12_31_424971 676 2 steps step NNS 10_1101-2020_12_31_424971 676 3 in in IN 10_1101-2020_12_31_424971 676 4 bacterial bacterial JJ 10_1101-2020_12_31_424971 676 5 translation translation NN 10_1101-2020_12_31_424971 676 6 initiation initiation NN 10_1101-2020_12_31_424971 676 7 visualized visualize VBN 10_1101-2020_12_31_424971 676 8 using use VBG 10_1101-2020_12_31_424971 676 9 time-1151 time-1151 NNP 10_1101-2020_12_31_424971 676 10 resolved resolve VBN 10_1101-2020_12_31_424971 676 11 cryo cryo JJ 10_1101-2020_12_31_424971 676 12 - - HYPH 10_1101-2020_12_31_424971 676 13 EM EM NNP 10_1101-2020_12_31_424971 676 14 . . . 10_1101-2020_12_31_424971 677 1 Nature nature NN 10_1101-2020_12_31_424971 677 2 570 570 CD 10_1101-2020_12_31_424971 677 3 , , , 10_1101-2020_12_31_424971 677 4 400 400 CD 10_1101-2020_12_31_424971 677 5 - - SYM 10_1101-2020_12_31_424971 677 6 404 404 CD 10_1101-2020_12_31_424971 677 7 ( ( -LRB- 10_1101-2020_12_31_424971 677 8 2019 2019 CD 10_1101-2020_12_31_424971 677 9 ) ) -RRB- 10_1101-2020_12_31_424971 677 10 . . . 10_1101-2020_12_31_424971 678 1 1152 1152 CD 10_1101-2020_12_31_424971 678 2 57 57 CD 10_1101-2020_12_31_424971 678 3 . . . 10_1101-2020_12_31_424971 679 1 Chen Chen NNP 10_1101-2020_12_31_424971 679 2 , , , 10_1101-2020_12_31_424971 679 3 B. B. NNP 10_1101-2020_12_31_424971 679 4 et et NNP 10_1101-2020_12_31_424971 679 5 al al NNP 10_1101-2020_12_31_424971 679 6 . . . 10_1101-2020_12_31_424971 680 1 Structural structural JJ 10_1101-2020_12_31_424971 680 2 dynamics dynamic NNS 10_1101-2020_12_31_424971 680 3 of of IN 10_1101-2020_12_31_424971 680 4 ribosome ribosome NNP 10_1101-2020_12_31_424971 680 5 subunit subunit NNP 10_1101-2020_12_31_424971 680 6 association association NNP 10_1101-2020_12_31_424971 680 7 studied study VBN 10_1101-2020_12_31_424971 680 8 by by IN 10_1101-2020_12_31_424971 680 9 mixing-1153 mixing-1153 JJ 10_1101-2020_12_31_424971 680 10 spraying spray VBG 10_1101-2020_12_31_424971 680 11 time time NN 10_1101-2020_12_31_424971 680 12 - - HYPH 10_1101-2020_12_31_424971 680 13 resolved resolve VBN 10_1101-2020_12_31_424971 680 14 cryogenic cryogenic JJ 10_1101-2020_12_31_424971 680 15 electron electron NN 10_1101-2020_12_31_424971 680 16 microscopy microscopy NN 10_1101-2020_12_31_424971 680 17 . . . 10_1101-2020_12_31_424971 681 1 Structure structure NN 10_1101-2020_12_31_424971 681 2 23 23 CD 10_1101-2020_12_31_424971 681 3 , , , 10_1101-2020_12_31_424971 681 4 1097 1097 CD 10_1101-2020_12_31_424971 681 5 - - SYM 10_1101-2020_12_31_424971 681 6 105 105 CD 10_1101-2020_12_31_424971 681 7 ( ( -LRB- 10_1101-2020_12_31_424971 681 8 2015 2015 CD 10_1101-2020_12_31_424971 681 9 ) ) -RRB- 10_1101-2020_12_31_424971 681 10 . . . 10_1101-2020_12_31_424971 682 1 1154 1154 CD 10_1101-2020_12_31_424971 682 2 58 58 CD 10_1101-2020_12_31_424971 682 3 . . . 10_1101-2020_12_31_424971 683 1 Reinkemeier Reinkemeier NNP 10_1101-2020_12_31_424971 683 2 , , , 10_1101-2020_12_31_424971 683 3 C.D. C.D. NNP 10_1101-2020_12_31_424971 683 4 , , , 10_1101-2020_12_31_424971 683 5 Girona Girona NNP 10_1101-2020_12_31_424971 683 6 , , , 10_1101-2020_12_31_424971 683 7 G.E. G.E. NNP 10_1101-2020_12_31_424971 684 1 & & CC 10_1101-2020_12_31_424971 684 2 Lemke Lemke NNP 10_1101-2020_12_31_424971 684 3 , , , 10_1101-2020_12_31_424971 684 4 E.A. E.A. NNP 10_1101-2020_12_31_424971 685 1 Designer designer NN 10_1101-2020_12_31_424971 685 2 membraneless membraneless NN 10_1101-2020_12_31_424971 685 3 organelles organelle NNS 10_1101-2020_12_31_424971 685 4 1155 1155 CD 10_1101-2020_12_31_424971 685 5 enable enable VBP 10_1101-2020_12_31_424971 685 6 codon codon NNP 10_1101-2020_12_31_424971 685 7 reassignment reassignment NN 10_1101-2020_12_31_424971 685 8 of of IN 10_1101-2020_12_31_424971 685 9 selected select VBN 10_1101-2020_12_31_424971 685 10 mRNAs mRNAs '' 10_1101-2020_12_31_424971 685 11 in in IN 10_1101-2020_12_31_424971 685 12 eukaryotes eukaryote NNS 10_1101-2020_12_31_424971 685 13 . . . 10_1101-2020_12_31_424971 686 1 Science science NN 10_1101-2020_12_31_424971 686 2 363(2019 363(2019 NNP 10_1101-2020_12_31_424971 686 3 ) ) -RRB- 10_1101-2020_12_31_424971 686 4 . . . 10_1101-2020_12_31_424971 687 1 1156 1156 CD 10_1101-2020_12_31_424971 687 2 59 59 CD 10_1101-2020_12_31_424971 687 3 . . . 10_1101-2020_12_31_424971 688 1 Datsenko Datsenko NNP 10_1101-2020_12_31_424971 688 2 , , , 10_1101-2020_12_31_424971 688 3 K.A. K.A. NNP 10_1101-2020_12_31_424971 689 1 & & CC 10_1101-2020_12_31_424971 689 2 Wanner Wanner NNP 10_1101-2020_12_31_424971 689 3 , , , 10_1101-2020_12_31_424971 689 4 B.L. B.L. NNP 10_1101-2020_12_31_424971 690 1 One one CD 10_1101-2020_12_31_424971 690 2 - - HYPH 10_1101-2020_12_31_424971 690 3 step step NN 10_1101-2020_12_31_424971 690 4 inactivation inactivation NN 10_1101-2020_12_31_424971 690 5 of of IN 10_1101-2020_12_31_424971 690 6 chromosomal chromosomal JJ 10_1101-2020_12_31_424971 690 7 genes gene NNS 10_1101-2020_12_31_424971 690 8 in in IN 10_1101-2020_12_31_424971 690 9 1157 1157 CD 10_1101-2020_12_31_424971 690 10 Escherichia Escherichia NNP 10_1101-2020_12_31_424971 690 11 coli coli NNS 10_1101-2020_12_31_424971 690 12 K-12 k-12 CD 10_1101-2020_12_31_424971 690 13 using use VBG 10_1101-2020_12_31_424971 690 14 PCR PCR NNP 10_1101-2020_12_31_424971 690 15 products product NNS 10_1101-2020_12_31_424971 690 16 . . . 10_1101-2020_12_31_424971 691 1 Proc Proc NNP 10_1101-2020_12_31_424971 691 2 Natl Natl NNP 10_1101-2020_12_31_424971 691 3 Acad Acad NNP 10_1101-2020_12_31_424971 691 4 Sci Sci NNP 10_1101-2020_12_31_424971 691 5 U U NNP 10_1101-2020_12_31_424971 691 6 S S NNP 10_1101-2020_12_31_424971 691 7 A A NNP 10_1101-2020_12_31_424971 691 8 97 97 CD 10_1101-2020_12_31_424971 691 9 , , , 10_1101-2020_12_31_424971 691 10 6640 6640 CD 10_1101-2020_12_31_424971 691 11 - - SYM 10_1101-2020_12_31_424971 691 12 5 5 CD 10_1101-2020_12_31_424971 691 13 ( ( -LRB- 10_1101-2020_12_31_424971 691 14 2000 2000 CD 10_1101-2020_12_31_424971 691 15 ) ) -RRB- 10_1101-2020_12_31_424971 691 16 . . . 10_1101-2020_12_31_424971 692 1 1158 1158 CD 10_1101-2020_12_31_424971 692 2 60 60 CD 10_1101-2020_12_31_424971 692 3 . . . 10_1101-2020_12_31_424971 693 1 Fei Fei NNP 10_1101-2020_12_31_424971 693 2 , , , 10_1101-2020_12_31_424971 693 3 J. J. NNP 10_1101-2020_12_31_424971 693 4 et et NNP 10_1101-2020_12_31_424971 693 5 al al NNP 10_1101-2020_12_31_424971 693 6 . . . 10_1101-2020_12_31_424971 694 1 A a DT 10_1101-2020_12_31_424971 694 2 highly highly RB 10_1101-2020_12_31_424971 694 3 purified purified JJ 10_1101-2020_12_31_424971 694 4 , , , 10_1101-2020_12_31_424971 694 5 fluorescently fluorescently RB 10_1101-2020_12_31_424971 694 6 labeled label VBN 10_1101-2020_12_31_424971 694 7 in in IN 10_1101-2020_12_31_424971 694 8 vitro vitro FW 10_1101-2020_12_31_424971 694 9 translation translation NN 10_1101-2020_12_31_424971 694 10 system system NN 10_1101-2020_12_31_424971 694 11 for for IN 10_1101-2020_12_31_424971 694 12 single-1159 single-1159 NN 10_1101-2020_12_31_424971 694 13 molecule molecule JJ 10_1101-2020_12_31_424971 694 14 studies study NNS 10_1101-2020_12_31_424971 694 15 of of IN 10_1101-2020_12_31_424971 694 16 protein protein NN 10_1101-2020_12_31_424971 694 17 synthesis synthesis NN 10_1101-2020_12_31_424971 694 18 . . . 10_1101-2020_12_31_424971 695 1 Methods Methods NNP 10_1101-2020_12_31_424971 695 2 Enzymol enzymol JJ 10_1101-2020_12_31_424971 695 3 472 472 CD 10_1101-2020_12_31_424971 695 4 , , , 10_1101-2020_12_31_424971 695 5 221 221 CD 10_1101-2020_12_31_424971 695 6 - - SYM 10_1101-2020_12_31_424971 695 7 59 59 CD 10_1101-2020_12_31_424971 695 8 ( ( -LRB- 10_1101-2020_12_31_424971 695 9 2010 2010 CD 10_1101-2020_12_31_424971 695 10 ) ) -RRB- 10_1101-2020_12_31_424971 695 11 . . . 10_1101-2020_12_31_424971 696 1 1160 1160 CD 10_1101-2020_12_31_424971 696 2 61 61 CD 10_1101-2020_12_31_424971 696 3 . . . 10_1101-2020_12_31_424971 697 1 Christian Christian NNP 10_1101-2020_12_31_424971 697 2 , , , 10_1101-2020_12_31_424971 697 3 T. T. NNP 10_1101-2020_12_31_424971 697 4 , , , 10_1101-2020_12_31_424971 697 5 Lahoud Lahoud NNP 10_1101-2020_12_31_424971 697 6 , , , 10_1101-2020_12_31_424971 697 7 G. G. NNP 10_1101-2020_12_31_424971 697 8 , , , 10_1101-2020_12_31_424971 697 9 Liu Liu NNP 10_1101-2020_12_31_424971 697 10 , , , 10_1101-2020_12_31_424971 697 11 C. C. NNP 10_1101-2020_12_31_424971 697 12 & & CC 10_1101-2020_12_31_424971 697 13 Hou Hou NNP 10_1101-2020_12_31_424971 697 14 , , , 10_1101-2020_12_31_424971 697 15 Y.M. Y.M. NNP 10_1101-2020_12_31_424971 698 1 Control control NN 10_1101-2020_12_31_424971 698 2 of of IN 10_1101-2020_12_31_424971 698 3 catalytic catalytic JJ 10_1101-2020_12_31_424971 698 4 cycle cycle NN 10_1101-2020_12_31_424971 698 5 by by IN 10_1101-2020_12_31_424971 698 6 a a DT 10_1101-2020_12_31_424971 698 7 pair pair NN 10_1101-2020_12_31_424971 698 8 of of IN 10_1101-2020_12_31_424971 698 9 1161 1161 CD 10_1101-2020_12_31_424971 698 10 analogous analogous JJ 10_1101-2020_12_31_424971 698 11 tRNA trna NN 10_1101-2020_12_31_424971 698 12 modification modification NN 10_1101-2020_12_31_424971 698 13 enzymes enzyme NNS 10_1101-2020_12_31_424971 698 14 . . . 10_1101-2020_12_31_424971 699 1 J J NNP 10_1101-2020_12_31_424971 699 2 Mol Mol NNP 10_1101-2020_12_31_424971 699 3 Biol Biol NNP 10_1101-2020_12_31_424971 699 4 400 400 CD 10_1101-2020_12_31_424971 699 5 , , , 10_1101-2020_12_31_424971 699 6 204 204 CD 10_1101-2020_12_31_424971 699 7 - - SYM 10_1101-2020_12_31_424971 699 8 17 17 CD 10_1101-2020_12_31_424971 699 9 ( ( -LRB- 10_1101-2020_12_31_424971 699 10 2010 2010 CD 10_1101-2020_12_31_424971 699 11 ) ) -RRB- 10_1101-2020_12_31_424971 699 12 . . . 10_1101-2020_12_31_424971 700 1 1162 1162 CD 10_1101-2020_12_31_424971 700 2 62 62 CD 10_1101-2020_12_31_424971 700 3 . . . 10_1101-2020_12_31_424971 701 1 Zhang Zhang NNP 10_1101-2020_12_31_424971 701 2 , , , 10_1101-2020_12_31_424971 701 3 C.M. C.M. NNP 10_1101-2020_12_31_424971 701 4 , , , 10_1101-2020_12_31_424971 701 5 Perona Perona NNP 10_1101-2020_12_31_424971 701 6 , , , 10_1101-2020_12_31_424971 701 7 J.J. J.J. NNP 10_1101-2020_12_31_424971 701 8 , , , 10_1101-2020_12_31_424971 701 9 Ryu Ryu NNP 10_1101-2020_12_31_424971 701 10 , , , 10_1101-2020_12_31_424971 701 11 K. K. NNP 10_1101-2020_12_31_424971 701 12 , , , 10_1101-2020_12_31_424971 701 13 Francklyn Francklyn NNP 10_1101-2020_12_31_424971 701 14 , , , 10_1101-2020_12_31_424971 701 15 C. C. NNP 10_1101-2020_12_31_424971 701 16 & & CC 10_1101-2020_12_31_424971 701 17 Hou Hou NNP 10_1101-2020_12_31_424971 701 18 , , , 10_1101-2020_12_31_424971 701 19 Y.M. Y.M. NNP 10_1101-2020_12_31_424971 702 1 Distinct distinct JJ 10_1101-2020_12_31_424971 702 2 kinetic kinetic JJ 10_1101-2020_12_31_424971 702 3 1163 1163 CD 10_1101-2020_12_31_424971 702 4 mechanisms mechanism NNS 10_1101-2020_12_31_424971 702 5 of of IN 10_1101-2020_12_31_424971 702 6 the the DT 10_1101-2020_12_31_424971 702 7 two two CD 10_1101-2020_12_31_424971 702 8 classes class NNS 10_1101-2020_12_31_424971 702 9 of of IN 10_1101-2020_12_31_424971 702 10 Aminoacyl Aminoacyl NNP 10_1101-2020_12_31_424971 702 11 - - HYPH 10_1101-2020_12_31_424971 702 12 tRNA trna NN 10_1101-2020_12_31_424971 702 13 synthetases synthetase NNS 10_1101-2020_12_31_424971 702 14 . . . 10_1101-2020_12_31_424971 703 1 J J NNP 10_1101-2020_12_31_424971 703 2 Mol Mol NNP 10_1101-2020_12_31_424971 703 3 Biol Biol NNP 10_1101-2020_12_31_424971 703 4 361 361 CD 10_1101-2020_12_31_424971 703 5 , , , 10_1101-2020_12_31_424971 703 6 300 300 CD 10_1101-2020_12_31_424971 703 7 - - SYM 10_1101-2020_12_31_424971 703 8 11 11 CD 10_1101-2020_12_31_424971 703 9 1164 1164 CD 10_1101-2020_12_31_424971 703 10 ( ( -LRB- 10_1101-2020_12_31_424971 703 11 2006 2006 CD 10_1101-2020_12_31_424971 703 12 ) ) -RRB- 10_1101-2020_12_31_424971 703 13 . . . 10_1101-2020_12_31_424971 704 1 1165 1165 CD 10_1101-2020_12_31_424971 704 2 63 63 CD 10_1101-2020_12_31_424971 704 3 . . . 10_1101-2020_12_31_424971 705 1 Peacock Peacock NNP 10_1101-2020_12_31_424971 705 2 , , , 10_1101-2020_12_31_424971 705 3 J.R. J.R. NNP 10_1101-2020_12_31_424971 705 4 et et NNP 10_1101-2020_12_31_424971 705 5 al al NNP 10_1101-2020_12_31_424971 705 6 . . . 10_1101-2020_12_31_424971 706 1 Amino amino JJ 10_1101-2020_12_31_424971 706 2 acid acid NN 10_1101-2020_12_31_424971 706 3 - - HYPH 10_1101-2020_12_31_424971 706 4 dependent dependent JJ 10_1101-2020_12_31_424971 706 5 stability stability NN 10_1101-2020_12_31_424971 706 6 of of IN 10_1101-2020_12_31_424971 706 7 the the DT 10_1101-2020_12_31_424971 706 8 acyl acyl NN 10_1101-2020_12_31_424971 706 9 linkage linkage NN 10_1101-2020_12_31_424971 706 10 in in IN 10_1101-2020_12_31_424971 706 11 aminoacyl-1166 aminoacyl-1166 NNP 10_1101-2020_12_31_424971 706 12 tRNA trna NN 10_1101-2020_12_31_424971 706 13 . . . 10_1101-2020_12_31_424971 707 1 RNA RNA NNP 10_1101-2020_12_31_424971 707 2 20 20 CD 10_1101-2020_12_31_424971 707 3 , , , 10_1101-2020_12_31_424971 707 4 758 758 CD 10_1101-2020_12_31_424971 707 5 - - SYM 10_1101-2020_12_31_424971 707 6 64 64 CD 10_1101-2020_12_31_424971 707 7 ( ( -LRB- 10_1101-2020_12_31_424971 707 8 2014 2014 CD 10_1101-2020_12_31_424971 707 9 ) ) -RRB- 10_1101-2020_12_31_424971 707 10 . . . 10_1101-2020_12_31_424971 708 1 1167 1167 CD 10_1101-2020_12_31_424971 708 2 .CC .CC : 10_1101-2020_12_31_424971 708 3 - - HYPH 10_1101-2020_12_31_424971 708 4 BY by IN 10_1101-2020_12_31_424971 708 5 - - HYPH 10_1101-2020_12_31_424971 708 6 NC NC NNP 10_1101-2020_12_31_424971 708 7 - - HYPH 10_1101-2020_12_31_424971 708 8 ND ND NNP 10_1101-2020_12_31_424971 708 9 4.0 4.0 CD 10_1101-2020_12_31_424971 708 10 International International NNP 10_1101-2020_12_31_424971 708 11 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 708 12 under under IN 10_1101-2020_12_31_424971 708 13 a a DT 10_1101-2020_12_31_424971 708 14 ( ( -LRB- 10_1101-2020_12_31_424971 708 15 which which WDT 10_1101-2020_12_31_424971 708 16 was be VBD 10_1101-2020_12_31_424971 708 17 not not RB 10_1101-2020_12_31_424971 708 18 certified certify VBN 10_1101-2020_12_31_424971 708 19 by by IN 10_1101-2020_12_31_424971 708 20 peer peer NN 10_1101-2020_12_31_424971 708 21 review review NN 10_1101-2020_12_31_424971 708 22 ) ) -RRB- 10_1101-2020_12_31_424971 708 23 is be VBZ 10_1101-2020_12_31_424971 708 24 the the DT 10_1101-2020_12_31_424971 708 25 author author NN 10_1101-2020_12_31_424971 708 26 / / SYM 10_1101-2020_12_31_424971 708 27 funder funder NN 10_1101-2020_12_31_424971 708 28 , , , 10_1101-2020_12_31_424971 708 29 who who WP 10_1101-2020_12_31_424971 708 30 has have VBZ 10_1101-2020_12_31_424971 708 31 granted grant VBN 10_1101-2020_12_31_424971 708 32 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 708 33 a a DT 10_1101-2020_12_31_424971 708 34 license license NN 10_1101-2020_12_31_424971 708 35 to to TO 10_1101-2020_12_31_424971 708 36 display display VB 10_1101-2020_12_31_424971 708 37 the the DT 10_1101-2020_12_31_424971 708 38 preprint preprint NN 10_1101-2020_12_31_424971 708 39 in in IN 10_1101-2020_12_31_424971 708 40 perpetuity perpetuity NN 10_1101-2020_12_31_424971 708 41 . . . 10_1101-2020_12_31_424971 709 1 It -PRON- PRP 10_1101-2020_12_31_424971 709 2 is be VBZ 10_1101-2020_12_31_424971 709 3 made make VBN 10_1101-2020_12_31_424971 709 4 The the DT 10_1101-2020_12_31_424971 709 5 copyright copyright NN 10_1101-2020_12_31_424971 709 6 holder holder NN 10_1101-2020_12_31_424971 709 7 for for IN 10_1101-2020_12_31_424971 709 8 this this DT 10_1101-2020_12_31_424971 709 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 709 10 version version NN 10_1101-2020_12_31_424971 709 11 posted post VBD 10_1101-2020_12_31_424971 709 12 January January NNP 10_1101-2020_12_31_424971 709 13 2 2 CD 10_1101-2020_12_31_424971 709 14 , , , 10_1101-2020_12_31_424971 709 15 2021 2021 CD 10_1101-2020_12_31_424971 709 16 . . . 10_1101-2020_12_31_424971 709 17 ; ; : 10_1101-2020_12_31_424971 709 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 709 19 : : : 10_1101-2020_12_31_424971 709 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 709 21 preprint preprint NN 10_1101-2020_12_31_424971 709 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 709 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 709 24 45 45 CD 10_1101-2020_12_31_424971 709 25 64 64 CD 10_1101-2020_12_31_424971 709 26 . . . 10_1101-2020_12_31_424971 710 1 Aitken Aitken NNP 10_1101-2020_12_31_424971 710 2 , , , 10_1101-2020_12_31_424971 710 3 C.E. C.E. NNP 10_1101-2020_12_31_424971 710 4 , , , 10_1101-2020_12_31_424971 710 5 Marshall Marshall NNP 10_1101-2020_12_31_424971 710 6 , , , 10_1101-2020_12_31_424971 710 7 R.A. R.A. NNP 10_1101-2020_12_31_424971 710 8 & & CC 10_1101-2020_12_31_424971 710 9 Puglisi Puglisi NNP 10_1101-2020_12_31_424971 710 10 , , , 10_1101-2020_12_31_424971 710 11 J.D. J.D. NNP 10_1101-2020_12_31_424971 711 1 An an DT 10_1101-2020_12_31_424971 711 2 oxygen oxygen NN 10_1101-2020_12_31_424971 711 3 scavenging scavenge VBG 10_1101-2020_12_31_424971 711 4 system system NN 10_1101-2020_12_31_424971 711 5 for for IN 10_1101-2020_12_31_424971 711 6 1168 1168 CD 10_1101-2020_12_31_424971 711 7 improvement improvement NN 10_1101-2020_12_31_424971 711 8 of of IN 10_1101-2020_12_31_424971 711 9 dye dye NN 10_1101-2020_12_31_424971 711 10 stability stability NN 10_1101-2020_12_31_424971 711 11 in in IN 10_1101-2020_12_31_424971 711 12 single single JJ 10_1101-2020_12_31_424971 711 13 - - HYPH 10_1101-2020_12_31_424971 711 14 molecule molecule NN 10_1101-2020_12_31_424971 711 15 fluorescence fluorescence NN 10_1101-2020_12_31_424971 711 16 experiments experiment NNS 10_1101-2020_12_31_424971 711 17 . . . 10_1101-2020_12_31_424971 712 1 Biophys Biophys NNP 10_1101-2020_12_31_424971 712 2 J J NNP 10_1101-2020_12_31_424971 712 3 94 94 CD 10_1101-2020_12_31_424971 712 4 , , , 10_1101-2020_12_31_424971 712 5 1169 1169 CD 10_1101-2020_12_31_424971 712 6 1826 1826 CD 10_1101-2020_12_31_424971 712 7 - - SYM 10_1101-2020_12_31_424971 712 8 35 35 CD 10_1101-2020_12_31_424971 712 9 ( ( -LRB- 10_1101-2020_12_31_424971 712 10 2008 2008 CD 10_1101-2020_12_31_424971 712 11 ) ) -RRB- 10_1101-2020_12_31_424971 712 12 . . . 10_1101-2020_12_31_424971 713 1 1170 1170 CD 10_1101-2020_12_31_424971 713 2 65 65 CD 10_1101-2020_12_31_424971 713 3 . . . 10_1101-2020_12_31_424971 714 1 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 714 2 , , , 10_1101-2020_12_31_424971 714 3 R.L. R.L. NNP 10_1101-2020_12_31_424971 714 4 , , , 10_1101-2020_12_31_424971 714 5 Jr. Jr. NNP 10_1101-2020_12_31_424971 714 6 , , , 10_1101-2020_12_31_424971 714 7 Chu Chu NNP 10_1101-2020_12_31_424971 714 8 , , , 10_1101-2020_12_31_424971 714 9 S. S. NNP 10_1101-2020_12_31_424971 714 10 & & CC 10_1101-2020_12_31_424971 714 11 Puglisi Puglisi NNP 10_1101-2020_12_31_424971 714 12 , , , 10_1101-2020_12_31_424971 714 13 J.D. J.D. NNP 10_1101-2020_12_31_424971 715 1 Thiostrepton Thiostrepton NNP 10_1101-2020_12_31_424971 715 2 inhibition inhibition NN 10_1101-2020_12_31_424971 715 3 of of IN 10_1101-2020_12_31_424971 715 4 tRNA trna NN 10_1101-2020_12_31_424971 715 5 delivery delivery NN 10_1101-2020_12_31_424971 715 6 to to IN 10_1101-2020_12_31_424971 715 7 the the DT 10_1101-2020_12_31_424971 715 8 1171 1171 CD 10_1101-2020_12_31_424971 715 9 ribosome ribosome VBD 10_1101-2020_12_31_424971 715 10 . . . 10_1101-2020_12_31_424971 716 1 RNA RNA NNP 10_1101-2020_12_31_424971 716 2 13 13 CD 10_1101-2020_12_31_424971 716 3 , , , 10_1101-2020_12_31_424971 716 4 2091 2091 CD 10_1101-2020_12_31_424971 716 5 - - SYM 10_1101-2020_12_31_424971 716 6 7 7 CD 10_1101-2020_12_31_424971 716 7 ( ( -LRB- 10_1101-2020_12_31_424971 716 8 2007 2007 CD 10_1101-2020_12_31_424971 716 9 ) ) -RRB- 10_1101-2020_12_31_424971 716 10 . . . 10_1101-2020_12_31_424971 717 1 1172 1172 CD 10_1101-2020_12_31_424971 717 2 66 66 CD 10_1101-2020_12_31_424971 717 3 . . . 10_1101-2020_12_31_424971 718 1 Desai Desai NNP 10_1101-2020_12_31_424971 718 2 , , , 10_1101-2020_12_31_424971 718 3 B.J. B.J. NNP 10_1101-2020_12_31_424971 719 1 & & CC 10_1101-2020_12_31_424971 719 2 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 719 3 , , , 10_1101-2020_12_31_424971 719 4 R.L. R.L. NNP 10_1101-2020_12_31_424971 719 5 , , , 10_1101-2020_12_31_424971 719 6 Jr. Jr. NNP 10_1101-2020_12_31_424971 719 7 Multiplexed Multiplexed NNP 10_1101-2020_12_31_424971 719 8 , , , 10_1101-2020_12_31_424971 719 9 bioorthogonal bioorthogonal JJ 10_1101-2020_12_31_424971 719 10 labeling labeling NN 10_1101-2020_12_31_424971 719 11 of of IN 10_1101-2020_12_31_424971 719 12 multicomponent multicomponent NN 10_1101-2020_12_31_424971 719 13 , , , 10_1101-2020_12_31_424971 719 14 1173 1173 CD 10_1101-2020_12_31_424971 719 15 biomolecular biomolecular JJ 10_1101-2020_12_31_424971 719 16 complexes complex NNS 10_1101-2020_12_31_424971 719 17 using use VBG 10_1101-2020_12_31_424971 719 18 genomically genomically RB 10_1101-2020_12_31_424971 719 19 encoded encode VBN 10_1101-2020_12_31_424971 719 20 , , , 10_1101-2020_12_31_424971 719 21 non non JJ 10_1101-2020_12_31_424971 719 22 - - JJ 10_1101-2020_12_31_424971 719 23 canonical canonical JJ 10_1101-2020_12_31_424971 719 24 amino amino NN 10_1101-2020_12_31_424971 719 25 acids acid NNS 10_1101-2020_12_31_424971 719 26 . . . 10_1101-2020_12_31_424971 720 1 1174 1174 CD 10_1101-2020_12_31_424971 720 2 bioRxiv biorxiv NN 10_1101-2020_12_31_424971 720 3 doi doi NN 10_1101-2020_12_31_424971 720 4 : : : 10_1101-2020_12_31_424971 720 5 10.1101/730465(2019 10.1101/730465(2019 LS 10_1101-2020_12_31_424971 720 6 ) ) -RRB- 10_1101-2020_12_31_424971 720 7 . . . 10_1101-2020_12_31_424971 721 1 1175 1175 CD 10_1101-2020_12_31_424971 721 2 67 67 CD 10_1101-2020_12_31_424971 721 3 . . . 10_1101-2020_12_31_424971 722 1 MacDougall MacDougall NNP 10_1101-2020_12_31_424971 722 2 , , , 10_1101-2020_12_31_424971 722 3 D.D. D.D. NNP 10_1101-2020_12_31_424971 723 1 & & CC 10_1101-2020_12_31_424971 723 2 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 723 3 , , , 10_1101-2020_12_31_424971 723 4 R.L. R.L. NNP 10_1101-2020_12_31_424971 723 5 , , , 10_1101-2020_12_31_424971 723 6 Jr. Jr. NNP 10_1101-2020_12_31_424971 724 1 Translation translation NN 10_1101-2020_12_31_424971 724 2 initiation initiation NN 10_1101-2020_12_31_424971 724 3 factor factor NN 10_1101-2020_12_31_424971 724 4 3 3 CD 10_1101-2020_12_31_424971 724 5 regulates regulate NNS 10_1101-2020_12_31_424971 724 6 switching switch VBG 10_1101-2020_12_31_424971 724 7 1176 1176 CD 10_1101-2020_12_31_424971 724 8 between between IN 10_1101-2020_12_31_424971 724 9 different different JJ 10_1101-2020_12_31_424971 724 10 modes mode NNS 10_1101-2020_12_31_424971 724 11 of of IN 10_1101-2020_12_31_424971 724 12 ribosomal ribosomal NNP 10_1101-2020_12_31_424971 724 13 subunit subunit NN 10_1101-2020_12_31_424971 724 14 joining joining NN 10_1101-2020_12_31_424971 724 15 . . . 10_1101-2020_12_31_424971 725 1 J J NNP 10_1101-2020_12_31_424971 725 2 Mol Mol NNP 10_1101-2020_12_31_424971 725 3 Biol Biol NNP 10_1101-2020_12_31_424971 725 4 427 427 CD 10_1101-2020_12_31_424971 725 5 , , , 10_1101-2020_12_31_424971 725 6 1801 1801 CD 10_1101-2020_12_31_424971 725 7 - - SYM 10_1101-2020_12_31_424971 725 8 18 18 CD 10_1101-2020_12_31_424971 725 9 ( ( -LRB- 10_1101-2020_12_31_424971 725 10 2015 2015 CD 10_1101-2020_12_31_424971 725 11 ) ) -RRB- 10_1101-2020_12_31_424971 725 12 . . . 10_1101-2020_12_31_424971 726 1 1177 1177 CD 10_1101-2020_12_31_424971 726 2 68 68 CD 10_1101-2020_12_31_424971 726 3 . . . 10_1101-2020_12_31_424971 727 1 Bronson Bronson NNP 10_1101-2020_12_31_424971 727 2 , , , 10_1101-2020_12_31_424971 727 3 J.E. J.E. NNP 10_1101-2020_12_31_424971 727 4 , , , 10_1101-2020_12_31_424971 727 5 Fei Fei NNP 10_1101-2020_12_31_424971 727 6 , , , 10_1101-2020_12_31_424971 727 7 J. J. NNP 10_1101-2020_12_31_424971 727 8 , , , 10_1101-2020_12_31_424971 727 9 Hofman Hofman NNP 10_1101-2020_12_31_424971 727 10 , , , 10_1101-2020_12_31_424971 727 11 J.M. J.M. NNP 10_1101-2020_12_31_424971 727 12 , , , 10_1101-2020_12_31_424971 727 13 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 727 14 , , , 10_1101-2020_12_31_424971 727 15 R.L. R.L. NNP 10_1101-2020_12_31_424971 727 16 , , , 10_1101-2020_12_31_424971 727 17 Jr. Jr. NNP 10_1101-2020_12_31_424971 727 18 & & CC 10_1101-2020_12_31_424971 727 19 Wiggins Wiggins NNP 10_1101-2020_12_31_424971 727 20 , , , 10_1101-2020_12_31_424971 727 21 C.H. C.H. NNP 10_1101-2020_12_31_424971 728 1 Learning learning NN 10_1101-2020_12_31_424971 728 2 rates rate NNS 10_1101-2020_12_31_424971 728 3 1178 1178 CD 10_1101-2020_12_31_424971 728 4 and and CC 10_1101-2020_12_31_424971 728 5 states state NNS 10_1101-2020_12_31_424971 728 6 from from IN 10_1101-2020_12_31_424971 728 7 biophysical biophysical JJ 10_1101-2020_12_31_424971 728 8 time time NN 10_1101-2020_12_31_424971 728 9 series series NNP 10_1101-2020_12_31_424971 728 10 : : : 10_1101-2020_12_31_424971 728 11 a a DT 10_1101-2020_12_31_424971 728 12 Bayesian bayesian JJ 10_1101-2020_12_31_424971 728 13 approach approach NN 10_1101-2020_12_31_424971 728 14 to to IN 10_1101-2020_12_31_424971 728 15 model model NN 10_1101-2020_12_31_424971 728 16 selection selection NN 10_1101-2020_12_31_424971 728 17 and and CC 10_1101-2020_12_31_424971 728 18 1179 1179 CD 10_1101-2020_12_31_424971 728 19 single single JJ 10_1101-2020_12_31_424971 728 20 - - HYPH 10_1101-2020_12_31_424971 728 21 molecule molecule NN 10_1101-2020_12_31_424971 728 22 FRET FRET NNP 10_1101-2020_12_31_424971 728 23 data datum NNS 10_1101-2020_12_31_424971 728 24 . . . 10_1101-2020_12_31_424971 729 1 Biophys Biophys NNP 10_1101-2020_12_31_424971 729 2 J J NNP 10_1101-2020_12_31_424971 729 3 97 97 CD 10_1101-2020_12_31_424971 729 4 , , , 10_1101-2020_12_31_424971 729 5 3196 3196 CD 10_1101-2020_12_31_424971 729 6 - - SYM 10_1101-2020_12_31_424971 729 7 205 205 CD 10_1101-2020_12_31_424971 729 8 ( ( -LRB- 10_1101-2020_12_31_424971 729 9 2009 2009 CD 10_1101-2020_12_31_424971 729 10 ) ) -RRB- 10_1101-2020_12_31_424971 729 11 . . . 10_1101-2020_12_31_424971 730 1 1180 1180 CD 10_1101-2020_12_31_424971 730 2 69 69 CD 10_1101-2020_12_31_424971 730 3 . . . 10_1101-2020_12_31_424971 731 1 Viterbi Viterbi NNP 10_1101-2020_12_31_424971 731 2 , , , 10_1101-2020_12_31_424971 731 3 A.J. A.J. NNP 10_1101-2020_12_31_424971 732 1 Error error NN 10_1101-2020_12_31_424971 732 2 bounds bound NNS 10_1101-2020_12_31_424971 732 3 for for IN 10_1101-2020_12_31_424971 732 4 convolutional convolutional JJ 10_1101-2020_12_31_424971 732 5 codes code NNS 10_1101-2020_12_31_424971 732 6 and and CC 10_1101-2020_12_31_424971 732 7 an an DT 10_1101-2020_12_31_424971 732 8 asymptotically asymptotically RB 10_1101-2020_12_31_424971 732 9 optimum optimum JJ 10_1101-2020_12_31_424971 732 10 1181 1181 CD 10_1101-2020_12_31_424971 732 11 decoding decode VBG 10_1101-2020_12_31_424971 732 12 algorithm algorithm NNP 10_1101-2020_12_31_424971 732 13 . . . 10_1101-2020_12_31_424971 733 1 IEEE IEEE NNP 10_1101-2020_12_31_424971 733 2 Trans Trans NNPS 10_1101-2020_12_31_424971 733 3 . . . 10_1101-2020_12_31_424971 734 1 Inform inform NN 10_1101-2020_12_31_424971 734 2 . . . 10_1101-2020_12_31_424971 735 1 Theory theory NN 10_1101-2020_12_31_424971 735 2 13 13 CD 10_1101-2020_12_31_424971 735 3 , , , 10_1101-2020_12_31_424971 735 4 260 260 CD 10_1101-2020_12_31_424971 735 5 - - HYPH 10_1101-2020_12_31_424971 735 6 269 269 CD 10_1101-2020_12_31_424971 735 7 ( ( -LRB- 10_1101-2020_12_31_424971 735 8 1967 1967 CD 10_1101-2020_12_31_424971 735 9 ) ) -RRB- 10_1101-2020_12_31_424971 735 10 . . . 10_1101-2020_12_31_424971 736 1 1182 1182 CD 10_1101-2020_12_31_424971 736 2 70 70 CD 10_1101-2020_12_31_424971 736 3 . . . 10_1101-2020_12_31_424971 737 1 Agirrezabala Agirrezabala NNP 10_1101-2020_12_31_424971 737 2 , , , 10_1101-2020_12_31_424971 737 3 X. X. NNP 10_1101-2020_12_31_424971 737 4 et et NNP 10_1101-2020_12_31_424971 737 5 al al NNP 10_1101-2020_12_31_424971 737 6 . . . 10_1101-2020_12_31_424971 738 1 Visualization visualization NN 10_1101-2020_12_31_424971 738 2 of of IN 10_1101-2020_12_31_424971 738 3 the the DT 10_1101-2020_12_31_424971 738 4 hybrid hybrid JJ 10_1101-2020_12_31_424971 738 5 state state NN 10_1101-2020_12_31_424971 738 6 of of IN 10_1101-2020_12_31_424971 738 7 tRNA trna NN 10_1101-2020_12_31_424971 738 8 binding bind VBG 10_1101-2020_12_31_424971 738 9 promoted promote VBN 10_1101-2020_12_31_424971 738 10 by by IN 10_1101-2020_12_31_424971 738 11 1183 1183 CD 10_1101-2020_12_31_424971 738 12 spontaneous spontaneous JJ 10_1101-2020_12_31_424971 738 13 ratcheting ratcheting NN 10_1101-2020_12_31_424971 738 14 of of IN 10_1101-2020_12_31_424971 738 15 the the DT 10_1101-2020_12_31_424971 738 16 ribosome ribosome NN 10_1101-2020_12_31_424971 738 17 . . . 10_1101-2020_12_31_424971 739 1 Mol Mol NNP 10_1101-2020_12_31_424971 739 2 Cell Cell NNP 10_1101-2020_12_31_424971 739 3 32 32 CD 10_1101-2020_12_31_424971 739 4 , , , 10_1101-2020_12_31_424971 739 5 190 190 CD 10_1101-2020_12_31_424971 739 6 - - SYM 10_1101-2020_12_31_424971 739 7 7 7 CD 10_1101-2020_12_31_424971 739 8 ( ( -LRB- 10_1101-2020_12_31_424971 739 9 2008 2008 CD 10_1101-2020_12_31_424971 739 10 ) ) -RRB- 10_1101-2020_12_31_424971 739 11 . . . 10_1101-2020_12_31_424971 740 1 1184 1184 CD 10_1101-2020_12_31_424971 740 2 1185 1185 CD 10_1101-2020_12_31_424971 740 3 1186 1186 CD 10_1101-2020_12_31_424971 740 4 .CC .CC : 10_1101-2020_12_31_424971 740 5 - - HYPH 10_1101-2020_12_31_424971 740 6 BY by IN 10_1101-2020_12_31_424971 740 7 - - HYPH 10_1101-2020_12_31_424971 740 8 NC NC NNP 10_1101-2020_12_31_424971 740 9 - - HYPH 10_1101-2020_12_31_424971 740 10 ND ND NNP 10_1101-2020_12_31_424971 740 11 4.0 4.0 CD 10_1101-2020_12_31_424971 740 12 International International NNP 10_1101-2020_12_31_424971 740 13 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 740 14 under under IN 10_1101-2020_12_31_424971 740 15 a a DT 10_1101-2020_12_31_424971 740 16 ( ( -LRB- 10_1101-2020_12_31_424971 740 17 which which WDT 10_1101-2020_12_31_424971 740 18 was be VBD 10_1101-2020_12_31_424971 740 19 not not RB 10_1101-2020_12_31_424971 740 20 certified certify VBN 10_1101-2020_12_31_424971 740 21 by by IN 10_1101-2020_12_31_424971 740 22 peer peer NN 10_1101-2020_12_31_424971 740 23 review review NN 10_1101-2020_12_31_424971 740 24 ) ) -RRB- 10_1101-2020_12_31_424971 740 25 is be VBZ 10_1101-2020_12_31_424971 740 26 the the DT 10_1101-2020_12_31_424971 740 27 author author NN 10_1101-2020_12_31_424971 740 28 / / SYM 10_1101-2020_12_31_424971 740 29 funder funder NN 10_1101-2020_12_31_424971 740 30 , , , 10_1101-2020_12_31_424971 740 31 who who WP 10_1101-2020_12_31_424971 740 32 has have VBZ 10_1101-2020_12_31_424971 740 33 granted grant VBN 10_1101-2020_12_31_424971 740 34 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 740 35 a a DT 10_1101-2020_12_31_424971 740 36 license license NN 10_1101-2020_12_31_424971 740 37 to to TO 10_1101-2020_12_31_424971 740 38 display display VB 10_1101-2020_12_31_424971 740 39 the the DT 10_1101-2020_12_31_424971 740 40 preprint preprint NN 10_1101-2020_12_31_424971 740 41 in in IN 10_1101-2020_12_31_424971 740 42 perpetuity perpetuity NN 10_1101-2020_12_31_424971 740 43 . . . 10_1101-2020_12_31_424971 741 1 It -PRON- PRP 10_1101-2020_12_31_424971 741 2 is be VBZ 10_1101-2020_12_31_424971 741 3 made make VBN 10_1101-2020_12_31_424971 741 4 The the DT 10_1101-2020_12_31_424971 741 5 copyright copyright NN 10_1101-2020_12_31_424971 741 6 holder holder NN 10_1101-2020_12_31_424971 741 7 for for IN 10_1101-2020_12_31_424971 741 8 this this DT 10_1101-2020_12_31_424971 741 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 741 10 version version NN 10_1101-2020_12_31_424971 741 11 posted post VBD 10_1101-2020_12_31_424971 741 12 January January NNP 10_1101-2020_12_31_424971 741 13 2 2 CD 10_1101-2020_12_31_424971 741 14 , , , 10_1101-2020_12_31_424971 741 15 2021 2021 CD 10_1101-2020_12_31_424971 741 16 . . . 10_1101-2020_12_31_424971 741 17 ; ; : 10_1101-2020_12_31_424971 741 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 741 19 : : : 10_1101-2020_12_31_424971 741 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 741 21 preprint preprint NN 10_1101-2020_12_31_424971 741 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 741 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 741 24 46 46 CD 10_1101-2020_12_31_424971 741 25 ACKNOWLEDGEMENTS acknowledgement NNS 10_1101-2020_12_31_424971 741 26 1187 1187 CD 10_1101-2020_12_31_424971 741 27 We -PRON- PRP 10_1101-2020_12_31_424971 741 28 thank thank VBP 10_1101-2020_12_31_424971 741 29 Dr. Dr. NNP 10_1101-2020_12_31_424971 741 30 Hajime Hajime NNP 10_1101-2020_12_31_424971 741 31 Tokuda Tokuda NNP 10_1101-2020_12_31_424971 741 32 for for IN 10_1101-2020_12_31_424971 741 33 rabbit rabbit NN 10_1101-2020_12_31_424971 741 34 polyclonal polyclonal JJ 10_1101-2020_12_31_424971 741 35 anti anti JJ 10_1101-2020_12_31_424971 741 36 - - JJ 10_1101-2020_12_31_424971 741 37 LolB lolb JJ 10_1101-2020_12_31_424971 741 38 antibodies antibody NNS 10_1101-2020_12_31_424971 741 39 , , , 10_1101-2020_12_31_424971 741 40 Dr. Dr. NNP 10_1101-2020_12_31_424971 741 41 Colin Colin NNP 10_1101-2020_12_31_424971 741 42 Kinz-1188 Kinz-1188 NNP 10_1101-2020_12_31_424971 741 43 Thompson Thompson NNP 10_1101-2020_12_31_424971 741 44 and and CC 10_1101-2020_12_31_424971 741 45 Korak Korak NNP 10_1101-2020_12_31_424971 741 46 Kumar Kumar NNP 10_1101-2020_12_31_424971 741 47 Ray Ray NNP 10_1101-2020_12_31_424971 741 48 for for IN 10_1101-2020_12_31_424971 741 49 help help NN 10_1101-2020_12_31_424971 741 50 with with IN 10_1101-2020_12_31_424971 741 51 smFRET smFRET NNP 10_1101-2020_12_31_424971 741 52 data datum NNS 10_1101-2020_12_31_424971 741 53 analysis analysis NN 10_1101-2020_12_31_424971 741 54 . . . 10_1101-2020_12_31_424971 742 1 R.L.G. R.L.G. NNP 10_1101-2020_12_31_424971 743 1 and and CC 10_1101-2020_12_31_424971 743 2 H.L. H.L. NNP 10_1101-2020_12_31_424971 744 1 thank thank VBP 10_1101-2020_12_31_424971 744 2 the the DT 10_1101-2020_12_31_424971 744 3 1189 1189 CD 10_1101-2020_12_31_424971 744 4 Columbia Columbia NNP 10_1101-2020_12_31_424971 744 5 University University NNP 10_1101-2020_12_31_424971 744 6 Precision Precision NNP 10_1101-2020_12_31_424971 744 7 Biomolecular Biomolecular NNP 10_1101-2020_12_31_424971 744 8 Characterization Characterization NNP 10_1101-2020_12_31_424971 744 9 Facility Facility NNP 10_1101-2020_12_31_424971 744 10 for for IN 10_1101-2020_12_31_424971 744 11 access access NN 10_1101-2020_12_31_424971 744 12 to to IN 10_1101-2020_12_31_424971 744 13 and and CC 10_1101-2020_12_31_424971 744 14 support support NN 10_1101-2020_12_31_424971 744 15 of of IN 10_1101-2020_12_31_424971 744 16 1190 1190 CD 10_1101-2020_12_31_424971 744 17 instrumentation instrumentation NN 10_1101-2020_12_31_424971 744 18 . . . 10_1101-2020_12_31_424971 745 1 This this DT 10_1101-2020_12_31_424971 745 2 work work NN 10_1101-2020_12_31_424971 745 3 was be VBD 10_1101-2020_12_31_424971 745 4 supported support VBN 10_1101-2020_12_31_424971 745 5 by by IN 10_1101-2020_12_31_424971 745 6 NIH NIH NNP 10_1101-2020_12_31_424971 745 7 grants grant NNS 10_1101-2020_12_31_424971 745 8 GM134931 gm134931 ADD 10_1101-2020_12_31_424971 745 9 to to IN 10_1101-2020_12_31_424971 745 10 Y Y NNP 10_1101-2020_12_31_424971 745 11 - - HYPH 10_1101-2020_12_31_424971 745 12 M.H. M.H. NNP 10_1101-2020_12_31_424971 746 1 and and CC 10_1101-2020_12_31_424971 746 2 GM119386 GM119386 NNP 10_1101-2020_12_31_424971 746 3 to to IN 10_1101-2020_12_31_424971 746 4 1191 1191 CD 10_1101-2020_12_31_424971 746 5 R.L.G. R.L.G. NNP 10_1101-2020_12_31_424971 746 6 , , , 10_1101-2020_12_31_424971 746 7 a a DT 10_1101-2020_12_31_424971 746 8 Charles Charles NNP 10_1101-2020_12_31_424971 746 9 H. H. NNP 10_1101-2020_12_31_424971 746 10 Revson Revson NNP 10_1101-2020_12_31_424971 746 11 Foundation Foundation NNP 10_1101-2020_12_31_424971 746 12 Postdoctoral Postdoctoral NNP 10_1101-2020_12_31_424971 746 13 Fellowship Fellowship NNP 10_1101-2020_12_31_424971 746 14 in in IN 10_1101-2020_12_31_424971 746 15 Biomedical Biomedical NNP 10_1101-2020_12_31_424971 746 16 Science Science NNP 10_1101-2020_12_31_424971 746 17 19 19 CD 10_1101-2020_12_31_424971 746 18 - - SYM 10_1101-2020_12_31_424971 746 19 24 24 CD 10_1101-2020_12_31_424971 746 20 to to IN 10_1101-2020_12_31_424971 746 21 1192 1192 CD 10_1101-2020_12_31_424971 746 22 H.L. H.L. NNP 10_1101-2020_12_31_424971 746 23 , , , 10_1101-2020_12_31_424971 746 24 a a DT 10_1101-2020_12_31_424971 746 25 Japanese japanese JJ 10_1101-2020_12_31_424971 746 26 JSPS JSPS NNP 10_1101-2020_12_31_424971 746 27 overseas overseas JJ 10_1101-2020_12_31_424971 746 28 postdoctoral postdoctoral JJ 10_1101-2020_12_31_424971 746 29 fellowship fellowship NN 10_1101-2020_12_31_424971 746 30 to to IN 10_1101-2020_12_31_424971 746 31 I.M. I.M. NNP 10_1101-2020_12_31_424971 746 32 , , , 10_1101-2020_12_31_424971 746 33 and and CC 10_1101-2020_12_31_424971 746 34 NSF NSF NNP 10_1101-2020_12_31_424971 746 35 grant grant NN 10_1101-2020_12_31_424971 746 36 CHE-1708759 CHE-1708759 NNP 10_1101-2020_12_31_424971 746 37 to to IN 10_1101-2020_12_31_424971 746 38 1193 1193 CD 10_1101-2020_12_31_424971 746 39 E.J.P. E.J.P. NNP 10_1101-2020_12_31_424971 747 1 1194 1194 CD 10_1101-2020_12_31_424971 747 2 1195 1195 CD 10_1101-2020_12_31_424971 747 3 AUTHOR AUTHOR NNP 10_1101-2020_12_31_424971 747 4 CONTRIBUTIONS CONTRIBUTIONS NNP 10_1101-2020_12_31_424971 747 5 1196 1196 CD 10_1101-2020_12_31_424971 747 6 H.G. H.G. NNP 10_1101-2020_12_31_424971 748 1 conceived conceive VBN 10_1101-2020_12_31_424971 748 2 of of IN 10_1101-2020_12_31_424971 748 3 and and CC 10_1101-2020_12_31_424971 748 4 performed perform VBN 10_1101-2020_12_31_424971 748 5 ensemble ensemble JJ 10_1101-2020_12_31_424971 748 6 rapid rapid JJ 10_1101-2020_12_31_424971 748 7 kinetic kinetic NNP 10_1101-2020_12_31_424971 748 8 assays assay NNS 10_1101-2020_12_31_424971 748 9 , , , 10_1101-2020_12_31_424971 748 10 R.L.G. R.L.G. NNP 10_1101-2020_12_31_424971 749 1 and and CC 10_1101-2020_12_31_424971 749 2 H.L. H.L. NNP 10_1101-2020_12_31_424971 750 1 conceived conceive VBD 10_1101-2020_12_31_424971 750 2 1197 1197 CD 10_1101-2020_12_31_424971 750 3 of of IN 10_1101-2020_12_31_424971 750 4 and and CC 10_1101-2020_12_31_424971 750 5 designed design VBD 10_1101-2020_12_31_424971 750 6 smFRET smFRET NNP 10_1101-2020_12_31_424971 750 7 assays assay NNS 10_1101-2020_12_31_424971 750 8 , , , 10_1101-2020_12_31_424971 750 9 H.L. H.L. NNP 10_1101-2020_12_31_424971 751 1 performed perform VBN 10_1101-2020_12_31_424971 751 2 smFRET smFRET NNP 10_1101-2020_12_31_424971 751 3 assays assay NNS 10_1101-2020_12_31_424971 751 4 , , , 10_1101-2020_12_31_424971 751 5 I.M. I.M. NNP 10_1101-2020_12_31_424971 752 1 performed perform VBN 10_1101-2020_12_31_424971 752 2 cell cell NN 10_1101-2020_12_31_424971 752 3 - - HYPH 10_1101-2020_12_31_424971 752 4 based base VBN 10_1101-2020_12_31_424971 752 5 1198 1198 CD 10_1101-2020_12_31_424971 752 6 reporter reporter NN 10_1101-2020_12_31_424971 752 7 assays assays RB 10_1101-2020_12_31_424971 752 8 , , , 10_1101-2020_12_31_424971 752 9 D.M.R. D.M.R. NNP 10_1101-2020_12_31_424971 753 1 and and CC 10_1101-2020_12_31_424971 753 2 E.J.P. E.J.P. NNP 10_1101-2020_12_31_424971 754 1 generated generate VBN 10_1101-2020_12_31_424971 754 2 aminoacyl aminoacyl NNP 10_1101-2020_12_31_424971 754 3 - - HYPH 10_1101-2020_12_31_424971 754 4 DBE DBE NNP 10_1101-2020_12_31_424971 754 5 derivatives derivative NNS 10_1101-2020_12_31_424971 754 6 , , , 10_1101-2020_12_31_424971 754 7 T.C. T.C. NNP 10_1101-2020_12_31_424971 755 1 performed perform VBN 10_1101-2020_12_31_424971 755 2 G37 G37 NNP 10_1101-2020_12_31_424971 755 3 1199 1199 CD 10_1101-2020_12_31_424971 755 4 methylation methylation NN 10_1101-2020_12_31_424971 755 5 and and CC 10_1101-2020_12_31_424971 755 6 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 755 7 assays assays RB 10_1101-2020_12_31_424971 755 8 , , , 10_1101-2020_12_31_424971 755 9 and and CC 10_1101-2020_12_31_424971 755 10 A.B.C. A.B.C. NNP 10_1101-2020_12_31_424971 756 1 and and CC 10_1101-2020_12_31_424971 756 2 G.B. G.B. NNP 10_1101-2020_12_31_424971 757 1 provided provide VBN 10_1101-2020_12_31_424971 757 2 E. E. NNP 10_1101-2020_12_31_424971 757 3 coli coli NNS 10_1101-2020_12_31_424971 757 4 70S 70S NNP 10_1101-2020_12_31_424971 757 5 ribosomes ribosome NNS 10_1101-2020_12_31_424971 757 6 . . . 10_1101-2020_12_31_424971 758 1 1200 1200 CD 10_1101-2020_12_31_424971 758 2 Y.M.H. Y.M.H. NNP 10_1101-2020_12_31_424971 759 1 and and CC 10_1101-2020_12_31_424971 759 2 R.L.G. R.L.G. NNP 10_1101-2020_12_31_424971 760 1 wrote write VBD 10_1101-2020_12_31_424971 760 2 the the DT 10_1101-2020_12_31_424971 760 3 manuscript manuscript NN 10_1101-2020_12_31_424971 760 4 . . . 10_1101-2020_12_31_424971 761 1 1201 1201 CD 10_1101-2020_12_31_424971 761 2 1202 1202 CD 10_1101-2020_12_31_424971 761 3 COMPETING COMPETING NNP 10_1101-2020_12_31_424971 761 4 FINANICAL FINANICAL NNP 10_1101-2020_12_31_424971 761 5 INTERESTS INTERESTS NNP 10_1101-2020_12_31_424971 761 6 1203 1203 CD 10_1101-2020_12_31_424971 761 7 The the DT 10_1101-2020_12_31_424971 761 8 authors author NNS 10_1101-2020_12_31_424971 761 9 declare declare VBP 10_1101-2020_12_31_424971 761 10 no no DT 10_1101-2020_12_31_424971 761 11 competing compete VBG 10_1101-2020_12_31_424971 761 12 interests interest NNS 10_1101-2020_12_31_424971 761 13 . . . 10_1101-2020_12_31_424971 762 1 1204 1204 CD 10_1101-2020_12_31_424971 762 2 1205 1205 CD 10_1101-2020_12_31_424971 762 3 CONTACT contact NN 10_1101-2020_12_31_424971 762 4 FOR for IN 10_1101-2020_12_31_424971 762 5 REAGENT REAGENT NNP 10_1101-2020_12_31_424971 762 6 AND and CC 10_1101-2020_12_31_424971 762 7 RESOURCE resource NN 10_1101-2020_12_31_424971 762 8 SHARING sharing NN 10_1101-2020_12_31_424971 762 9 1206 1206 CD 10_1101-2020_12_31_424971 762 10 Further further JJ 10_1101-2020_12_31_424971 762 11 information information NN 10_1101-2020_12_31_424971 762 12 and and CC 10_1101-2020_12_31_424971 762 13 requests request NNS 10_1101-2020_12_31_424971 762 14 for for IN 10_1101-2020_12_31_424971 762 15 resources resource NNS 10_1101-2020_12_31_424971 762 16 and and CC 10_1101-2020_12_31_424971 762 17 reagents reagent NNS 10_1101-2020_12_31_424971 762 18 should should MD 10_1101-2020_12_31_424971 762 19 be be VB 10_1101-2020_12_31_424971 762 20 directed direct VBN 10_1101-2020_12_31_424971 762 21 to to IN 10_1101-2020_12_31_424971 762 22 and and CC 10_1101-2020_12_31_424971 762 23 will will MD 10_1101-2020_12_31_424971 762 24 be be VB 10_1101-2020_12_31_424971 762 25 1207 1207 CD 10_1101-2020_12_31_424971 762 26 fulfilled fulfil VBN 10_1101-2020_12_31_424971 762 27 by by IN 10_1101-2020_12_31_424971 762 28 the the DT 10_1101-2020_12_31_424971 762 29 lead lead NN 10_1101-2020_12_31_424971 762 30 contacts contact NNS 10_1101-2020_12_31_424971 762 31 Ruben Ruben NNP 10_1101-2020_12_31_424971 762 32 L. L. NNP 10_1101-2020_12_31_424971 762 33 Gonzalez Gonzalez NNP 10_1101-2020_12_31_424971 762 34 , , , 10_1101-2020_12_31_424971 762 35 Jr. Jr. NNP 10_1101-2020_12_31_424971 762 36 ( ( -LRB- 10_1101-2020_12_31_424971 762 37 rlg2118@columbia.edu rlg2118@columbia.edu NNP 10_1101-2020_12_31_424971 762 38 ) ) -RRB- 10_1101-2020_12_31_424971 762 39 and and CC 10_1101-2020_12_31_424971 762 40 Ya Ya NNP 10_1101-2020_12_31_424971 762 41 - - HYPH 10_1101-2020_12_31_424971 762 42 Ming Ming NNP 10_1101-2020_12_31_424971 762 43 Hou Hou NNP 10_1101-2020_12_31_424971 762 44 1208 1208 CD 10_1101-2020_12_31_424971 762 45 ( ( -LRB- 10_1101-2020_12_31_424971 762 46 ya-ming.hou@jefferson.edu ya-ming.hou@jefferson.edu NNP 10_1101-2020_12_31_424971 762 47 ) ) -RRB- 10_1101-2020_12_31_424971 762 48 . . . 10_1101-2020_12_31_424971 763 1 1209 1209 CD 10_1101-2020_12_31_424971 763 2 1210 1210 CD 10_1101-2020_12_31_424971 763 3 1211 1211 CD 10_1101-2020_12_31_424971 763 4 1212 1212 CD 10_1101-2020_12_31_424971 763 5 1213 1213 CD 10_1101-2020_12_31_424971 763 6 1214 1214 CD 10_1101-2020_12_31_424971 763 7 .CC .CC : 10_1101-2020_12_31_424971 763 8 - - : 10_1101-2020_12_31_424971 763 9 BY by IN 10_1101-2020_12_31_424971 763 10 - - HYPH 10_1101-2020_12_31_424971 763 11 NC NC NNP 10_1101-2020_12_31_424971 763 12 - - HYPH 10_1101-2020_12_31_424971 763 13 ND ND NNP 10_1101-2020_12_31_424971 763 14 4.0 4.0 CD 10_1101-2020_12_31_424971 763 15 International International NNP 10_1101-2020_12_31_424971 763 16 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 763 17 under under IN 10_1101-2020_12_31_424971 763 18 a a DT 10_1101-2020_12_31_424971 763 19 ( ( -LRB- 10_1101-2020_12_31_424971 763 20 which which WDT 10_1101-2020_12_31_424971 763 21 was be VBD 10_1101-2020_12_31_424971 763 22 not not RB 10_1101-2020_12_31_424971 763 23 certified certify VBN 10_1101-2020_12_31_424971 763 24 by by IN 10_1101-2020_12_31_424971 763 25 peer peer NN 10_1101-2020_12_31_424971 763 26 review review NN 10_1101-2020_12_31_424971 763 27 ) ) -RRB- 10_1101-2020_12_31_424971 763 28 is be VBZ 10_1101-2020_12_31_424971 763 29 the the DT 10_1101-2020_12_31_424971 763 30 author author NN 10_1101-2020_12_31_424971 763 31 / / SYM 10_1101-2020_12_31_424971 763 32 funder funder NN 10_1101-2020_12_31_424971 763 33 , , , 10_1101-2020_12_31_424971 763 34 who who WP 10_1101-2020_12_31_424971 763 35 has have VBZ 10_1101-2020_12_31_424971 763 36 granted grant VBN 10_1101-2020_12_31_424971 763 37 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 763 38 a a DT 10_1101-2020_12_31_424971 763 39 license license NN 10_1101-2020_12_31_424971 763 40 to to TO 10_1101-2020_12_31_424971 763 41 display display VB 10_1101-2020_12_31_424971 763 42 the the DT 10_1101-2020_12_31_424971 763 43 preprint preprint NN 10_1101-2020_12_31_424971 763 44 in in IN 10_1101-2020_12_31_424971 763 45 perpetuity perpetuity NN 10_1101-2020_12_31_424971 763 46 . . . 10_1101-2020_12_31_424971 764 1 It -PRON- PRP 10_1101-2020_12_31_424971 764 2 is be VBZ 10_1101-2020_12_31_424971 764 3 made make VBN 10_1101-2020_12_31_424971 764 4 The the DT 10_1101-2020_12_31_424971 764 5 copyright copyright NN 10_1101-2020_12_31_424971 764 6 holder holder NN 10_1101-2020_12_31_424971 764 7 for for IN 10_1101-2020_12_31_424971 764 8 this this DT 10_1101-2020_12_31_424971 764 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 764 10 version version NN 10_1101-2020_12_31_424971 764 11 posted post VBD 10_1101-2020_12_31_424971 764 12 January January NNP 10_1101-2020_12_31_424971 764 13 2 2 CD 10_1101-2020_12_31_424971 764 14 , , , 10_1101-2020_12_31_424971 764 15 2021 2021 CD 10_1101-2020_12_31_424971 764 16 . . . 10_1101-2020_12_31_424971 764 17 ; ; : 10_1101-2020_12_31_424971 764 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 764 19 : : : 10_1101-2020_12_31_424971 764 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 764 21 preprint preprint NN 10_1101-2020_12_31_424971 764 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 764 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 764 24 47 47 CD 10_1101-2020_12_31_424971 764 25 FIGURE FIGURE NNP 10_1101-2020_12_31_424971 764 26 LEGENDS legend NNS 10_1101-2020_12_31_424971 764 27 1215 1215 CD 10_1101-2020_12_31_424971 764 28 1216 1216 CD 10_1101-2020_12_31_424971 764 29 Figure Figure NNP 10_1101-2020_12_31_424971 764 30 1 1 CD 10_1101-2020_12_31_424971 764 31 . . . 10_1101-2020_12_31_424971 765 1 Methylation methylation NN 10_1101-2020_12_31_424971 765 2 and and CC 10_1101-2020_12_31_424971 765 3 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 765 4 of of IN 10_1101-2020_12_31_424971 765 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 765 6 and and CC 10_1101-2020_12_31_424971 765 7 ProL. ProL. . 10_1101-2020_12_31_424971 766 1 a a DT 10_1101-2020_12_31_424971 766 2 Sequence Sequence NNP 10_1101-2020_12_31_424971 766 3 and and CC 10_1101-2020_12_31_424971 766 4 secondary secondary JJ 10_1101-2020_12_31_424971 766 5 1217 1217 CD 10_1101-2020_12_31_424971 766 6 structure structure NN 10_1101-2020_12_31_424971 766 7 of of IN 10_1101-2020_12_31_424971 766 8 native native JJ 10_1101-2020_12_31_424971 766 9 - - HYPH 10_1101-2020_12_31_424971 766 10 state state NN 10_1101-2020_12_31_424971 766 11 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 766 12 , , , 10_1101-2020_12_31_424971 766 13 showing show VBG 10_1101-2020_12_31_424971 766 14 the the DT 10_1101-2020_12_31_424971 766 15 N1-methylated n1-methylate VBN 10_1101-2020_12_31_424971 766 16 G37 G37 NNP 10_1101-2020_12_31_424971 766 17 in in IN 10_1101-2020_12_31_424971 766 18 red red NN 10_1101-2020_12_31_424971 766 19 and and CC 10_1101-2020_12_31_424971 766 20 the the DT 10_1101-2020_12_31_424971 766 21 G37a g37a NN 10_1101-2020_12_31_424971 766 22 insertion insertion NN 10_1101-2020_12_31_424971 766 23 to to IN 10_1101-2020_12_31_424971 766 24 1218 1218 CD 10_1101-2020_12_31_424971 766 25 ProL ProL NNP 10_1101-2020_12_31_424971 766 26 in in IN 10_1101-2020_12_31_424971 766 27 blue blue NNP 10_1101-2020_12_31_424971 766 28 . . . 10_1101-2020_12_31_424971 767 1 b b LS 10_1101-2020_12_31_424971 767 2 RNase RNase NNP 10_1101-2020_12_31_424971 767 3 T1 T1 NNP 10_1101-2020_12_31_424971 767 4 cleavage cleavage NN 10_1101-2020_12_31_424971 767 5 inhibition inhibition NN 10_1101-2020_12_31_424971 767 6 assays assays RB 10_1101-2020_12_31_424971 767 7 of of IN 10_1101-2020_12_31_424971 767 8 TrmD TrmD NNP 10_1101-2020_12_31_424971 767 9 - - HYPH 10_1101-2020_12_31_424971 767 10 methylated methylate VBN 10_1101-2020_12_31_424971 767 11 G37-state G37-state NNP 10_1101-2020_12_31_424971 767 12 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 767 13 1219 1219 CD 10_1101-2020_12_31_424971 767 14 transcript transcript NN 10_1101-2020_12_31_424971 767 15 confirm confirm VB 10_1101-2020_12_31_424971 767 16 the the DT 10_1101-2020_12_31_424971 767 17 presence presence NN 10_1101-2020_12_31_424971 767 18 of of IN 10_1101-2020_12_31_424971 767 19 m1G37 m1G37 NNP 10_1101-2020_12_31_424971 767 20 and and CC 10_1101-2020_12_31_424971 767 21 m1G37a m1g37a ADD 10_1101-2020_12_31_424971 767 22 . . . 10_1101-2020_12_31_424971 768 1 Cleavage cleavage NN 10_1101-2020_12_31_424971 768 2 products product NNS 10_1101-2020_12_31_424971 768 3 are be VBP 10_1101-2020_12_31_424971 768 4 marked mark VBN 10_1101-2020_12_31_424971 768 5 by by IN 10_1101-2020_12_31_424971 768 6 the the DT 10_1101-2020_12_31_424971 768 7 1220 1220 CD 10_1101-2020_12_31_424971 768 8 nucleotide nucleotide JJ 10_1101-2020_12_31_424971 768 9 positions position NNS 10_1101-2020_12_31_424971 768 10 of of IN 10_1101-2020_12_31_424971 768 11 Gs Gs NNP 10_1101-2020_12_31_424971 768 12 . . . 10_1101-2020_12_31_424971 769 1 L l NN 10_1101-2020_12_31_424971 769 2 : : : 10_1101-2020_12_31_424971 769 3 the the DT 10_1101-2020_12_31_424971 769 4 molecular molecular JJ 10_1101-2020_12_31_424971 769 5 ladder ladder NN 10_1101-2020_12_31_424971 769 6 of of IN 10_1101-2020_12_31_424971 769 7 tRNA trna NN 10_1101-2020_12_31_424971 769 8 fragments fragment NNS 10_1101-2020_12_31_424971 769 9 generated generate VBN 10_1101-2020_12_31_424971 769 10 from from IN 10_1101-2020_12_31_424971 769 11 alkali alkali JJ 10_1101-2020_12_31_424971 769 12 1221 1221 CD 10_1101-2020_12_31_424971 769 13 hydrolysis hydrolysis NN 10_1101-2020_12_31_424971 769 14 . . . 10_1101-2020_12_31_424971 770 1 c c LS 10_1101-2020_12_31_424971 770 2 Primer Primer NNP 10_1101-2020_12_31_424971 770 3 extension extension NN 10_1101-2020_12_31_424971 770 4 inhibition inhibition NN 10_1101-2020_12_31_424971 770 5 assays assays RB 10_1101-2020_12_31_424971 770 6 identify identify VBP 10_1101-2020_12_31_424971 770 7 m1G37 m1G37 NNP 10_1101-2020_12_31_424971 770 8 in in IN 10_1101-2020_12_31_424971 770 9 native native JJ 10_1101-2020_12_31_424971 770 10 - - HYPH 10_1101-2020_12_31_424971 770 11 state state NN 10_1101-2020_12_31_424971 770 12 SufB2 sufb2 NN 10_1101-2020_12_31_424971 770 13 . . . 10_1101-2020_12_31_424971 771 1 Red red JJ 10_1101-2020_12_31_424971 771 2 and and CC 10_1101-2020_12_31_424971 771 3 1222 1222 CD 10_1101-2020_12_31_424971 771 4 blue blue JJ 10_1101-2020_12_31_424971 771 5 arrows arrow NNS 10_1101-2020_12_31_424971 771 6 indicate indicate VBP 10_1101-2020_12_31_424971 771 7 positions position NNS 10_1101-2020_12_31_424971 771 8 of of IN 10_1101-2020_12_31_424971 771 9 primer primer JJ 10_1101-2020_12_31_424971 771 10 extension extension NN 10_1101-2020_12_31_424971 771 11 inhibition inhibition NN 10_1101-2020_12_31_424971 771 12 products product NNS 10_1101-2020_12_31_424971 771 13 at at IN 10_1101-2020_12_31_424971 771 14 the the DT 10_1101-2020_12_31_424971 771 15 methylated methylated JJ 10_1101-2020_12_31_424971 771 16 G37 G37 NNP 10_1101-2020_12_31_424971 771 17 and and CC 10_1101-2020_12_31_424971 771 18 1223 1223 CD 10_1101-2020_12_31_424971 771 19 G37a G37a NNS 10_1101-2020_12_31_424971 771 20 , , , 10_1101-2020_12_31_424971 771 21 respectively respectively RB 10_1101-2020_12_31_424971 771 22 , , , 10_1101-2020_12_31_424971 771 23 which which WDT 10_1101-2020_12_31_424971 771 24 are be VBP 10_1101-2020_12_31_424971 771 25 offset offset VBN 10_1101-2020_12_31_424971 771 26 by by IN 10_1101-2020_12_31_424971 771 27 one one CD 10_1101-2020_12_31_424971 771 28 nucleotide nucleotide JJ 10_1101-2020_12_31_424971 771 29 relative relative NNP 10_1101-2020_12_31_424971 771 30 to to IN 10_1101-2020_12_31_424971 771 31 ProL. ProL. . 10_1101-2020_12_31_424971 772 1 The the DT 10_1101-2020_12_31_424971 772 2 first first JJ 10_1101-2020_12_31_424971 772 3 primer primer JJ 10_1101-2020_12_31_424971 772 4 extension extension NN 10_1101-2020_12_31_424971 772 5 1224 1224 CD 10_1101-2020_12_31_424971 772 6 inhibition inhibition NN 10_1101-2020_12_31_424971 772 7 product product NN 10_1101-2020_12_31_424971 772 8 for for IN 10_1101-2020_12_31_424971 772 9 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 772 10 corresponds correspond NNS 10_1101-2020_12_31_424971 772 11 to to IN 10_1101-2020_12_31_424971 772 12 m1G37a m1g37a ADD 10_1101-2020_12_31_424971 772 13 , , , 10_1101-2020_12_31_424971 772 14 the the DT 10_1101-2020_12_31_424971 772 15 second second JJ 10_1101-2020_12_31_424971 772 16 corresponds correspond NNS 10_1101-2020_12_31_424971 772 17 to to TO 10_1101-2020_12_31_424971 772 18 m1G37 m1g37 VB 10_1101-2020_12_31_424971 772 19 , , , 10_1101-2020_12_31_424971 772 20 while while IN 10_1101-2020_12_31_424971 772 21 the the DT 10_1101-2020_12_31_424971 772 22 1225 1225 CD 10_1101-2020_12_31_424971 772 23 primer primer NNP 10_1101-2020_12_31_424971 772 24 extension extension NN 10_1101-2020_12_31_424971 772 25 inhibition inhibition NN 10_1101-2020_12_31_424971 772 26 product product NN 10_1101-2020_12_31_424971 772 27 for for IN 10_1101-2020_12_31_424971 772 28 ProL ProL NNP 10_1101-2020_12_31_424971 772 29 corresponds correspond NNS 10_1101-2020_12_31_424971 772 30 to to IN 10_1101-2020_12_31_424971 772 31 m1G37 m1g37 VB 10_1101-2020_12_31_424971 772 32 . . . 10_1101-2020_12_31_424971 773 1 Due due IN 10_1101-2020_12_31_424971 773 2 to to IN 10_1101-2020_12_31_424971 773 3 the the DT 10_1101-2020_12_31_424971 773 4 propensity propensity NN 10_1101-2020_12_31_424971 773 5 of of IN 10_1101-2020_12_31_424971 773 6 1226 1226 CD 10_1101-2020_12_31_424971 773 7 primer primer NN 10_1101-2020_12_31_424971 773 8 extension extension NN 10_1101-2020_12_31_424971 773 9 to to TO 10_1101-2020_12_31_424971 773 10 make make VB 10_1101-2020_12_31_424971 773 11 multiple multiple JJ 10_1101-2020_12_31_424971 773 12 stops stop NNS 10_1101-2020_12_31_424971 773 13 on on IN 10_1101-2020_12_31_424971 773 14 a a DT 10_1101-2020_12_31_424971 773 15 long long JJ 10_1101-2020_12_31_424971 773 16 transcript transcript NN 10_1101-2020_12_31_424971 773 17 of of IN 10_1101-2020_12_31_424971 773 18 tRNA tRNA NNP 10_1101-2020_12_31_424971 773 19 , , , 10_1101-2020_12_31_424971 773 20 the the DT 10_1101-2020_12_31_424971 773 21 read read VB 10_1101-2020_12_31_424971 773 22 - - HYPH 10_1101-2020_12_31_424971 773 23 through through RP 10_1101-2020_12_31_424971 773 24 primer primer NN 10_1101-2020_12_31_424971 773 25 1227 1227 CD 10_1101-2020_12_31_424971 773 26 extension extension NN 10_1101-2020_12_31_424971 773 27 product product NN 10_1101-2020_12_31_424971 773 28 ( ( -LRB- 10_1101-2020_12_31_424971 773 29 54 54 CD 10_1101-2020_12_31_424971 773 30 - - SYM 10_1101-2020_12_31_424971 773 31 55 55 CD 10_1101-2020_12_31_424971 773 32 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 773 33 ) ) -RRB- 10_1101-2020_12_31_424971 773 34 had have VBD 10_1101-2020_12_31_424971 773 35 a a DT 10_1101-2020_12_31_424971 773 36 reduced reduce VBN 10_1101-2020_12_31_424971 773 37 intensity intensity NN 10_1101-2020_12_31_424971 773 38 relative relative JJ 10_1101-2020_12_31_424971 773 39 to to IN 10_1101-2020_12_31_424971 773 40 the the DT 10_1101-2020_12_31_424971 773 41 primer primer NNP 10_1101-2020_12_31_424971 773 42 extension extension NN 10_1101-2020_12_31_424971 773 43 1228 1228 CD 10_1101-2020_12_31_424971 773 44 inhibition inhibition NN 10_1101-2020_12_31_424971 773 45 products product NNS 10_1101-2020_12_31_424971 773 46 ( ( -LRB- 10_1101-2020_12_31_424971 773 47 21 21 CD 10_1101-2020_12_31_424971 773 48 - - SYM 10_1101-2020_12_31_424971 773 49 22 22 CD 10_1101-2020_12_31_424971 773 50 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 773 51 ) ) -RRB- 10_1101-2020_12_31_424971 773 52 . . . 10_1101-2020_12_31_424971 774 1 Molecular molecular JJ 10_1101-2020_12_31_424971 774 2 size size NN 10_1101-2020_12_31_424971 774 3 markers marker NNS 10_1101-2020_12_31_424971 774 4 are be VBP 10_1101-2020_12_31_424971 774 5 provided provide VBN 10_1101-2020_12_31_424971 774 6 by by IN 10_1101-2020_12_31_424971 774 7 the the DT 10_1101-2020_12_31_424971 774 8 primer primer NN 10_1101-2020_12_31_424971 774 9 alone alone RB 10_1101-2020_12_31_424971 774 10 1229 1229 CD 10_1101-2020_12_31_424971 774 11 ( ( -LRB- 10_1101-2020_12_31_424971 774 12 17 17 CD 10_1101-2020_12_31_424971 774 13 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 774 14 ) ) -RRB- 10_1101-2020_12_31_424971 774 15 and and CC 10_1101-2020_12_31_424971 774 16 the the DT 10_1101-2020_12_31_424971 774 17 run run VB 10_1101-2020_12_31_424971 774 18 - - HYPH 10_1101-2020_12_31_424971 774 19 off off RP 10_1101-2020_12_31_424971 774 20 products product NNS 10_1101-2020_12_31_424971 774 21 ( ( -LRB- 10_1101-2020_12_31_424971 774 22 54 54 CD 10_1101-2020_12_31_424971 774 23 - - SYM 10_1101-2020_12_31_424971 774 24 55 55 CD 10_1101-2020_12_31_424971 774 25 nucleotides nucleotide NNS 10_1101-2020_12_31_424971 774 26 ) ) -RRB- 10_1101-2020_12_31_424971 774 27 . . . 10_1101-2020_12_31_424971 775 1 d d LS 10_1101-2020_12_31_424971 775 2 TrmD TrmD NNP 10_1101-2020_12_31_424971 775 3 - - HYPH 10_1101-2020_12_31_424971 775 4 catalyzed catalyze VBN 10_1101-2020_12_31_424971 775 5 N1 n1 NN 10_1101-2020_12_31_424971 775 6 methylation methylation NN 10_1101-2020_12_31_424971 775 7 1230 1230 CD 10_1101-2020_12_31_424971 775 8 of of IN 10_1101-2020_12_31_424971 775 9 G37-state G37-state NNP 10_1101-2020_12_31_424971 775 10 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 775 11 and and CC 10_1101-2020_12_31_424971 775 12 ProL ProL NNP 10_1101-2020_12_31_424971 775 13 as as IN 10_1101-2020_12_31_424971 775 14 a a DT 10_1101-2020_12_31_424971 775 15 function function NN 10_1101-2020_12_31_424971 775 16 of of IN 10_1101-2020_12_31_424971 775 17 time time NN 10_1101-2020_12_31_424971 775 18 . . . 10_1101-2020_12_31_424971 776 1 e e NNP 10_1101-2020_12_31_424971 776 2 , , , 10_1101-2020_12_31_424971 776 3 f f NNP 10_1101-2020_12_31_424971 776 4 ProRS ProRS NNP 10_1101-2020_12_31_424971 776 5 - - HYPH 10_1101-2020_12_31_424971 776 6 catalyzed catalyzed JJ 10_1101-2020_12_31_424971 776 7 aminoacylation aminoacylation NN 10_1101-2020_12_31_424971 776 8 . . . 10_1101-2020_12_31_424971 777 1 e e LS 10_1101-2020_12_31_424971 777 2 1231 1231 CD 10_1101-2020_12_31_424971 777 3 Aminoacylation Aminoacylation NNP 10_1101-2020_12_31_424971 777 4 of of IN 10_1101-2020_12_31_424971 777 5 native native JJ 10_1101-2020_12_31_424971 777 6 - - HYPH 10_1101-2020_12_31_424971 777 7 state state NN 10_1101-2020_12_31_424971 777 8 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 777 9 and and CC 10_1101-2020_12_31_424971 777 10 ProL. ProL. . 10_1101-2020_12_31_424971 778 1 f f LS 10_1101-2020_12_31_424971 778 2 Aminoacylation Aminoacylation NNP 10_1101-2020_12_31_424971 778 3 of of IN 10_1101-2020_12_31_424971 778 4 G37-state G37-state NNP 10_1101-2020_12_31_424971 778 5 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 778 6 and and CC 10_1101-2020_12_31_424971 778 7 ProL ProL NNP 10_1101-2020_12_31_424971 778 8 1232 1232 CD 10_1101-2020_12_31_424971 778 9 as as IN 10_1101-2020_12_31_424971 778 10 a a DT 10_1101-2020_12_31_424971 778 11 function function NN 10_1101-2020_12_31_424971 778 12 of of IN 10_1101-2020_12_31_424971 778 13 time time NN 10_1101-2020_12_31_424971 778 14 . . . 10_1101-2020_12_31_424971 779 1 In in IN 10_1101-2020_12_31_424971 779 2 panels panels NNP 10_1101-2020_12_31_424971 779 3 b b NNP 10_1101-2020_12_31_424971 779 4 , , , 10_1101-2020_12_31_424971 779 5 c c NNP 10_1101-2020_12_31_424971 779 6 , , , 10_1101-2020_12_31_424971 779 7 gels gel NNS 10_1101-2020_12_31_424971 779 8 were be VBD 10_1101-2020_12_31_424971 779 9 performed perform VBN 10_1101-2020_12_31_424971 779 10 three three CD 10_1101-2020_12_31_424971 779 11 times time NNS 10_1101-2020_12_31_424971 779 12 with with IN 10_1101-2020_12_31_424971 779 13 similar similar JJ 10_1101-2020_12_31_424971 779 14 results result NNS 10_1101-2020_12_31_424971 779 15 , , , 10_1101-2020_12_31_424971 779 16 while while IN 10_1101-2020_12_31_424971 779 17 in in IN 10_1101-2020_12_31_424971 779 18 1233 1233 CD 10_1101-2020_12_31_424971 779 19 panels panel NNS 10_1101-2020_12_31_424971 779 20 d d NNP 10_1101-2020_12_31_424971 779 21 - - HYPH 10_1101-2020_12_31_424971 779 22 f f NNP 10_1101-2020_12_31_424971 779 23 , , , 10_1101-2020_12_31_424971 779 24 the the DT 10_1101-2020_12_31_424971 779 25 bars bar NNS 10_1101-2020_12_31_424971 779 26 are be VBP 10_1101-2020_12_31_424971 779 27 SD sd NN 10_1101-2020_12_31_424971 779 28 of of IN 10_1101-2020_12_31_424971 779 29 three three CD 10_1101-2020_12_31_424971 779 30 independent independent NN 10_1101-2020_12_31_424971 779 31 ( ( -LRB- 10_1101-2020_12_31_424971 779 32 n n NN 10_1101-2020_12_31_424971 779 33 = = SYM 10_1101-2020_12_31_424971 779 34 3 3 LS 10_1101-2020_12_31_424971 779 35 ) ) -RRB- 10_1101-2020_12_31_424971 779 36 experiments experiment NNS 10_1101-2020_12_31_424971 779 37 , , , 10_1101-2020_12_31_424971 779 38 and and CC 10_1101-2020_12_31_424971 779 39 the the DT 10_1101-2020_12_31_424971 779 40 data datum NNS 10_1101-2020_12_31_424971 779 41 are be VBP 10_1101-2020_12_31_424971 779 42 presented present VBN 10_1101-2020_12_31_424971 779 43 1234 1234 CD 10_1101-2020_12_31_424971 779 44 as as IN 10_1101-2020_12_31_424971 779 45 mean mean JJ 10_1101-2020_12_31_424971 779 46 values value NNS 10_1101-2020_12_31_424971 779 47 ± ± CD 10_1101-2020_12_31_424971 779 48 SD sd NN 10_1101-2020_12_31_424971 779 49 . . . 10_1101-2020_12_31_424971 780 1 1235 1235 CD 10_1101-2020_12_31_424971 780 2 1236 1236 CD 10_1101-2020_12_31_424971 780 3 Figure Figure NNP 10_1101-2020_12_31_424971 780 4 2 2 CD 10_1101-2020_12_31_424971 780 5 . . . 10_1101-2020_12_31_424971 781 1 SufB2-induced sufb2-induced XX 10_1101-2020_12_31_424971 781 2 +1 +1 NNS 10_1101-2020_12_31_424971 781 3 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 781 4 and and CC 10_1101-2020_12_31_424971 781 5 genome genome JJ 10_1101-2020_12_31_424971 781 6 recoding recoding NN 10_1101-2020_12_31_424971 781 7 . . . 10_1101-2020_12_31_424971 782 1 a a DT 10_1101-2020_12_31_424971 782 2 The The NNP 10_1101-2020_12_31_424971 782 3 +1-frameshifting +1-frameshifting NNP 10_1101-2020_12_31_424971 782 4 1237 1237 CD 10_1101-2020_12_31_424971 782 5 efficiency efficiency NN 10_1101-2020_12_31_424971 782 6 in in IN 10_1101-2020_12_31_424971 782 7 cell cell NN 10_1101-2020_12_31_424971 782 8 - - HYPH 10_1101-2020_12_31_424971 782 9 based base VBN 10_1101-2020_12_31_424971 782 10 lacZ lacz JJ 10_1101-2020_12_31_424971 782 11 assay assay NNP 10_1101-2020_12_31_424971 782 12 for for IN 10_1101-2020_12_31_424971 782 13 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 782 14 and and CC 10_1101-2020_12_31_424971 782 15 ProL ProL NNP 10_1101-2020_12_31_424971 782 16 strains strain VBZ 10_1101-2020_12_31_424971 782 17 in in IN 10_1101-2020_12_31_424971 782 18 m1G37 m1g37 JJ 10_1101-2020_12_31_424971 782 19 + + SYM 10_1101-2020_12_31_424971 782 20 and and CC 10_1101-2020_12_31_424971 782 21 m1G37 m1g37 NN 10_1101-2020_12_31_424971 782 22 – – : 10_1101-2020_12_31_424971 782 23 conditions condition NNS 10_1101-2020_12_31_424971 782 24 . . . 10_1101-2020_12_31_424971 783 1 1238 1238 CD 10_1101-2020_12_31_424971 783 2 The the DT 10_1101-2020_12_31_424971 783 3 bars bar NNS 10_1101-2020_12_31_424971 783 4 in in IN 10_1101-2020_12_31_424971 783 5 the the DT 10_1101-2020_12_31_424971 783 6 graph graph NN 10_1101-2020_12_31_424971 783 7 are be VBP 10_1101-2020_12_31_424971 783 8 SD sd NN 10_1101-2020_12_31_424971 783 9 of of IN 10_1101-2020_12_31_424971 783 10 four four CD 10_1101-2020_12_31_424971 783 11 , , , 10_1101-2020_12_31_424971 783 12 five five CD 10_1101-2020_12_31_424971 783 13 , , , 10_1101-2020_12_31_424971 783 14 or or CC 10_1101-2020_12_31_424971 783 15 six six CD 10_1101-2020_12_31_424971 783 16 independent independent JJ 10_1101-2020_12_31_424971 783 17 ( ( -LRB- 10_1101-2020_12_31_424971 783 18 n n NN 10_1101-2020_12_31_424971 783 19 = = SYM 10_1101-2020_12_31_424971 783 20 4 4 CD 10_1101-2020_12_31_424971 783 21 , , , 10_1101-2020_12_31_424971 783 22 5 5 CD 10_1101-2020_12_31_424971 783 23 , , , 10_1101-2020_12_31_424971 783 24 or or CC 10_1101-2020_12_31_424971 783 25 6 6 CD 10_1101-2020_12_31_424971 783 26 ) ) -RRB- 10_1101-2020_12_31_424971 783 27 biological biological JJ 10_1101-2020_12_31_424971 783 28 repeats repeat NNS 10_1101-2020_12_31_424971 783 29 , , , 10_1101-2020_12_31_424971 783 30 1239 1239 CD 10_1101-2020_12_31_424971 783 31 and and CC 10_1101-2020_12_31_424971 783 32 the the DT 10_1101-2020_12_31_424971 783 33 data datum NNS 10_1101-2020_12_31_424971 783 34 are be VBP 10_1101-2020_12_31_424971 783 35 mean mean JJ 10_1101-2020_12_31_424971 783 36 values value NNS 10_1101-2020_12_31_424971 783 37 ± ± CD 10_1101-2020_12_31_424971 783 38 SD sd NN 10_1101-2020_12_31_424971 783 39 . . . 10_1101-2020_12_31_424971 784 1 b b LS 10_1101-2020_12_31_424971 784 2 The the DT 10_1101-2020_12_31_424971 784 3 difference difference NN 10_1101-2020_12_31_424971 784 4 in in IN 10_1101-2020_12_31_424971 784 5 the the DT 10_1101-2020_12_31_424971 784 6 ratio ratio NN 10_1101-2020_12_31_424971 784 7 of of IN 10_1101-2020_12_31_424971 784 8 protein protein NN 10_1101-2020_12_31_424971 784 9 synthesis synthesis NN 10_1101-2020_12_31_424971 784 10 of of IN 10_1101-2020_12_31_424971 784 11 lolB lolB NNP 10_1101-2020_12_31_424971 784 12 to to IN 10_1101-2020_12_31_424971 784 13 1240 1240 CD 10_1101-2020_12_31_424971 784 14 cysS cys NNS 10_1101-2020_12_31_424971 784 15 for for IN 10_1101-2020_12_31_424971 784 16 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 784 17 and and CC 10_1101-2020_12_31_424971 784 18 ProL ProL NNP 10_1101-2020_12_31_424971 784 19 strains strain VBZ 10_1101-2020_12_31_424971 784 20 in in IN 10_1101-2020_12_31_424971 784 21 m1G37 m1g37 JJ 10_1101-2020_12_31_424971 784 22 + + SYM 10_1101-2020_12_31_424971 784 23 and and CC 10_1101-2020_12_31_424971 784 24 m1G37 m1g37 NN 10_1101-2020_12_31_424971 784 25 – – : 10_1101-2020_12_31_424971 784 26 conditions condition NNS 10_1101-2020_12_31_424971 784 27 relative relative JJ 10_1101-2020_12_31_424971 784 28 to to IN 10_1101-2020_12_31_424971 784 29 ProL ProL NNP 10_1101-2020_12_31_424971 784 30 in in IN 10_1101-2020_12_31_424971 784 31 the the DT 10_1101-2020_12_31_424971 784 32 m1G37 m1g37 NN 10_1101-2020_12_31_424971 784 33 + + CD 10_1101-2020_12_31_424971 784 34 1241 1241 CD 10_1101-2020_12_31_424971 784 35 .CC .cc SYM 10_1101-2020_12_31_424971 784 36 - - HYPH 10_1101-2020_12_31_424971 784 37 BY by IN 10_1101-2020_12_31_424971 784 38 - - HYPH 10_1101-2020_12_31_424971 784 39 NC NC NNP 10_1101-2020_12_31_424971 784 40 - - HYPH 10_1101-2020_12_31_424971 784 41 ND ND NNP 10_1101-2020_12_31_424971 784 42 4.0 4.0 CD 10_1101-2020_12_31_424971 784 43 International International NNP 10_1101-2020_12_31_424971 784 44 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 784 45 under under IN 10_1101-2020_12_31_424971 784 46 a a DT 10_1101-2020_12_31_424971 784 47 ( ( -LRB- 10_1101-2020_12_31_424971 784 48 which which WDT 10_1101-2020_12_31_424971 784 49 was be VBD 10_1101-2020_12_31_424971 784 50 not not RB 10_1101-2020_12_31_424971 784 51 certified certify VBN 10_1101-2020_12_31_424971 784 52 by by IN 10_1101-2020_12_31_424971 784 53 peer peer NN 10_1101-2020_12_31_424971 784 54 review review NN 10_1101-2020_12_31_424971 784 55 ) ) -RRB- 10_1101-2020_12_31_424971 784 56 is be VBZ 10_1101-2020_12_31_424971 784 57 the the DT 10_1101-2020_12_31_424971 784 58 author author NN 10_1101-2020_12_31_424971 784 59 / / SYM 10_1101-2020_12_31_424971 784 60 funder funder NN 10_1101-2020_12_31_424971 784 61 , , , 10_1101-2020_12_31_424971 784 62 who who WP 10_1101-2020_12_31_424971 784 63 has have VBZ 10_1101-2020_12_31_424971 784 64 granted grant VBN 10_1101-2020_12_31_424971 784 65 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 784 66 a a DT 10_1101-2020_12_31_424971 784 67 license license NN 10_1101-2020_12_31_424971 784 68 to to TO 10_1101-2020_12_31_424971 784 69 display display VB 10_1101-2020_12_31_424971 784 70 the the DT 10_1101-2020_12_31_424971 784 71 preprint preprint NN 10_1101-2020_12_31_424971 784 72 in in IN 10_1101-2020_12_31_424971 784 73 perpetuity perpetuity NN 10_1101-2020_12_31_424971 784 74 . . . 10_1101-2020_12_31_424971 785 1 It -PRON- PRP 10_1101-2020_12_31_424971 785 2 is be VBZ 10_1101-2020_12_31_424971 785 3 made make VBN 10_1101-2020_12_31_424971 785 4 The the DT 10_1101-2020_12_31_424971 785 5 copyright copyright NN 10_1101-2020_12_31_424971 785 6 holder holder NN 10_1101-2020_12_31_424971 785 7 for for IN 10_1101-2020_12_31_424971 785 8 this this DT 10_1101-2020_12_31_424971 785 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 785 10 version version NN 10_1101-2020_12_31_424971 785 11 posted post VBD 10_1101-2020_12_31_424971 785 12 January January NNP 10_1101-2020_12_31_424971 785 13 2 2 CD 10_1101-2020_12_31_424971 785 14 , , , 10_1101-2020_12_31_424971 785 15 2021 2021 CD 10_1101-2020_12_31_424971 785 16 . . . 10_1101-2020_12_31_424971 785 17 ; ; : 10_1101-2020_12_31_424971 785 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 785 19 : : : 10_1101-2020_12_31_424971 785 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 785 21 preprint preprint NN 10_1101-2020_12_31_424971 785 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 785 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 785 24 48 48 CD 10_1101-2020_12_31_424971 785 25 condition condition NN 10_1101-2020_12_31_424971 785 26 . . . 10_1101-2020_12_31_424971 786 1 c c NNP 10_1101-2020_12_31_424971 786 2 Measurements measurement NNS 10_1101-2020_12_31_424971 786 3 underlying underlie VBG 10_1101-2020_12_31_424971 786 4 the the DT 10_1101-2020_12_31_424971 786 5 bar bar NN 10_1101-2020_12_31_424971 786 6 plots plot NNS 10_1101-2020_12_31_424971 786 7 in in IN 10_1101-2020_12_31_424971 786 8 panel panel NN 10_1101-2020_12_31_424971 786 9 b. b. NNP 10_1101-2020_12_31_424971 787 1 Each each DT 10_1101-2020_12_31_424971 787 2 ratio ratio NN 10_1101-2020_12_31_424971 787 3 was be VBD 10_1101-2020_12_31_424971 787 4 measured measure VBN 10_1101-2020_12_31_424971 787 5 directly directly RB 10_1101-2020_12_31_424971 787 6 1242 1242 CD 10_1101-2020_12_31_424971 787 7 and and CC 10_1101-2020_12_31_424971 787 8 the the DT 10_1101-2020_12_31_424971 787 9 ratio ratio NN 10_1101-2020_12_31_424971 787 10 of of IN 10_1101-2020_12_31_424971 787 11 ProL ProL NNP 10_1101-2020_12_31_424971 787 12 in in IN 10_1101-2020_12_31_424971 787 13 the the DT 10_1101-2020_12_31_424971 787 14 m1G37 m1g37 NN 10_1101-2020_12_31_424971 787 15 + + CC 10_1101-2020_12_31_424971 787 16 condition condition NN 10_1101-2020_12_31_424971 787 17 was be VBD 10_1101-2020_12_31_424971 787 18 normalized normalize VBN 10_1101-2020_12_31_424971 787 19 to to IN 10_1101-2020_12_31_424971 787 20 1.0 1.0 CD 10_1101-2020_12_31_424971 787 21 . . . 10_1101-2020_12_31_424971 788 1 The the DT 10_1101-2020_12_31_424971 788 2 difference difference NN 10_1101-2020_12_31_424971 788 3 of of IN 10_1101-2020_12_31_424971 788 4 each each DT 10_1101-2020_12_31_424971 788 5 ratio ratio NN 10_1101-2020_12_31_424971 788 6 1243 1243 CD 10_1101-2020_12_31_424971 788 7 relative relative JJ 10_1101-2020_12_31_424971 788 8 to to IN 10_1101-2020_12_31_424971 788 9 the the DT 10_1101-2020_12_31_424971 788 10 normalized normalized JJ 10_1101-2020_12_31_424971 788 11 ratio ratio NN 10_1101-2020_12_31_424971 788 12 represented represent VBD 10_1101-2020_12_31_424971 788 13 the the DT 10_1101-2020_12_31_424971 788 14 +1-frameshifting +1-frameshifte VBG 10_1101-2020_12_31_424971 788 15 efficiency efficiency NN 10_1101-2020_12_31_424971 788 16 at at IN 10_1101-2020_12_31_424971 788 17 the the DT 10_1101-2020_12_31_424971 788 18 CCC CCC NNP 10_1101-2020_12_31_424971 788 19 - - HYPH 10_1101-2020_12_31_424971 788 20 C C NNP 10_1101-2020_12_31_424971 788 21 motif motif NN 10_1101-2020_12_31_424971 788 22 at at IN 10_1101-2020_12_31_424971 788 23 1244 1244 CD 10_1101-2020_12_31_424971 788 24 the the DT 10_1101-2020_12_31_424971 788 25 2nd 2nd JJ 10_1101-2020_12_31_424971 788 26 codon codon NN 10_1101-2020_12_31_424971 788 27 of of IN 10_1101-2020_12_31_424971 788 28 lolB. lolb. NN 10_1101-2020_12_31_424971 789 1 The the DT 10_1101-2020_12_31_424971 789 2 bars bar NNS 10_1101-2020_12_31_424971 789 3 in in IN 10_1101-2020_12_31_424971 789 4 the the DT 10_1101-2020_12_31_424971 789 5 graph graph NN 10_1101-2020_12_31_424971 789 6 are be VBP 10_1101-2020_12_31_424971 789 7 SD sd NN 10_1101-2020_12_31_424971 789 8 of of IN 10_1101-2020_12_31_424971 789 9 three three CD 10_1101-2020_12_31_424971 789 10 independent independent NN 10_1101-2020_12_31_424971 789 11 ( ( -LRB- 10_1101-2020_12_31_424971 789 12 n n NN 10_1101-2020_12_31_424971 789 13 = = SYM 10_1101-2020_12_31_424971 789 14 3 3 LS 10_1101-2020_12_31_424971 789 15 ) ) -RRB- 10_1101-2020_12_31_424971 789 16 biological biological JJ 10_1101-2020_12_31_424971 789 17 repeats repeat NNS 10_1101-2020_12_31_424971 789 18 , , , 10_1101-2020_12_31_424971 789 19 1245 1245 CD 10_1101-2020_12_31_424971 789 20 and and CC 10_1101-2020_12_31_424971 789 21 the the DT 10_1101-2020_12_31_424971 789 22 data datum NNS 10_1101-2020_12_31_424971 789 23 are be VBP 10_1101-2020_12_31_424971 789 24 mean mean JJ 10_1101-2020_12_31_424971 789 25 values value NNS 10_1101-2020_12_31_424971 789 26 ± ± CD 10_1101-2020_12_31_424971 789 27 SD sd NN 10_1101-2020_12_31_424971 789 28 . . . 10_1101-2020_12_31_424971 790 1 In in IN 10_1101-2020_12_31_424971 790 2 a a DT 10_1101-2020_12_31_424971 790 3 , , , 10_1101-2020_12_31_424971 790 4 b b NN 10_1101-2020_12_31_424971 790 5 , , , 10_1101-2020_12_31_424971 790 6 decoding decoding NN 10_1101-2020_12_31_424971 790 7 of of IN 10_1101-2020_12_31_424971 790 8 the the DT 10_1101-2020_12_31_424971 790 9 CCC CCC NNP 10_1101-2020_12_31_424971 790 10 - - HYPH 10_1101-2020_12_31_424971 790 11 C C NNP 10_1101-2020_12_31_424971 790 12 motif motif NN 10_1101-2020_12_31_424971 790 13 was be VBD 10_1101-2020_12_31_424971 790 14 mediated mediate VBN 10_1101-2020_12_31_424971 790 15 by by IN 10_1101-2020_12_31_424971 790 16 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 790 17 1246 1246 CD 10_1101-2020_12_31_424971 790 18 and and CC 10_1101-2020_12_31_424971 790 19 ProM ProM NNP 10_1101-2020_12_31_424971 790 20 in in IN 10_1101-2020_12_31_424971 790 21 the the DT 10_1101-2020_12_31_424971 790 22 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 790 23 strain strain NN 10_1101-2020_12_31_424971 790 24 , , , 10_1101-2020_12_31_424971 790 25 and and CC 10_1101-2020_12_31_424971 790 26 by by IN 10_1101-2020_12_31_424971 790 27 ProL ProL NNP 10_1101-2020_12_31_424971 790 28 and and CC 10_1101-2020_12_31_424971 790 29 ProM ProM NNP 10_1101-2020_12_31_424971 790 30 in in IN 10_1101-2020_12_31_424971 790 31 the the DT 10_1101-2020_12_31_424971 790 32 ProL ProL NNP 10_1101-2020_12_31_424971 790 33 strain strain NN 10_1101-2020_12_31_424971 790 34 , , , 10_1101-2020_12_31_424971 790 35 where where WRB 10_1101-2020_12_31_424971 790 36 the the DT 10_1101-2020_12_31_424971 790 37 presence presence NN 10_1101-2020_12_31_424971 790 38 of of IN 10_1101-2020_12_31_424971 790 39 1247 1247 CD 10_1101-2020_12_31_424971 790 40 ProM ProM NNP 10_1101-2020_12_31_424971 790 41 ensured ensure VBD 10_1101-2020_12_31_424971 790 42 no no DT 10_1101-2020_12_31_424971 790 43 vacancy vacancy NN 10_1101-2020_12_31_424971 790 44 at at IN 10_1101-2020_12_31_424971 790 45 the the DT 10_1101-2020_12_31_424971 790 46 CCC CCC NNP 10_1101-2020_12_31_424971 790 47 - - HYPH 10_1101-2020_12_31_424971 790 48 C C NNP 10_1101-2020_12_31_424971 790 49 motif motif NN 10_1101-2020_12_31_424971 790 50 . . . 10_1101-2020_12_31_424971 791 1 The the DT 10_1101-2020_12_31_424971 791 2 increased increased JJ 10_1101-2020_12_31_424971 791 3 +1 +1 NNS 10_1101-2020_12_31_424971 791 4 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 791 5 in in IN 10_1101-2020_12_31_424971 791 6 the the DT 10_1101-2020_12_31_424971 791 7 m1G37 m1g37 NN 10_1101-2020_12_31_424971 791 8 – – : 10_1101-2020_12_31_424971 791 9 1248 1248 CD 10_1101-2020_12_31_424971 791 10 condition condition NN 10_1101-2020_12_31_424971 791 11 vs. vs. IN 10_1101-2020_12_31_424971 791 12 the the DT 10_1101-2020_12_31_424971 791 13 m1G37 m1g37 NN 10_1101-2020_12_31_424971 791 14 + + SYM 10_1101-2020_12_31_424971 791 15 condition condition NN 10_1101-2020_12_31_424971 791 16 indicates indicate VBZ 10_1101-2020_12_31_424971 791 17 that that IN 10_1101-2020_12_31_424971 791 18 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 791 19 and and CC 10_1101-2020_12_31_424971 791 20 ProL ProL NNP 10_1101-2020_12_31_424971 791 21 are be VBP 10_1101-2020_12_31_424971 791 22 each each DT 10_1101-2020_12_31_424971 791 23 an an DT 10_1101-2020_12_31_424971 791 24 active active JJ 10_1101-2020_12_31_424971 791 25 determinant determinant NN 10_1101-2020_12_31_424971 791 26 1249 1249 CD 10_1101-2020_12_31_424971 791 27 in in IN 10_1101-2020_12_31_424971 791 28 decoding decode VBG 10_1101-2020_12_31_424971 791 29 the the DT 10_1101-2020_12_31_424971 791 30 CCC CCC NNP 10_1101-2020_12_31_424971 791 31 - - HYPH 10_1101-2020_12_31_424971 791 32 C C NNP 10_1101-2020_12_31_424971 791 33 motif motif NN 10_1101-2020_12_31_424971 791 34 . . . 10_1101-2020_12_31_424971 792 1 d d NNP 10_1101-2020_12_31_424971 792 2 SufB2-mediated SufB2-mediated NNP 10_1101-2020_12_31_424971 792 3 insertion insertion NN 10_1101-2020_12_31_424971 792 4 of of IN 10_1101-2020_12_31_424971 792 5 non non JJ 10_1101-2020_12_31_424971 792 6 - - JJ 10_1101-2020_12_31_424971 792 7 proteinogenic proteinogenic JJ 10_1101-2020_12_31_424971 792 8 amino amino JJ 10_1101-2020_12_31_424971 792 9 acids acid NNS 10_1101-2020_12_31_424971 792 10 at at IN 10_1101-2020_12_31_424971 792 11 1250 1250 CD 10_1101-2020_12_31_424971 792 12 the the DT 10_1101-2020_12_31_424971 792 13 CCC CCC NNP 10_1101-2020_12_31_424971 792 14 - - HYPH 10_1101-2020_12_31_424971 792 15 C C NNP 10_1101-2020_12_31_424971 792 16 motif motif NN 10_1101-2020_12_31_424971 792 17 in in IN 10_1101-2020_12_31_424971 792 18 the the DT 10_1101-2020_12_31_424971 792 19 5th 5th JJ 10_1101-2020_12_31_424971 792 20 codon codon NN 10_1101-2020_12_31_424971 792 21 position position NN 10_1101-2020_12_31_424971 792 22 of of IN 10_1101-2020_12_31_424971 792 23 folA fola NN 10_1101-2020_12_31_424971 792 24 using use VBG 10_1101-2020_12_31_424971 792 25 [ [ -LRB- 10_1101-2020_12_31_424971 792 26 35S]-Met 35s]-met CD 10_1101-2020_12_31_424971 792 27 - - HYPH 10_1101-2020_12_31_424971 792 28 dependent dependent JJ 10_1101-2020_12_31_424971 792 29 in in FW 10_1101-2020_12_31_424971 792 30 vitro vitro FW 10_1101-2020_12_31_424971 792 31 translation translation NN 10_1101-2020_12_31_424971 792 32 . . . 10_1101-2020_12_31_424971 793 1 1251 1251 CD 10_1101-2020_12_31_424971 793 2 Reporters reporter NNS 10_1101-2020_12_31_424971 793 3 of of IN 10_1101-2020_12_31_424971 793 4 folA folA , 10_1101-2020_12_31_424971 793 5 are be VBP 10_1101-2020_12_31_424971 793 6 denoted denote VBN 10_1101-2020_12_31_424971 793 7 by by IN 10_1101-2020_12_31_424971 793 8 + + NNP 10_1101-2020_12_31_424971 793 9 / / SYM 10_1101-2020_12_31_424971 793 10 – – : 10_1101-2020_12_31_424971 793 11 CCC CCC NNP 10_1101-2020_12_31_424971 793 12 - - HYPH 10_1101-2020_12_31_424971 793 13 C C NNP 10_1101-2020_12_31_424971 793 14 , , , 10_1101-2020_12_31_424971 793 15 where where WRB 10_1101-2020_12_31_424971 793 16 “ " `` 10_1101-2020_12_31_424971 793 17 + + CC 10_1101-2020_12_31_424971 793 18 ” " '' 10_1101-2020_12_31_424971 793 19 and and CC 10_1101-2020_12_31_424971 793 20 “ " `` 10_1101-2020_12_31_424971 793 21 – – : 10_1101-2020_12_31_424971 793 22 ” " '' 10_1101-2020_12_31_424971 793 23 indicate indicate VBP 10_1101-2020_12_31_424971 793 24 constructs construct NNS 10_1101-2020_12_31_424971 793 25 with with IN 10_1101-2020_12_31_424971 793 26 and and CC 10_1101-2020_12_31_424971 793 27 1252 1252 CD 10_1101-2020_12_31_424971 793 28 without without IN 10_1101-2020_12_31_424971 793 29 the the DT 10_1101-2020_12_31_424971 793 30 CCC CCC NNP 10_1101-2020_12_31_424971 793 31 - - HYPH 10_1101-2020_12_31_424971 793 32 C C NNP 10_1101-2020_12_31_424971 793 33 motif motif NN 10_1101-2020_12_31_424971 793 34 . . . 10_1101-2020_12_31_424971 794 1 SDS SDS NNP 10_1101-2020_12_31_424971 794 2 - - HYPH 10_1101-2020_12_31_424971 794 3 PAGE PAGE NNP 10_1101-2020_12_31_424971 794 4 analysis analysis NN 10_1101-2020_12_31_424971 794 5 identifies identify VBZ 10_1101-2020_12_31_424971 794 6 full full JJ 10_1101-2020_12_31_424971 794 7 - - HYPH 10_1101-2020_12_31_424971 794 8 length length NN 10_1101-2020_12_31_424971 794 9 DHFR DHFR NNP 10_1101-2020_12_31_424971 794 10 resulting result VBG 10_1101-2020_12_31_424971 794 11 from from IN 10_1101-2020_12_31_424971 794 12 a a DT 10_1101-2020_12_31_424971 794 13 +1 +1 NN 10_1101-2020_12_31_424971 794 14 - - SYM 10_1101-2020_12_31_424971 794 15 1253 1253 CD 10_1101-2020_12_31_424971 794 16 frameshift frameshift NN 10_1101-2020_12_31_424971 794 17 event event NN 10_1101-2020_12_31_424971 794 18 at at IN 10_1101-2020_12_31_424971 794 19 the the DT 10_1101-2020_12_31_424971 794 20 CCC CCC NNP 10_1101-2020_12_31_424971 794 21 - - HYPH 10_1101-2020_12_31_424971 794 22 C C NNP 10_1101-2020_12_31_424971 794 23 motif motif NN 10_1101-2020_12_31_424971 794 24 by by IN 10_1101-2020_12_31_424971 794 25 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 794 26 pre pre JJ 10_1101-2020_12_31_424971 794 27 - - JJ 10_1101-2020_12_31_424971 794 28 aminoacylated aminoacylate VBN 10_1101-2020_12_31_424971 794 29 with with IN 10_1101-2020_12_31_424971 794 30 the the DT 10_1101-2020_12_31_424971 794 31 amino amino JJ 10_1101-2020_12_31_424971 794 32 acid acid NN 10_1101-2020_12_31_424971 794 33 shown show VBN 10_1101-2020_12_31_424971 794 34 at at IN 10_1101-2020_12_31_424971 794 35 1254 1254 CD 10_1101-2020_12_31_424971 794 36 the the DT 10_1101-2020_12_31_424971 794 37 top top NN 10_1101-2020_12_31_424971 794 38 of of IN 10_1101-2020_12_31_424971 794 39 each each DT 10_1101-2020_12_31_424971 794 40 lane lane NN 10_1101-2020_12_31_424971 794 41 , , , 10_1101-2020_12_31_424971 794 42 a a DT 10_1101-2020_12_31_424971 794 43 DC DC NNP 10_1101-2020_12_31_424971 794 44 fragment fragment NN 10_1101-2020_12_31_424971 794 45 resulting result VBG 10_1101-2020_12_31_424971 794 46 from from IN 10_1101-2020_12_31_424971 794 47 lack lack NN 10_1101-2020_12_31_424971 794 48 of of IN 10_1101-2020_12_31_424971 794 49 the the DT 10_1101-2020_12_31_424971 794 50 +1-frameshift +1-frameshift NN 10_1101-2020_12_31_424971 794 51 event event NN 10_1101-2020_12_31_424971 794 52 , , , 10_1101-2020_12_31_424971 794 53 and and CC 10_1101-2020_12_31_424971 794 54 a a DT 10_1101-2020_12_31_424971 794 55 DN DN NNP 10_1101-2020_12_31_424971 794 56 1255 1255 CD 10_1101-2020_12_31_424971 794 57 fragment fragment NN 10_1101-2020_12_31_424971 794 58 resulting result VBG 10_1101-2020_12_31_424971 794 59 from from IN 10_1101-2020_12_31_424971 794 60 translation translation NN 10_1101-2020_12_31_424971 794 61 initiation initiation NN 10_1101-2020_12_31_424971 794 62 at at IN 10_1101-2020_12_31_424971 794 63 the the DT 10_1101-2020_12_31_424971 794 64 AUG AUG NNP 10_1101-2020_12_31_424971 794 65 codon codon NN 10_1101-2020_12_31_424971 794 66 likely likely RB 10_1101-2020_12_31_424971 794 67 at at IN 10_1101-2020_12_31_424971 794 68 position position NN 10_1101-2020_12_31_424971 794 69 17 17 CD 10_1101-2020_12_31_424971 794 70 or or CC 10_1101-2020_12_31_424971 794 71 21 21 CD 10_1101-2020_12_31_424971 794 72 1256 1256 CD 10_1101-2020_12_31_424971 794 73 downstream downstream RB 10_1101-2020_12_31_424971 794 74 from from IN 10_1101-2020_12_31_424971 794 75 the the DT 10_1101-2020_12_31_424971 794 76 CCC CCC NNP 10_1101-2020_12_31_424971 794 77 - - HYPH 10_1101-2020_12_31_424971 794 78 C C NNP 10_1101-2020_12_31_424971 794 79 motif motif NN 10_1101-2020_12_31_424971 794 80 . . . 10_1101-2020_12_31_424971 795 1 Gel gel NN 10_1101-2020_12_31_424971 795 2 samples sample NNS 10_1101-2020_12_31_424971 795 3 were be VBD 10_1101-2020_12_31_424971 795 4 derived derive VBN 10_1101-2020_12_31_424971 795 5 from from IN 10_1101-2020_12_31_424971 795 6 the the DT 10_1101-2020_12_31_424971 795 7 same same JJ 10_1101-2020_12_31_424971 795 8 experiment experiment NN 10_1101-2020_12_31_424971 795 9 , , , 10_1101-2020_12_31_424971 795 10 which which WDT 10_1101-2020_12_31_424971 795 11 1257 1257 CD 10_1101-2020_12_31_424971 795 12 was be VBD 10_1101-2020_12_31_424971 795 13 performed perform VBN 10_1101-2020_12_31_424971 795 14 five five CD 10_1101-2020_12_31_424971 795 15 times time NNS 10_1101-2020_12_31_424971 795 16 with with IN 10_1101-2020_12_31_424971 795 17 similar similar JJ 10_1101-2020_12_31_424971 795 18 results result NNS 10_1101-2020_12_31_424971 795 19 . . . 10_1101-2020_12_31_424971 796 1 Gels gel NNS 10_1101-2020_12_31_424971 796 2 for for IN 10_1101-2020_12_31_424971 796 3 each each DT 10_1101-2020_12_31_424971 796 4 experiment experiment NN 10_1101-2020_12_31_424971 796 5 were be VBD 10_1101-2020_12_31_424971 796 6 processed process VBN 10_1101-2020_12_31_424971 796 7 in in IN 10_1101-2020_12_31_424971 796 8 parallel parallel NN 10_1101-2020_12_31_424971 796 9 . . . 10_1101-2020_12_31_424971 797 1 1258 1258 CD 10_1101-2020_12_31_424971 797 2 Lane Lane NNP 10_1101-2020_12_31_424971 797 3 1 1 CD 10_1101-2020_12_31_424971 797 4 : : : 10_1101-2020_12_31_424971 797 5 full full JJ 10_1101-2020_12_31_424971 797 6 - - HYPH 10_1101-2020_12_31_424971 797 7 length length NN 10_1101-2020_12_31_424971 797 8 DHFR DHFR NNP 10_1101-2020_12_31_424971 797 9 as as IN 10_1101-2020_12_31_424971 797 10 the the DT 10_1101-2020_12_31_424971 797 11 molecular molecular JJ 10_1101-2020_12_31_424971 797 12 marker marker NN 10_1101-2020_12_31_424971 797 13 ; ; : 10_1101-2020_12_31_424971 797 14 deacyl deacyl NNP 10_1101-2020_12_31_424971 797 15 : : : 10_1101-2020_12_31_424971 797 16 deacylated deacylate VBN 10_1101-2020_12_31_424971 797 17 tRNA tRNA NNP 10_1101-2020_12_31_424971 797 18 . . . 10_1101-2020_12_31_424971 798 1 1259 1259 CD 10_1101-2020_12_31_424971 798 2 1260 1260 CD 10_1101-2020_12_31_424971 798 3 Figure Figure NNP 10_1101-2020_12_31_424971 798 4 3 3 CD 10_1101-2020_12_31_424971 798 5 . . . 10_1101-2020_12_31_424971 799 1 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 799 2 uses use VBZ 10_1101-2020_12_31_424971 799 3 a a DT 10_1101-2020_12_31_424971 799 4 triplet triplet NN 10_1101-2020_12_31_424971 799 5 anticodon anticodon NN 10_1101-2020_12_31_424971 799 6 - - HYPH 10_1101-2020_12_31_424971 799 7 codon codon NN 10_1101-2020_12_31_424971 799 8 pairing pairing NN 10_1101-2020_12_31_424971 799 9 scheme scheme NN 10_1101-2020_12_31_424971 799 10 at at IN 10_1101-2020_12_31_424971 799 11 the the DT 10_1101-2020_12_31_424971 799 12 A A NNP 10_1101-2020_12_31_424971 799 13 site site NN 10_1101-2020_12_31_424971 799 14 . . . 10_1101-2020_12_31_424971 800 1 a a DT 10_1101-2020_12_31_424971 800 2 GTP GTP NNP 10_1101-2020_12_31_424971 800 3 1261 1261 CD 10_1101-2020_12_31_424971 800 4 hydrolysis hydrolysis NN 10_1101-2020_12_31_424971 800 5 by by IN 10_1101-2020_12_31_424971 800 6 EF EF NNP 10_1101-2020_12_31_424971 800 7 - - HYPH 10_1101-2020_12_31_424971 800 8 Tu Tu NNP 10_1101-2020_12_31_424971 800 9 as as IN 10_1101-2020_12_31_424971 800 10 a a DT 10_1101-2020_12_31_424971 800 11 function function NN 10_1101-2020_12_31_424971 800 12 of of IN 10_1101-2020_12_31_424971 800 13 time time NN 10_1101-2020_12_31_424971 800 14 for for IN 10_1101-2020_12_31_424971 800 15 delivery delivery NN 10_1101-2020_12_31_424971 800 16 of of IN 10_1101-2020_12_31_424971 800 17 G37- G37- NNP 10_1101-2020_12_31_424971 800 18 or or CC 10_1101-2020_12_31_424971 800 19 native native JJ 10_1101-2020_12_31_424971 800 20 - - HYPH 10_1101-2020_12_31_424971 800 21 state state NN 10_1101-2020_12_31_424971 800 22 SufB2- sufb2- NN 10_1101-2020_12_31_424971 800 23 or or CC 10_1101-2020_12_31_424971 800 24 ProL ProL NNP 10_1101-2020_12_31_424971 800 25 - - HYPH 10_1101-2020_12_31_424971 800 26 TC TC NNP 10_1101-2020_12_31_424971 800 27 1262 1262 CD 10_1101-2020_12_31_424971 800 28 to to IN 10_1101-2020_12_31_424971 800 29 the the DT 10_1101-2020_12_31_424971 800 30 A A NNP 10_1101-2020_12_31_424971 800 31 site site NN 10_1101-2020_12_31_424971 800 32 of of IN 10_1101-2020_12_31_424971 800 33 a a DT 10_1101-2020_12_31_424971 800 34 70S 70S NNP 10_1101-2020_12_31_424971 800 35 IC IC NNP 10_1101-2020_12_31_424971 800 36 . . . 10_1101-2020_12_31_424971 801 1 Although although IN 10_1101-2020_12_31_424971 801 2 the the DT 10_1101-2020_12_31_424971 801 3 concentration concentration NN 10_1101-2020_12_31_424971 801 4 of of IN 10_1101-2020_12_31_424971 801 5 TCs tc NNS 10_1101-2020_12_31_424971 801 6 was be VBD 10_1101-2020_12_31_424971 801 7 limiting limit VBG 10_1101-2020_12_31_424971 801 8 , , , 10_1101-2020_12_31_424971 801 9 which which WDT 10_1101-2020_12_31_424971 801 10 would would MD 10_1101-2020_12_31_424971 801 11 limit limit VB 10_1101-2020_12_31_424971 801 12 the the DT 10_1101-2020_12_31_424971 801 13 1263 1263 CD 10_1101-2020_12_31_424971 801 14 rate rate NN 10_1101-2020_12_31_424971 801 15 of of IN 10_1101-2020_12_31_424971 801 16 binding binding NN 10_1101-2020_12_31_424971 801 17 of of IN 10_1101-2020_12_31_424971 801 18 TCs tc NNS 10_1101-2020_12_31_424971 801 19 to to IN 10_1101-2020_12_31_424971 801 20 the the DT 10_1101-2020_12_31_424971 801 21 70S 70S NNP 10_1101-2020_12_31_424971 801 22 IC IC NNP 10_1101-2020_12_31_424971 801 23 , , , 10_1101-2020_12_31_424971 801 24 the the DT 10_1101-2020_12_31_424971 801 25 observed observed JJ 10_1101-2020_12_31_424971 801 26 differences difference NNS 10_1101-2020_12_31_424971 801 27 in in IN 10_1101-2020_12_31_424971 801 28 the the DT 10_1101-2020_12_31_424971 801 29 yield yield NN 10_1101-2020_12_31_424971 801 30 of of IN 10_1101-2020_12_31_424971 801 31 GTPase GTPase NNP 10_1101-2020_12_31_424971 801 32 activity activity NN 10_1101-2020_12_31_424971 801 33 1264 1264 CD 10_1101-2020_12_31_424971 801 34 indicated indicate VBD 10_1101-2020_12_31_424971 801 35 that that IN 10_1101-2020_12_31_424971 801 36 binding bind VBG 10_1101-2020_12_31_424971 801 37 was be VBD 10_1101-2020_12_31_424971 801 38 not not RB 10_1101-2020_12_31_424971 801 39 the the DT 10_1101-2020_12_31_424971 801 40 sole sole JJ 10_1101-2020_12_31_424971 801 41 determinant determinant NN 10_1101-2020_12_31_424971 801 42 , , , 10_1101-2020_12_31_424971 801 43 but but CC 10_1101-2020_12_31_424971 801 44 that that IN 10_1101-2020_12_31_424971 801 45 other other JJ 10_1101-2020_12_31_424971 801 46 factors factor NNS 10_1101-2020_12_31_424971 801 47 , , , 10_1101-2020_12_31_424971 801 48 such such JJ 10_1101-2020_12_31_424971 801 49 as as IN 10_1101-2020_12_31_424971 801 50 the the DT 10_1101-2020_12_31_424971 801 51 identity identity NN 10_1101-2020_12_31_424971 801 52 1265 1265 CD 10_1101-2020_12_31_424971 801 53 and and CC 10_1101-2020_12_31_424971 801 54 the the DT 10_1101-2020_12_31_424971 801 55 methylation methylation NN 10_1101-2020_12_31_424971 801 56 state state NN 10_1101-2020_12_31_424971 801 57 of of IN 10_1101-2020_12_31_424971 801 58 the the DT 10_1101-2020_12_31_424971 801 59 tRNA trna NN 10_1101-2020_12_31_424971 801 60 , , , 10_1101-2020_12_31_424971 801 61 affected affect VBD 10_1101-2020_12_31_424971 801 62 the the DT 10_1101-2020_12_31_424971 801 63 GTPase GTPase NNP 10_1101-2020_12_31_424971 801 64 activity activity NN 10_1101-2020_12_31_424971 801 65 . . . 10_1101-2020_12_31_424971 802 1 b b LS 10_1101-2020_12_31_424971 802 2 Dipeptide Dipeptide NNP 10_1101-2020_12_31_424971 802 3 fMP fmp CD 10_1101-2020_12_31_424971 802 4 formation formation NN 10_1101-2020_12_31_424971 802 5 1266 1266 CD 10_1101-2020_12_31_424971 802 6 as as IN 10_1101-2020_12_31_424971 802 7 a a DT 10_1101-2020_12_31_424971 802 8 function function NN 10_1101-2020_12_31_424971 802 9 of of IN 10_1101-2020_12_31_424971 802 10 time time NN 10_1101-2020_12_31_424971 802 11 for for IN 10_1101-2020_12_31_424971 802 12 delivery delivery NN 10_1101-2020_12_31_424971 802 13 of of IN 10_1101-2020_12_31_424971 802 14 G37- G37- NNP 10_1101-2020_12_31_424971 802 15 or or CC 10_1101-2020_12_31_424971 802 16 native native JJ 10_1101-2020_12_31_424971 802 17 - - HYPH 10_1101-2020_12_31_424971 802 18 state state NN 10_1101-2020_12_31_424971 802 19 SufB2- sufb2- NN 10_1101-2020_12_31_424971 802 20 or or CC 10_1101-2020_12_31_424971 802 21 ProL ProL NNP 10_1101-2020_12_31_424971 802 22 - - HYPH 10_1101-2020_12_31_424971 802 23 TC TC NNP 10_1101-2020_12_31_424971 802 24 to to IN 10_1101-2020_12_31_424971 802 25 the the DT 10_1101-2020_12_31_424971 802 26 A A NNP 10_1101-2020_12_31_424971 802 27 site site NN 10_1101-2020_12_31_424971 802 28 of of IN 10_1101-2020_12_31_424971 802 29 a a DT 10_1101-2020_12_31_424971 802 30 70S 70S NNP 10_1101-2020_12_31_424971 802 31 1267 1267 CD 10_1101-2020_12_31_424971 802 32 .CC .cc SYM 10_1101-2020_12_31_424971 802 33 - - HYPH 10_1101-2020_12_31_424971 802 34 BY by IN 10_1101-2020_12_31_424971 802 35 - - HYPH 10_1101-2020_12_31_424971 802 36 NC NC NNP 10_1101-2020_12_31_424971 802 37 - - HYPH 10_1101-2020_12_31_424971 802 38 ND ND NNP 10_1101-2020_12_31_424971 802 39 4.0 4.0 CD 10_1101-2020_12_31_424971 802 40 International International NNP 10_1101-2020_12_31_424971 802 41 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 802 42 under under IN 10_1101-2020_12_31_424971 802 43 a a DT 10_1101-2020_12_31_424971 802 44 ( ( -LRB- 10_1101-2020_12_31_424971 802 45 which which WDT 10_1101-2020_12_31_424971 802 46 was be VBD 10_1101-2020_12_31_424971 802 47 not not RB 10_1101-2020_12_31_424971 802 48 certified certify VBN 10_1101-2020_12_31_424971 802 49 by by IN 10_1101-2020_12_31_424971 802 50 peer peer NN 10_1101-2020_12_31_424971 802 51 review review NN 10_1101-2020_12_31_424971 802 52 ) ) -RRB- 10_1101-2020_12_31_424971 802 53 is be VBZ 10_1101-2020_12_31_424971 802 54 the the DT 10_1101-2020_12_31_424971 802 55 author author NN 10_1101-2020_12_31_424971 802 56 / / SYM 10_1101-2020_12_31_424971 802 57 funder funder NN 10_1101-2020_12_31_424971 802 58 , , , 10_1101-2020_12_31_424971 802 59 who who WP 10_1101-2020_12_31_424971 802 60 has have VBZ 10_1101-2020_12_31_424971 802 61 granted grant VBN 10_1101-2020_12_31_424971 802 62 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 802 63 a a DT 10_1101-2020_12_31_424971 802 64 license license NN 10_1101-2020_12_31_424971 802 65 to to TO 10_1101-2020_12_31_424971 802 66 display display VB 10_1101-2020_12_31_424971 802 67 the the DT 10_1101-2020_12_31_424971 802 68 preprint preprint NN 10_1101-2020_12_31_424971 802 69 in in IN 10_1101-2020_12_31_424971 802 70 perpetuity perpetuity NN 10_1101-2020_12_31_424971 802 71 . . . 10_1101-2020_12_31_424971 803 1 It -PRON- PRP 10_1101-2020_12_31_424971 803 2 is be VBZ 10_1101-2020_12_31_424971 803 3 made make VBN 10_1101-2020_12_31_424971 803 4 The the DT 10_1101-2020_12_31_424971 803 5 copyright copyright NN 10_1101-2020_12_31_424971 803 6 holder holder NN 10_1101-2020_12_31_424971 803 7 for for IN 10_1101-2020_12_31_424971 803 8 this this DT 10_1101-2020_12_31_424971 803 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 803 10 version version NN 10_1101-2020_12_31_424971 803 11 posted post VBD 10_1101-2020_12_31_424971 803 12 January January NNP 10_1101-2020_12_31_424971 803 13 2 2 CD 10_1101-2020_12_31_424971 803 14 , , , 10_1101-2020_12_31_424971 803 15 2021 2021 CD 10_1101-2020_12_31_424971 803 16 . . . 10_1101-2020_12_31_424971 803 17 ; ; : 10_1101-2020_12_31_424971 803 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 803 19 : : : 10_1101-2020_12_31_424971 803 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 803 21 preprint preprint NN 10_1101-2020_12_31_424971 803 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NNP 10_1101-2020_12_31_424971 803 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 803 24 49 49 CD 10_1101-2020_12_31_424971 803 25 IC IC NNP 10_1101-2020_12_31_424971 803 26 . . . 10_1101-2020_12_31_424971 804 1 Due due IN 10_1101-2020_12_31_424971 804 2 to to IN 10_1101-2020_12_31_424971 804 3 the the DT 10_1101-2020_12_31_424971 804 4 limiting limit VBG 10_1101-2020_12_31_424971 804 5 concentration concentration NN 10_1101-2020_12_31_424971 804 6 of of IN 10_1101-2020_12_31_424971 804 7 the the DT 10_1101-2020_12_31_424971 804 8 70S 70S NNP 10_1101-2020_12_31_424971 804 9 IC IC NNP 10_1101-2020_12_31_424971 804 10 , , , 10_1101-2020_12_31_424971 804 11 which which WDT 10_1101-2020_12_31_424971 804 12 did do VBD 10_1101-2020_12_31_424971 804 13 not not RB 10_1101-2020_12_31_424971 804 14 include include VB 10_1101-2020_12_31_424971 804 15 the the DT 10_1101-2020_12_31_424971 804 16 tRNA trna NN 10_1101-2020_12_31_424971 804 17 substrate substrate NN 10_1101-2020_12_31_424971 804 18 , , , 10_1101-2020_12_31_424971 804 19 the the DT 10_1101-2020_12_31_424971 804 20 1268 1268 CD 10_1101-2020_12_31_424971 804 21 yield yield NN 10_1101-2020_12_31_424971 804 22 of of IN 10_1101-2020_12_31_424971 804 23 di- di- JJ 10_1101-2020_12_31_424971 804 24 or or CC 10_1101-2020_12_31_424971 804 25 tri tri JJ 10_1101-2020_12_31_424971 804 26 - - HYPH 10_1101-2020_12_31_424971 804 27 peptide peptide JJ 10_1101-2020_12_31_424971 804 28 formation formation NN 10_1101-2020_12_31_424971 804 29 assays assays RB 10_1101-2020_12_31_424971 804 30 was be VBD 10_1101-2020_12_31_424971 804 31 constant constant JJ 10_1101-2020_12_31_424971 804 32 even even RB 10_1101-2020_12_31_424971 804 33 with with IN 10_1101-2020_12_31_424971 804 34 different different JJ 10_1101-2020_12_31_424971 804 35 tRNAs trna NNS 10_1101-2020_12_31_424971 804 36 in in IN 10_1101-2020_12_31_424971 804 37 TCs tc NNS 10_1101-2020_12_31_424971 804 38 . . . 10_1101-2020_12_31_424971 805 1 c c LS 10_1101-2020_12_31_424971 805 2 The the DT 10_1101-2020_12_31_424971 805 3 1269 1269 CD 10_1101-2020_12_31_424971 805 4 yield yield NN 10_1101-2020_12_31_424971 805 5 of of IN 10_1101-2020_12_31_424971 805 6 fMP fMP NNS 10_1101-2020_12_31_424971 805 7 and and CC 10_1101-2020_12_31_424971 805 8 fMR fMR NNS 10_1101-2020_12_31_424971 805 9 in in IN 10_1101-2020_12_31_424971 805 10 dipeptide dipeptide NN 10_1101-2020_12_31_424971 805 11 formation formation NN 10_1101-2020_12_31_424971 805 12 assays assays RB 10_1101-2020_12_31_424971 805 13 in in IN 10_1101-2020_12_31_424971 805 14 which which WDT 10_1101-2020_12_31_424971 805 15 equimolar equimolar JJ 10_1101-2020_12_31_424971 805 16 mixtures mixture NNS 10_1101-2020_12_31_424971 805 17 of of IN 10_1101-2020_12_31_424971 805 18 native native JJ 10_1101-2020_12_31_424971 805 19 - - HYPH 10_1101-2020_12_31_424971 805 20 state state NN 10_1101-2020_12_31_424971 805 21 1270 1270 CD 10_1101-2020_12_31_424971 805 22 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 805 23 , , , 10_1101-2020_12_31_424971 805 24 carrying carry VBG 10_1101-2020_12_31_424971 805 25 Pro Pro NNP 10_1101-2020_12_31_424971 805 26 and/or and/or CC 10_1101-2020_12_31_424971 805 27 Arg Arg NNP 10_1101-2020_12_31_424971 805 28 , , , 10_1101-2020_12_31_424971 805 29 and/or and/or CC 10_1101-2020_12_31_424971 805 30 native native JJ 10_1101-2020_12_31_424971 805 31 - - HYPH 10_1101-2020_12_31_424971 805 32 state state NN 10_1101-2020_12_31_424971 805 33 ProL ProL NNP 10_1101-2020_12_31_424971 805 34 - - HYPH 10_1101-2020_12_31_424971 805 35 TC TC NNP 10_1101-2020_12_31_424971 805 36 , , , 10_1101-2020_12_31_424971 805 37 carrying carry VBG 10_1101-2020_12_31_424971 805 38 Pro Pro NNP 10_1101-2020_12_31_424971 805 39 and/or and/or CC 10_1101-2020_12_31_424971 805 40 Arg Arg NNP 10_1101-2020_12_31_424971 805 41 , , , 10_1101-2020_12_31_424971 805 42 are be VBP 10_1101-2020_12_31_424971 805 43 1271 1271 CD 10_1101-2020_12_31_424971 805 44 delivered deliver VBN 10_1101-2020_12_31_424971 805 45 to to IN 10_1101-2020_12_31_424971 805 46 70S 70S NNP 10_1101-2020_12_31_424971 805 47 ICs ic NNS 10_1101-2020_12_31_424971 805 48 . . . 10_1101-2020_12_31_424971 806 1 The the DT 10_1101-2020_12_31_424971 806 2 mRNA mrna NN 10_1101-2020_12_31_424971 806 3 in in IN 10_1101-2020_12_31_424971 806 4 70S 70S NNP 10_1101-2020_12_31_424971 806 5 ICs ic NNS 10_1101-2020_12_31_424971 806 6 in in IN 10_1101-2020_12_31_424971 806 7 ( ( -LRB- 10_1101-2020_12_31_424971 806 8 A A NNP 10_1101-2020_12_31_424971 806 9 - - HYPH 10_1101-2020_12_31_424971 806 10 C C NNP 10_1101-2020_12_31_424971 806 11 ) ) -RRB- 10_1101-2020_12_31_424971 806 12 is be VBZ 10_1101-2020_12_31_424971 806 13 AUG AUG NNP 10_1101-2020_12_31_424971 806 14 - - HYPH 10_1101-2020_12_31_424971 806 15 CCC CCC NNP 10_1101-2020_12_31_424971 806 16 - - HYPH 10_1101-2020_12_31_424971 806 17 CGU CGU NNP 10_1101-2020_12_31_424971 806 18 - - HYPH 10_1101-2020_12_31_424971 806 19 U. U. NNP 10_1101-2020_12_31_424971 807 1 d d LS 10_1101-2020_12_31_424971 807 2 Dipeptide Dipeptide NNP 10_1101-2020_12_31_424971 807 3 formation formation NN 10_1101-2020_12_31_424971 807 4 1272 1272 CD 10_1101-2020_12_31_424971 807 5 rate rate NN 10_1101-2020_12_31_424971 807 6 kfMP kfMP NNP 10_1101-2020_12_31_424971 807 7 , , , 10_1101-2020_12_31_424971 807 8 obs obs NN 10_1101-2020_12_31_424971 807 9 for for IN 10_1101-2020_12_31_424971 807 10 delivery delivery NN 10_1101-2020_12_31_424971 807 11 of of IN 10_1101-2020_12_31_424971 807 12 G37-state G37-state NNP 10_1101-2020_12_31_424971 807 13 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 807 14 to to IN 10_1101-2020_12_31_424971 807 15 70S 70S NNP 10_1101-2020_12_31_424971 807 16 ICs ic NNS 10_1101-2020_12_31_424971 807 17 containing contain VBG 10_1101-2020_12_31_424971 807 18 sequence sequence NN 10_1101-2020_12_31_424971 807 19 variants variant NNS 10_1101-2020_12_31_424971 807 20 of of IN 10_1101-2020_12_31_424971 807 21 the the DT 10_1101-2020_12_31_424971 807 22 1273 1273 CD 10_1101-2020_12_31_424971 807 23 CCC CCC NNP 10_1101-2020_12_31_424971 807 24 - - HYPH 10_1101-2020_12_31_424971 807 25 C C NNP 10_1101-2020_12_31_424971 807 26 motif motif NN 10_1101-2020_12_31_424971 807 27 in in IN 10_1101-2020_12_31_424971 807 28 the the DT 10_1101-2020_12_31_424971 807 29 A A NNP 10_1101-2020_12_31_424971 807 30 site site NN 10_1101-2020_12_31_424971 807 31 . . . 10_1101-2020_12_31_424971 808 1 In in IN 10_1101-2020_12_31_424971 808 2 panels panel NNS 10_1101-2020_12_31_424971 808 3 a a XX 10_1101-2020_12_31_424971 808 4 , , , 10_1101-2020_12_31_424971 808 5 b b NN 10_1101-2020_12_31_424971 808 6 , , , 10_1101-2020_12_31_424971 808 7 the the DT 10_1101-2020_12_31_424971 808 8 bars bar NNS 10_1101-2020_12_31_424971 808 9 in in IN 10_1101-2020_12_31_424971 808 10 the the DT 10_1101-2020_12_31_424971 808 11 graphs graph NNS 10_1101-2020_12_31_424971 808 12 are be VBP 10_1101-2020_12_31_424971 808 13 SD sd NN 10_1101-2020_12_31_424971 808 14 of of IN 10_1101-2020_12_31_424971 808 15 three three CD 10_1101-2020_12_31_424971 808 16 independent independent NN 10_1101-2020_12_31_424971 808 17 ( ( -LRB- 10_1101-2020_12_31_424971 808 18 n n IN 10_1101-2020_12_31_424971 808 19 1274 1274 CD 10_1101-2020_12_31_424971 808 20 = = SYM 10_1101-2020_12_31_424971 808 21 3 3 LS 10_1101-2020_12_31_424971 808 22 ) ) -RRB- 10_1101-2020_12_31_424971 808 23 experiments experiment NNS 10_1101-2020_12_31_424971 808 24 , , , 10_1101-2020_12_31_424971 808 25 in in IN 10_1101-2020_12_31_424971 808 26 panel panel NN 10_1101-2020_12_31_424971 808 27 c c NNP 10_1101-2020_12_31_424971 808 28 , , , 10_1101-2020_12_31_424971 808 29 the the DT 10_1101-2020_12_31_424971 808 30 bars bar NNS 10_1101-2020_12_31_424971 808 31 in in IN 10_1101-2020_12_31_424971 808 32 the the DT 10_1101-2020_12_31_424971 808 33 graphs graph NNS 10_1101-2020_12_31_424971 808 34 are be VBP 10_1101-2020_12_31_424971 808 35 SD sd NN 10_1101-2020_12_31_424971 808 36 of of IN 10_1101-2020_12_31_424971 808 37 four four CD 10_1101-2020_12_31_424971 808 38 independent independent NN 10_1101-2020_12_31_424971 808 39 ( ( -LRB- 10_1101-2020_12_31_424971 808 40 n n JJ 10_1101-2020_12_31_424971 808 41 = = SYM 10_1101-2020_12_31_424971 808 42 4 4 LS 10_1101-2020_12_31_424971 808 43 ) ) -RRB- 10_1101-2020_12_31_424971 808 44 experiments experiment NNS 10_1101-2020_12_31_424971 808 45 , , , 10_1101-2020_12_31_424971 808 46 1275 1275 CD 10_1101-2020_12_31_424971 808 47 and and CC 10_1101-2020_12_31_424971 808 48 in in IN 10_1101-2020_12_31_424971 808 49 panel panel NN 10_1101-2020_12_31_424971 808 50 d d NN 10_1101-2020_12_31_424971 808 51 , , , 10_1101-2020_12_31_424971 808 52 the the DT 10_1101-2020_12_31_424971 808 53 bars bar NNS 10_1101-2020_12_31_424971 808 54 in in IN 10_1101-2020_12_31_424971 808 55 the the DT 10_1101-2020_12_31_424971 808 56 graphs graph NNS 10_1101-2020_12_31_424971 808 57 are be VBP 10_1101-2020_12_31_424971 808 58 SD sd NN 10_1101-2020_12_31_424971 808 59 of of IN 10_1101-2020_12_31_424971 808 60 three three CD 10_1101-2020_12_31_424971 808 61 or or CC 10_1101-2020_12_31_424971 808 62 four four CD 10_1101-2020_12_31_424971 808 63 independent independent NN 10_1101-2020_12_31_424971 808 64 ( ( -LRB- 10_1101-2020_12_31_424971 808 65 n n NN 10_1101-2020_12_31_424971 808 66 = = SYM 10_1101-2020_12_31_424971 808 67 3 3 CD 10_1101-2020_12_31_424971 808 68 or or CC 10_1101-2020_12_31_424971 808 69 4 4 CD 10_1101-2020_12_31_424971 808 70 ) ) -RRB- 10_1101-2020_12_31_424971 808 71 experiments experiment NNS 10_1101-2020_12_31_424971 808 72 . . . 10_1101-2020_12_31_424971 809 1 1276 1276 CD 10_1101-2020_12_31_424971 809 2 All all DT 10_1101-2020_12_31_424971 809 3 data datum NNS 10_1101-2020_12_31_424971 809 4 are be VBP 10_1101-2020_12_31_424971 809 5 presented present VBN 10_1101-2020_12_31_424971 809 6 as as IN 10_1101-2020_12_31_424971 809 7 mean mean JJ 10_1101-2020_12_31_424971 809 8 values value NNS 10_1101-2020_12_31_424971 809 9 ± ± CD 10_1101-2020_12_31_424971 809 10 SD sd NN 10_1101-2020_12_31_424971 809 11 . . . 10_1101-2020_12_31_424971 810 1 ∆t ∆t LS 10_1101-2020_12_31_424971 810 2 : : : 10_1101-2020_12_31_424971 810 3 a a DT 10_1101-2020_12_31_424971 810 4 time time NN 10_1101-2020_12_31_424971 810 5 interval interval NN 10_1101-2020_12_31_424971 810 6 , , , 10_1101-2020_12_31_424971 810 7 ND ND NNP 10_1101-2020_12_31_424971 810 8 : : : 10_1101-2020_12_31_424971 810 9 not not RB 10_1101-2020_12_31_424971 810 10 detected detect VBN 10_1101-2020_12_31_424971 810 11 . . . 10_1101-2020_12_31_424971 811 1 1277 1277 CD 10_1101-2020_12_31_424971 811 2 1278 1278 CD 10_1101-2020_12_31_424971 811 3 Figure Figure NNP 10_1101-2020_12_31_424971 811 4 4 4 CD 10_1101-2020_12_31_424971 811 5 . . . 10_1101-2020_12_31_424971 812 1 Plasticity plasticity NN 10_1101-2020_12_31_424971 812 2 of of IN 10_1101-2020_12_31_424971 812 3 SufB2-induced SufB2-induced NNP 10_1101-2020_12_31_424971 812 4 +1 +1 NNS 10_1101-2020_12_31_424971 812 5 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 812 6 . . . 10_1101-2020_12_31_424971 813 1 a a DT 10_1101-2020_12_31_424971 813 2 fMP fMP NNP 10_1101-2020_12_31_424971 813 3 formation formation NN 10_1101-2020_12_31_424971 813 4 as as IN 10_1101-2020_12_31_424971 813 5 a a DT 10_1101-2020_12_31_424971 813 6 function function NN 10_1101-2020_12_31_424971 813 7 of of IN 10_1101-2020_12_31_424971 813 8 time time NN 10_1101-2020_12_31_424971 813 9 1279 1279 CD 10_1101-2020_12_31_424971 813 10 upon upon IN 10_1101-2020_12_31_424971 813 11 delivery delivery NN 10_1101-2020_12_31_424971 813 12 of of IN 10_1101-2020_12_31_424971 813 13 the the DT 10_1101-2020_12_31_424971 813 14 G37C G37C NNP 10_1101-2020_12_31_424971 813 15 variant variant NN 10_1101-2020_12_31_424971 813 16 of of IN 10_1101-2020_12_31_424971 813 17 G37-state G37-state NNP 10_1101-2020_12_31_424971 813 18 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 813 19 to to IN 10_1101-2020_12_31_424971 813 20 the the DT 10_1101-2020_12_31_424971 813 21 A A NNP 10_1101-2020_12_31_424971 813 22 site site NN 10_1101-2020_12_31_424971 813 23 of of IN 10_1101-2020_12_31_424971 813 24 a a DT 10_1101-2020_12_31_424971 813 25 70S 70S NNP 10_1101-2020_12_31_424971 813 26 IC IC NNP 10_1101-2020_12_31_424971 813 27 , , , 10_1101-2020_12_31_424971 813 28 allowing allow VBG 10_1101-2020_12_31_424971 813 29 1280 1280 CD 10_1101-2020_12_31_424971 813 30 nucleotides nucleotide VBZ 10_1101-2020_12_31_424971 813 31 34 34 CD 10_1101-2020_12_31_424971 813 32 - - SYM 10_1101-2020_12_31_424971 813 33 36 36 CD 10_1101-2020_12_31_424971 813 34 to to TO 10_1101-2020_12_31_424971 813 35 pair pair VB 10_1101-2020_12_31_424971 813 36 with with IN 10_1101-2020_12_31_424971 813 37 a a DT 10_1101-2020_12_31_424971 813 38 CCC CCC NNP 10_1101-2020_12_31_424971 813 39 - - HYPH 10_1101-2020_12_31_424971 813 40 C C NNP 10_1101-2020_12_31_424971 813 41 motif motif NN 10_1101-2020_12_31_424971 813 42 at at IN 10_1101-2020_12_31_424971 813 43 the the DT 10_1101-2020_12_31_424971 813 44 A A NNP 10_1101-2020_12_31_424971 813 45 site site NN 10_1101-2020_12_31_424971 813 46 . . . 10_1101-2020_12_31_424971 814 1 b b NN 10_1101-2020_12_31_424971 814 2 fMP fMP NNP 10_1101-2020_12_31_424971 814 3 formation formation NN 10_1101-2020_12_31_424971 814 4 as as IN 10_1101-2020_12_31_424971 814 5 a a DT 10_1101-2020_12_31_424971 814 6 function function NN 10_1101-2020_12_31_424971 814 7 of of IN 10_1101-2020_12_31_424971 814 8 time time NN 10_1101-2020_12_31_424971 814 9 1281 1281 CD 10_1101-2020_12_31_424971 814 10 upon upon IN 10_1101-2020_12_31_424971 814 11 delivery delivery NN 10_1101-2020_12_31_424971 814 12 of of IN 10_1101-2020_12_31_424971 814 13 the the DT 10_1101-2020_12_31_424971 814 14 G34C G34C NNP 10_1101-2020_12_31_424971 814 15 variant variant NN 10_1101-2020_12_31_424971 814 16 of of IN 10_1101-2020_12_31_424971 814 17 G37-state G37-state NNP 10_1101-2020_12_31_424971 814 18 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 814 19 to to IN 10_1101-2020_12_31_424971 814 20 the the DT 10_1101-2020_12_31_424971 814 21 A A NNP 10_1101-2020_12_31_424971 814 22 site site NN 10_1101-2020_12_31_424971 814 23 of of IN 10_1101-2020_12_31_424971 814 24 a a DT 10_1101-2020_12_31_424971 814 25 70S 70S NNP 10_1101-2020_12_31_424971 814 26 IC IC NNP 10_1101-2020_12_31_424971 814 27 , , , 10_1101-2020_12_31_424971 814 28 allowing allow VBG 10_1101-2020_12_31_424971 814 29 1282 1282 CD 10_1101-2020_12_31_424971 814 30 nucleotides nucleotide VBZ 10_1101-2020_12_31_424971 814 31 35 35 CD 10_1101-2020_12_31_424971 814 32 - - SYM 10_1101-2020_12_31_424971 814 33 37 37 CD 10_1101-2020_12_31_424971 814 34 to to TO 10_1101-2020_12_31_424971 814 35 pair pair VB 10_1101-2020_12_31_424971 814 36 with with IN 10_1101-2020_12_31_424971 814 37 a a DT 10_1101-2020_12_31_424971 814 38 CCC CCC NNP 10_1101-2020_12_31_424971 814 39 - - HYPH 10_1101-2020_12_31_424971 814 40 C C NNP 10_1101-2020_12_31_424971 814 41 motif motif NN 10_1101-2020_12_31_424971 814 42 . . . 10_1101-2020_12_31_424971 815 1 c c LS 10_1101-2020_12_31_424971 815 2 - - HYPH 10_1101-2020_12_31_424971 815 3 f f NNP 10_1101-2020_12_31_424971 815 4 Results Results NNP 10_1101-2020_12_31_424971 815 5 of of IN 10_1101-2020_12_31_424971 815 6 fMPV fmpv NN 10_1101-2020_12_31_424971 815 7 formation formation NN 10_1101-2020_12_31_424971 815 8 assays assays RB 10_1101-2020_12_31_424971 815 9 in in IN 10_1101-2020_12_31_424971 815 10 which which WDT 10_1101-2020_12_31_424971 815 11 1283 1283 CD 10_1101-2020_12_31_424971 815 12 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 815 13 is be VBZ 10_1101-2020_12_31_424971 815 14 delivered deliver VBN 10_1101-2020_12_31_424971 815 15 to to IN 10_1101-2020_12_31_424971 815 16 an an DT 10_1101-2020_12_31_424971 815 17 A A NNP 10_1101-2020_12_31_424971 815 18 site site NN 10_1101-2020_12_31_424971 815 19 programmed program VBN 10_1101-2020_12_31_424971 815 20 with with IN 10_1101-2020_12_31_424971 815 21 a a DT 10_1101-2020_12_31_424971 815 22 quadruplet quadruplet NN 10_1101-2020_12_31_424971 815 23 codon codon NN 10_1101-2020_12_31_424971 815 24 at at IN 10_1101-2020_12_31_424971 815 25 the the DT 10_1101-2020_12_31_424971 815 26 2nd 2nd JJ 10_1101-2020_12_31_424971 815 27 position position NN 10_1101-2020_12_31_424971 815 28 and and CC 10_1101-2020_12_31_424971 815 29 1284 1284 CD 10_1101-2020_12_31_424971 815 30 sequences sequence NNS 10_1101-2020_12_31_424971 815 31 of of IN 10_1101-2020_12_31_424971 815 32 the the DT 10_1101-2020_12_31_424971 815 33 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 815 34 anticodon anticodon NN 10_1101-2020_12_31_424971 815 35 loop loop NN 10_1101-2020_12_31_424971 815 36 and/or and/or CC 10_1101-2020_12_31_424971 815 37 quadruplet quadruplet NN 10_1101-2020_12_31_424971 815 38 codon codon NNP 10_1101-2020_12_31_424971 815 39 are be VBP 10_1101-2020_12_31_424971 815 40 varied varied JJ 10_1101-2020_12_31_424971 815 41 . . . 10_1101-2020_12_31_424971 816 1 Yields yield NNS 10_1101-2020_12_31_424971 816 2 of of IN 10_1101-2020_12_31_424971 816 3 fMPV fMPV NNP 10_1101-2020_12_31_424971 816 4 1285 1285 CD 10_1101-2020_12_31_424971 816 5 formation formation NN 10_1101-2020_12_31_424971 816 6 represent represent NN 10_1101-2020_12_31_424971 816 7 +1 +1 : 10_1101-2020_12_31_424971 816 8 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 816 9 during during IN 10_1101-2020_12_31_424971 816 10 translocation translocation NN 10_1101-2020_12_31_424971 816 11 of of IN 10_1101-2020_12_31_424971 816 12 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 816 13 from from IN 10_1101-2020_12_31_424971 816 14 the the DT 10_1101-2020_12_31_424971 816 15 A A NNP 10_1101-2020_12_31_424971 816 16 site site NN 10_1101-2020_12_31_424971 816 17 to to IN 10_1101-2020_12_31_424971 816 18 the the DT 10_1101-2020_12_31_424971 816 19 P p NN 10_1101-2020_12_31_424971 816 20 site site NN 10_1101-2020_12_31_424971 816 21 . . . 10_1101-2020_12_31_424971 817 1 1286 1286 CD 10_1101-2020_12_31_424971 817 2 Possible possible JJ 10_1101-2020_12_31_424971 817 3 +1-frame +1-frame VBZ 10_1101-2020_12_31_424971 817 4 anticodon anticodon NN 10_1101-2020_12_31_424971 817 5 - - HYPH 10_1101-2020_12_31_424971 817 6 codon codon NN 10_1101-2020_12_31_424971 817 7 pairing pair VBG 10_1101-2020_12_31_424971 817 8 schemes scheme NNS 10_1101-2020_12_31_424971 817 9 of of IN 10_1101-2020_12_31_424971 817 10 SufB2 sufb2 NN 10_1101-2020_12_31_424971 817 11 during during IN 10_1101-2020_12_31_424971 817 12 translocation translocation NN 10_1101-2020_12_31_424971 817 13 : : : 10_1101-2020_12_31_424971 817 14 c c NN 10_1101-2020_12_31_424971 817 15 G37-state G37-state NNP 10_1101-2020_12_31_424971 817 16 1287 1287 CD 10_1101-2020_12_31_424971 817 17 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 817 18 capable capable JJ 10_1101-2020_12_31_424971 817 19 of of IN 10_1101-2020_12_31_424971 817 20 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 817 21 at at IN 10_1101-2020_12_31_424971 817 22 a a DT 10_1101-2020_12_31_424971 817 23 CCC CCC NNP 10_1101-2020_12_31_424971 817 24 - - HYPH 10_1101-2020_12_31_424971 817 25 C C NNP 10_1101-2020_12_31_424971 817 26 motif motif NN 10_1101-2020_12_31_424971 817 27 via via IN 10_1101-2020_12_31_424971 817 28 quadruplet quadruplet NN 10_1101-2020_12_31_424971 817 29 pairing pairing NN 10_1101-2020_12_31_424971 817 30 and/or and/or CC 10_1101-2020_12_31_424971 817 31 triplet triplet NN 10_1101-2020_12_31_424971 817 32 slippage slippage NN 10_1101-2020_12_31_424971 817 33 , , , 10_1101-2020_12_31_424971 817 34 d d NNP 10_1101-2020_12_31_424971 817 35 1288 1288 CD 10_1101-2020_12_31_424971 817 36 G37C G37C NNP 10_1101-2020_12_31_424971 817 37 variant variant NN 10_1101-2020_12_31_424971 817 38 of of IN 10_1101-2020_12_31_424971 817 39 G37-state G37-state NNP 10_1101-2020_12_31_424971 817 40 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 817 41 capable capable JJ 10_1101-2020_12_31_424971 817 42 of of IN 10_1101-2020_12_31_424971 817 43 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 817 44 at at IN 10_1101-2020_12_31_424971 817 45 a a DT 10_1101-2020_12_31_424971 817 46 GCC GCC NNP 10_1101-2020_12_31_424971 817 47 - - HYPH 10_1101-2020_12_31_424971 817 48 C c NN 10_1101-2020_12_31_424971 817 49 motif motif NN 10_1101-2020_12_31_424971 817 50 via via IN 10_1101-2020_12_31_424971 817 51 quadruplet quadruplet NN 10_1101-2020_12_31_424971 817 52 pairing pair VBG 10_1101-2020_12_31_424971 817 53 1289 1289 CD 10_1101-2020_12_31_424971 817 54 and/or and/or CC 10_1101-2020_12_31_424971 817 55 triplet triplet NN 10_1101-2020_12_31_424971 817 56 slippage slippage NN 10_1101-2020_12_31_424971 817 57 , , , 10_1101-2020_12_31_424971 817 58 e e LS 10_1101-2020_12_31_424971 817 59 m1G37-state m1G37-state NNP 10_1101-2020_12_31_424971 817 60 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 817 61 capable capable JJ 10_1101-2020_12_31_424971 817 62 of of IN 10_1101-2020_12_31_424971 817 63 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 817 64 at at IN 10_1101-2020_12_31_424971 817 65 a a DT 10_1101-2020_12_31_424971 817 66 CCC CCC NNP 10_1101-2020_12_31_424971 817 67 - - HYPH 10_1101-2020_12_31_424971 817 68 C C NNP 10_1101-2020_12_31_424971 817 69 motif motif NN 10_1101-2020_12_31_424971 817 70 via via IN 10_1101-2020_12_31_424971 817 71 only only RB 10_1101-2020_12_31_424971 817 72 1290 1290 CD 10_1101-2020_12_31_424971 817 73 triplet triplet NN 10_1101-2020_12_31_424971 817 74 slippage slippage NN 10_1101-2020_12_31_424971 817 75 , , , 10_1101-2020_12_31_424971 817 76 and and CC 10_1101-2020_12_31_424971 817 77 f f NNP 10_1101-2020_12_31_424971 817 78 G37C G37C NNP 10_1101-2020_12_31_424971 817 79 variant variant NN 10_1101-2020_12_31_424971 817 80 of of IN 10_1101-2020_12_31_424971 817 81 G37-state G37-state NNP 10_1101-2020_12_31_424971 817 82 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 817 83 capable capable JJ 10_1101-2020_12_31_424971 817 84 of of IN 10_1101-2020_12_31_424971 817 85 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 817 86 at at IN 10_1101-2020_12_31_424971 817 87 a a DT 10_1101-2020_12_31_424971 817 88 CCC CCC NNP 10_1101-2020_12_31_424971 817 89 - - HYPH 10_1101-2020_12_31_424971 817 90 C C NNP 10_1101-2020_12_31_424971 817 91 motif motif NN 10_1101-2020_12_31_424971 817 92 1291 1291 CD 10_1101-2020_12_31_424971 817 93 via via IN 10_1101-2020_12_31_424971 817 94 only only JJ 10_1101-2020_12_31_424971 817 95 triplet triplet NN 10_1101-2020_12_31_424971 817 96 slippage slippage NN 10_1101-2020_12_31_424971 817 97 . . . 10_1101-2020_12_31_424971 818 1 In in IN 10_1101-2020_12_31_424971 818 2 panels panel NNS 10_1101-2020_12_31_424971 818 3 a a XX 10_1101-2020_12_31_424971 818 4 , , , 10_1101-2020_12_31_424971 818 5 b b NN 10_1101-2020_12_31_424971 818 6 , , , 10_1101-2020_12_31_424971 818 7 the the DT 10_1101-2020_12_31_424971 818 8 bars bar NNS 10_1101-2020_12_31_424971 818 9 in in IN 10_1101-2020_12_31_424971 818 10 the the DT 10_1101-2020_12_31_424971 818 11 graphs graph NNS 10_1101-2020_12_31_424971 818 12 are be VBP 10_1101-2020_12_31_424971 818 13 SD sd NN 10_1101-2020_12_31_424971 818 14 of of IN 10_1101-2020_12_31_424971 818 15 three three CD 10_1101-2020_12_31_424971 818 16 ( ( -LRB- 10_1101-2020_12_31_424971 818 17 n n NN 10_1101-2020_12_31_424971 818 18 = = SYM 10_1101-2020_12_31_424971 818 19 3 3 LS 10_1101-2020_12_31_424971 818 20 ) ) -RRB- 10_1101-2020_12_31_424971 818 21 independent independent JJ 10_1101-2020_12_31_424971 818 22 1292 1292 CD 10_1101-2020_12_31_424971 818 23 experiments experiment NNS 10_1101-2020_12_31_424971 818 24 , , , 10_1101-2020_12_31_424971 818 25 and and CC 10_1101-2020_12_31_424971 818 26 the the DT 10_1101-2020_12_31_424971 818 27 data datum NNS 10_1101-2020_12_31_424971 818 28 are be VBP 10_1101-2020_12_31_424971 818 29 presented present VBN 10_1101-2020_12_31_424971 818 30 as as IN 10_1101-2020_12_31_424971 818 31 mean mean JJ 10_1101-2020_12_31_424971 818 32 values value NNS 10_1101-2020_12_31_424971 818 33 ± ± CD 10_1101-2020_12_31_424971 818 34 SD sd NN 10_1101-2020_12_31_424971 818 35 . . . 10_1101-2020_12_31_424971 819 1 ∆t ∆t LS 10_1101-2020_12_31_424971 819 2 : : : 10_1101-2020_12_31_424971 819 3 a a DT 10_1101-2020_12_31_424971 819 4 time time NN 10_1101-2020_12_31_424971 819 5 interval interval NN 10_1101-2020_12_31_424971 819 6 . . . 10_1101-2020_12_31_424971 820 1 1293 1293 CD 10_1101-2020_12_31_424971 820 2 .CC .CC : 10_1101-2020_12_31_424971 820 3 - - : 10_1101-2020_12_31_424971 820 4 BY by IN 10_1101-2020_12_31_424971 820 5 - - HYPH 10_1101-2020_12_31_424971 820 6 NC NC NNP 10_1101-2020_12_31_424971 820 7 - - HYPH 10_1101-2020_12_31_424971 820 8 ND ND NNP 10_1101-2020_12_31_424971 820 9 4.0 4.0 CD 10_1101-2020_12_31_424971 820 10 International International NNP 10_1101-2020_12_31_424971 820 11 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 820 12 under under IN 10_1101-2020_12_31_424971 820 13 a a DT 10_1101-2020_12_31_424971 820 14 ( ( -LRB- 10_1101-2020_12_31_424971 820 15 which which WDT 10_1101-2020_12_31_424971 820 16 was be VBD 10_1101-2020_12_31_424971 820 17 not not RB 10_1101-2020_12_31_424971 820 18 certified certify VBN 10_1101-2020_12_31_424971 820 19 by by IN 10_1101-2020_12_31_424971 820 20 peer peer NN 10_1101-2020_12_31_424971 820 21 review review NN 10_1101-2020_12_31_424971 820 22 ) ) -RRB- 10_1101-2020_12_31_424971 820 23 is be VBZ 10_1101-2020_12_31_424971 820 24 the the DT 10_1101-2020_12_31_424971 820 25 author author NN 10_1101-2020_12_31_424971 820 26 / / SYM 10_1101-2020_12_31_424971 820 27 funder funder NN 10_1101-2020_12_31_424971 820 28 , , , 10_1101-2020_12_31_424971 820 29 who who WP 10_1101-2020_12_31_424971 820 30 has have VBZ 10_1101-2020_12_31_424971 820 31 granted grant VBN 10_1101-2020_12_31_424971 820 32 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 820 33 a a DT 10_1101-2020_12_31_424971 820 34 license license NN 10_1101-2020_12_31_424971 820 35 to to TO 10_1101-2020_12_31_424971 820 36 display display VB 10_1101-2020_12_31_424971 820 37 the the DT 10_1101-2020_12_31_424971 820 38 preprint preprint NN 10_1101-2020_12_31_424971 820 39 in in IN 10_1101-2020_12_31_424971 820 40 perpetuity perpetuity NN 10_1101-2020_12_31_424971 820 41 . . . 10_1101-2020_12_31_424971 821 1 It -PRON- PRP 10_1101-2020_12_31_424971 821 2 is be VBZ 10_1101-2020_12_31_424971 821 3 made make VBN 10_1101-2020_12_31_424971 821 4 The the DT 10_1101-2020_12_31_424971 821 5 copyright copyright NN 10_1101-2020_12_31_424971 821 6 holder holder NN 10_1101-2020_12_31_424971 821 7 for for IN 10_1101-2020_12_31_424971 821 8 this this DT 10_1101-2020_12_31_424971 821 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 821 10 version version NN 10_1101-2020_12_31_424971 821 11 posted post VBD 10_1101-2020_12_31_424971 821 12 January January NNP 10_1101-2020_12_31_424971 821 13 2 2 CD 10_1101-2020_12_31_424971 821 14 , , , 10_1101-2020_12_31_424971 821 15 2021 2021 CD 10_1101-2020_12_31_424971 821 16 . . . 10_1101-2020_12_31_424971 821 17 ; ; : 10_1101-2020_12_31_424971 821 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 821 19 : : : 10_1101-2020_12_31_424971 821 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 821 21 preprint preprint NN 10_1101-2020_12_31_424971 821 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 821 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 821 24 50 50 CD 10_1101-2020_12_31_424971 821 25 1294 1294 CD 10_1101-2020_12_31_424971 821 26 Figure Figure NNP 10_1101-2020_12_31_424971 821 27 5 5 CD 10_1101-2020_12_31_424971 821 28 . . . 10_1101-2020_12_31_424971 822 1 SufB2 sufb2 NN 10_1101-2020_12_31_424971 822 2 shifts shift NNS 10_1101-2020_12_31_424971 822 3 to to IN 10_1101-2020_12_31_424971 822 4 the the DT 10_1101-2020_12_31_424971 822 5 +1-frame +1-frame NNS 10_1101-2020_12_31_424971 822 6 during during IN 10_1101-2020_12_31_424971 822 7 translocation translocation NN 10_1101-2020_12_31_424971 822 8 . . . 10_1101-2020_12_31_424971 823 1 a a DT 10_1101-2020_12_31_424971 823 2 Relative relative JJ 10_1101-2020_12_31_424971 823 3 fMPV fmpv NN 10_1101-2020_12_31_424971 823 4 and and CC 10_1101-2020_12_31_424971 823 5 fMPR fmpr JJ 10_1101-2020_12_31_424971 823 6 1295 1295 CD 10_1101-2020_12_31_424971 823 7 formation formation NN 10_1101-2020_12_31_424971 823 8 as as IN 10_1101-2020_12_31_424971 823 9 a a DT 10_1101-2020_12_31_424971 823 10 function function NN 10_1101-2020_12_31_424971 823 11 of of IN 10_1101-2020_12_31_424971 823 12 time time NN 10_1101-2020_12_31_424971 823 13 upon upon IN 10_1101-2020_12_31_424971 823 14 rapid rapid JJ 10_1101-2020_12_31_424971 823 15 delivery delivery NN 10_1101-2020_12_31_424971 823 16 of of IN 10_1101-2020_12_31_424971 823 17 EF EF NNP 10_1101-2020_12_31_424971 823 18 - - HYPH 10_1101-2020_12_31_424971 823 19 G G NNP 10_1101-2020_12_31_424971 823 20 and and CC 10_1101-2020_12_31_424971 823 21 an an DT 10_1101-2020_12_31_424971 823 22 equimolar equimolar JJ 10_1101-2020_12_31_424971 823 23 mixture mixture NN 10_1101-2020_12_31_424971 823 24 of of IN 10_1101-2020_12_31_424971 823 25 G37-state G37-state NNP 10_1101-2020_12_31_424971 823 26 1296 1296 CD 10_1101-2020_12_31_424971 823 27 SufB2- sufb2- CD 10_1101-2020_12_31_424971 823 28 , , , 10_1101-2020_12_31_424971 823 29 tRNAVal- tRNAVal- NNP 10_1101-2020_12_31_424971 823 30 , , , 10_1101-2020_12_31_424971 823 31 and and CC 10_1101-2020_12_31_424971 823 32 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 823 33 - - HYPH 10_1101-2020_12_31_424971 823 34 TCs TCs NNP 10_1101-2020_12_31_424971 823 35 to to IN 10_1101-2020_12_31_424971 823 36 70S 70S NNP 10_1101-2020_12_31_424971 823 37 ICs ic NNS 10_1101-2020_12_31_424971 823 38 carrying carry VBG 10_1101-2020_12_31_424971 823 39 a a DT 10_1101-2020_12_31_424971 823 40 CCC CCC NNP 10_1101-2020_12_31_424971 823 41 - - HYPH 10_1101-2020_12_31_424971 823 42 C C NNP 10_1101-2020_12_31_424971 823 43 motif motif NN 10_1101-2020_12_31_424971 823 44 in in IN 10_1101-2020_12_31_424971 823 45 the the DT 10_1101-2020_12_31_424971 823 46 A A NNP 10_1101-2020_12_31_424971 823 47 site site NN 10_1101-2020_12_31_424971 823 48 . . . 10_1101-2020_12_31_424971 824 1 b b LS 10_1101-2020_12_31_424971 824 2 Relative Relative NNP 10_1101-2020_12_31_424971 824 3 1297 1297 CD 10_1101-2020_12_31_424971 824 4 fMPV fmpv NN 10_1101-2020_12_31_424971 824 5 and and CC 10_1101-2020_12_31_424971 824 6 fMPR fmpr JJ 10_1101-2020_12_31_424971 824 7 formation formation NN 10_1101-2020_12_31_424971 824 8 as as IN 10_1101-2020_12_31_424971 824 9 a a DT 10_1101-2020_12_31_424971 824 10 function function NN 10_1101-2020_12_31_424971 824 11 of of IN 10_1101-2020_12_31_424971 824 12 time time NN 10_1101-2020_12_31_424971 824 13 when when WRB 10_1101-2020_12_31_424971 824 14 a a DT 10_1101-2020_12_31_424971 824 15 defined define VBN 10_1101-2020_12_31_424971 824 16 time time NN 10_1101-2020_12_31_424971 824 17 interval interval NN 10_1101-2020_12_31_424971 824 18 is be VBZ 10_1101-2020_12_31_424971 824 19 introduced introduce VBN 10_1101-2020_12_31_424971 824 20 between between IN 10_1101-2020_12_31_424971 824 21 1298 1298 CD 10_1101-2020_12_31_424971 824 22 delivery delivery NN 10_1101-2020_12_31_424971 824 23 of of IN 10_1101-2020_12_31_424971 824 24 G37-state G37-state NNP 10_1101-2020_12_31_424971 824 25 SufB2-TC SufB2-TC NNP 10_1101-2020_12_31_424971 824 26 and and CC 10_1101-2020_12_31_424971 824 27 EF EF NNP 10_1101-2020_12_31_424971 824 28 - - HYPH 10_1101-2020_12_31_424971 824 29 G G NNP 10_1101-2020_12_31_424971 824 30 and and CC 10_1101-2020_12_31_424971 824 31 delivery delivery NN 10_1101-2020_12_31_424971 824 32 of of IN 10_1101-2020_12_31_424971 824 33 an an DT 10_1101-2020_12_31_424971 824 34 equimolar equimolar JJ 10_1101-2020_12_31_424971 824 35 mixture mixture NN 10_1101-2020_12_31_424971 824 36 of of IN 10_1101-2020_12_31_424971 824 37 tRNAArg- trnaarg- NN 10_1101-2020_12_31_424971 824 38 and and CC 10_1101-2020_12_31_424971 824 39 1299 1299 CD 10_1101-2020_12_31_424971 824 40 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 824 41 - - HYPH 10_1101-2020_12_31_424971 824 42 TCs TCs NNP 10_1101-2020_12_31_424971 824 43 . . . 10_1101-2020_12_31_424971 825 1 c c LS 10_1101-2020_12_31_424971 825 2 Relative relative JJ 10_1101-2020_12_31_424971 825 3 fMPV fmpv NN 10_1101-2020_12_31_424971 825 4 and and CC 10_1101-2020_12_31_424971 825 5 fMPR fmpr JJ 10_1101-2020_12_31_424971 825 6 formation formation NN 10_1101-2020_12_31_424971 825 7 after after IN 10_1101-2020_12_31_424971 825 8 reacting react VBG 10_1101-2020_12_31_424971 825 9 fMP fmp XX 10_1101-2020_12_31_424971 825 10 - - HYPH 10_1101-2020_12_31_424971 825 11 POST POST AFX 10_1101-2020_12_31_424971 825 12 complexes complex NNS 10_1101-2020_12_31_424971 825 13 with with IN 10_1101-2020_12_31_424971 825 14 a a DT 10_1101-2020_12_31_424971 825 15 1300 1300 CD 10_1101-2020_12_31_424971 825 16 mixture mixture NN 10_1101-2020_12_31_424971 825 17 of of IN 10_1101-2020_12_31_424971 825 18 tRNAVal- tRNAVal- NNP 10_1101-2020_12_31_424971 825 19 and and CC 10_1101-2020_12_31_424971 825 20 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 825 21 - - HYPH 10_1101-2020_12_31_424971 825 22 TCs TCs NNP 10_1101-2020_12_31_424971 825 23 based base VBN 10_1101-2020_12_31_424971 825 24 on on IN 10_1101-2020_12_31_424971 825 25 the the DT 10_1101-2020_12_31_424971 825 26 time time NN 10_1101-2020_12_31_424971 825 27 courses course NNS 10_1101-2020_12_31_424971 825 28 in in IN 10_1101-2020_12_31_424971 825 29 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 825 30 Figures Figures NNP 10_1101-2020_12_31_424971 825 31 2d 2d NNP 10_1101-2020_12_31_424971 825 32 - - HYPH 10_1101-2020_12_31_424971 825 33 f f NN 10_1101-2020_12_31_424971 825 34 . . . 10_1101-2020_12_31_424971 825 35 1301 1301 CD 10_1101-2020_12_31_424971 825 36 d d NNP 10_1101-2020_12_31_424971 825 37 fMPV fMPV NNP 10_1101-2020_12_31_424971 825 38 formation formation NN 10_1101-2020_12_31_424971 825 39 as as IN 10_1101-2020_12_31_424971 825 40 a a DT 10_1101-2020_12_31_424971 825 41 function function NN 10_1101-2020_12_31_424971 825 42 of of IN 10_1101-2020_12_31_424971 825 43 time time NN 10_1101-2020_12_31_424971 825 44 upon upon IN 10_1101-2020_12_31_424971 825 45 rapid rapid JJ 10_1101-2020_12_31_424971 825 46 delivery delivery NN 10_1101-2020_12_31_424971 825 47 of of IN 10_1101-2020_12_31_424971 825 48 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 825 49 - - HYPH 10_1101-2020_12_31_424971 825 50 TC TC NNP 10_1101-2020_12_31_424971 825 51 to to IN 10_1101-2020_12_31_424971 825 52 an an DT 10_1101-2020_12_31_424971 825 53 fMP fmp CD 10_1101-2020_12_31_424971 825 54 - - HYPH 10_1101-2020_12_31_424971 825 55 POST POST NNP 10_1101-2020_12_31_424971 825 56 1302 1302 CD 10_1101-2020_12_31_424971 825 57 complex complex NN 10_1101-2020_12_31_424971 825 58 carrying carry VBG 10_1101-2020_12_31_424971 825 59 a a DT 10_1101-2020_12_31_424971 825 60 CCC CCC NNP 10_1101-2020_12_31_424971 825 61 - - HYPH 10_1101-2020_12_31_424971 825 62 N n CD 10_1101-2020_12_31_424971 825 63 motif motif NN 10_1101-2020_12_31_424971 825 64 in in IN 10_1101-2020_12_31_424971 825 65 the the DT 10_1101-2020_12_31_424971 825 66 A A NNP 10_1101-2020_12_31_424971 825 67 site site NN 10_1101-2020_12_31_424971 825 68 . . . 10_1101-2020_12_31_424971 826 1 e e LS 10_1101-2020_12_31_424971 826 2 Relative relative JJ 10_1101-2020_12_31_424971 826 3 fMPV fmpv NN 10_1101-2020_12_31_424971 826 4 and and CC 10_1101-2020_12_31_424971 826 5 fMPS fmps NN 10_1101-2020_12_31_424971 826 6 formation formation NN 10_1101-2020_12_31_424971 826 7 as as IN 10_1101-2020_12_31_424971 826 8 a a DT 10_1101-2020_12_31_424971 826 9 function function NN 10_1101-2020_12_31_424971 826 10 1303 1303 CD 10_1101-2020_12_31_424971 826 11 of of IN 10_1101-2020_12_31_424971 826 12 time time NN 10_1101-2020_12_31_424971 826 13 upon upon IN 10_1101-2020_12_31_424971 826 14 rapid rapid JJ 10_1101-2020_12_31_424971 826 15 delivery delivery NN 10_1101-2020_12_31_424971 826 16 of of IN 10_1101-2020_12_31_424971 826 17 an an DT 10_1101-2020_12_31_424971 826 18 equimolar equimolar JJ 10_1101-2020_12_31_424971 826 19 mixture mixture NN 10_1101-2020_12_31_424971 826 20 of of IN 10_1101-2020_12_31_424971 826 21 tRNAVal- tRNAVal- NNP 10_1101-2020_12_31_424971 826 22 and and CC 10_1101-2020_12_31_424971 826 23 tRNASer tRNASer NNP 10_1101-2020_12_31_424971 826 24 - - HYPH 10_1101-2020_12_31_424971 826 25 TCs tc NNS 10_1101-2020_12_31_424971 826 26 to to IN 10_1101-2020_12_31_424971 826 27 an an DT 10_1101-2020_12_31_424971 826 28 fMP fMP NNP 10_1101-2020_12_31_424971 826 29 - - HYPH 10_1101-2020_12_31_424971 826 30 POST POST NNP 10_1101-2020_12_31_424971 826 31 1304 1304 CD 10_1101-2020_12_31_424971 826 32 complex complex NN 10_1101-2020_12_31_424971 826 33 carrying carry VBG 10_1101-2020_12_31_424971 826 34 a a DT 10_1101-2020_12_31_424971 826 35 CCC CCC NNP 10_1101-2020_12_31_424971 826 36 - - HYPH 10_1101-2020_12_31_424971 826 37 A a DT 10_1101-2020_12_31_424971 826 38 motif motif NN 10_1101-2020_12_31_424971 826 39 in in IN 10_1101-2020_12_31_424971 826 40 the the DT 10_1101-2020_12_31_424971 826 41 A A NNP 10_1101-2020_12_31_424971 826 42 site site NN 10_1101-2020_12_31_424971 826 43 . . . 10_1101-2020_12_31_424971 827 1 In in IN 10_1101-2020_12_31_424971 827 2 panels panel NNS 10_1101-2020_12_31_424971 827 3 a a DT 10_1101-2020_12_31_424971 827 4 - - HYPH 10_1101-2020_12_31_424971 827 5 e e NN 10_1101-2020_12_31_424971 827 6 , , , 10_1101-2020_12_31_424971 827 7 the the DT 10_1101-2020_12_31_424971 827 8 bars bar NNS 10_1101-2020_12_31_424971 827 9 are be VBP 10_1101-2020_12_31_424971 827 10 SD sd NN 10_1101-2020_12_31_424971 827 11 of of IN 10_1101-2020_12_31_424971 827 12 three three CD 10_1101-2020_12_31_424971 827 13 ( ( -LRB- 10_1101-2020_12_31_424971 827 14 n n NN 10_1101-2020_12_31_424971 827 15 = = SYM 10_1101-2020_12_31_424971 827 16 3 3 CD 10_1101-2020_12_31_424971 827 17 ) ) -RRB- 10_1101-2020_12_31_424971 827 18 1305 1305 CD 10_1101-2020_12_31_424971 827 19 independent independent JJ 10_1101-2020_12_31_424971 827 20 experiments experiment NNS 10_1101-2020_12_31_424971 827 21 and and CC 10_1101-2020_12_31_424971 827 22 the the DT 10_1101-2020_12_31_424971 827 23 data datum NNS 10_1101-2020_12_31_424971 827 24 are be VBP 10_1101-2020_12_31_424971 827 25 presented present VBN 10_1101-2020_12_31_424971 827 26 as as IN 10_1101-2020_12_31_424971 827 27 mean mean JJ 10_1101-2020_12_31_424971 827 28 values value NNS 10_1101-2020_12_31_424971 827 29 ± ± CD 10_1101-2020_12_31_424971 827 30 SD sd NN 10_1101-2020_12_31_424971 827 31 . . . 10_1101-2020_12_31_424971 828 1 Arg arg NN 10_1101-2020_12_31_424971 828 2 : : : 10_1101-2020_12_31_424971 828 3 arginyl arginyl NN 10_1101-2020_12_31_424971 828 4 - - : 10_1101-2020_12_31_424971 828 5 tRNAArg tRNAArg NNP 10_1101-2020_12_31_424971 828 6 ; ; : 10_1101-2020_12_31_424971 828 7 1306 1306 CD 10_1101-2020_12_31_424971 828 8 Val Val NNP 10_1101-2020_12_31_424971 828 9 : : : 10_1101-2020_12_31_424971 828 10 valyl valyl NN 10_1101-2020_12_31_424971 828 11 - - HYPH 10_1101-2020_12_31_424971 828 12 tRNAVal tRNAVal NNP 10_1101-2020_12_31_424971 828 13 . . . 10_1101-2020_12_31_424971 829 1 1307 1307 CD 10_1101-2020_12_31_424971 829 2 1308 1308 CD 10_1101-2020_12_31_424971 829 3 Figure Figure NNP 10_1101-2020_12_31_424971 829 4 6 6 CD 10_1101-2020_12_31_424971 829 5 . . . 10_1101-2020_12_31_424971 830 1 SufB2 sufb2 NN 10_1101-2020_12_31_424971 830 2 interferes interfere VBZ 10_1101-2020_12_31_424971 830 3 with with IN 10_1101-2020_12_31_424971 830 4 elongation elongation NN 10_1101-2020_12_31_424971 830 5 complex complex JJ 10_1101-2020_12_31_424971 830 6 dynamics dynamic NNS 10_1101-2020_12_31_424971 830 7 during during IN 10_1101-2020_12_31_424971 830 8 late late JJ 10_1101-2020_12_31_424971 830 9 steps step NNS 10_1101-2020_12_31_424971 830 10 of of IN 10_1101-2020_12_31_424971 830 11 1309 1309 CD 10_1101-2020_12_31_424971 830 12 translocation translocation NN 10_1101-2020_12_31_424971 830 13 . . . 10_1101-2020_12_31_424971 831 1 a a LS 10_1101-2020_12_31_424971 831 2 - - HYPH 10_1101-2020_12_31_424971 831 3 c c NNP 10_1101-2020_12_31_424971 831 4 Cartoon Cartoon NNP 10_1101-2020_12_31_424971 831 5 representation representation NN 10_1101-2020_12_31_424971 831 6 of of IN 10_1101-2020_12_31_424971 831 7 elongation elongation NN 10_1101-2020_12_31_424971 831 8 as as IN 10_1101-2020_12_31_424971 831 9 a a DT 10_1101-2020_12_31_424971 831 10 G37-state G37-state NNP 10_1101-2020_12_31_424971 831 11 SufB2- sufb2- NN 10_1101-2020_12_31_424971 831 12 or or CC 10_1101-2020_12_31_424971 831 13 ProL ProL NNP 10_1101-2020_12_31_424971 831 14 - - HYPH 10_1101-2020_12_31_424971 831 15 TC TC NNP 10_1101-2020_12_31_424971 831 16 is be VBZ 10_1101-2020_12_31_424971 831 17 1310 1310 CD 10_1101-2020_12_31_424971 831 18 delivered deliver VBN 10_1101-2020_12_31_424971 831 19 to to IN 10_1101-2020_12_31_424971 831 20 the the DT 10_1101-2020_12_31_424971 831 21 A A NNP 10_1101-2020_12_31_424971 831 22 site site NN 10_1101-2020_12_31_424971 831 23 of of IN 10_1101-2020_12_31_424971 831 24 a a DT 10_1101-2020_12_31_424971 831 25 bL9(Cy3)- bl9(cy3)- NN 10_1101-2020_12_31_424971 831 26 and and CC 10_1101-2020_12_31_424971 831 27 uL1(Cy5)-labeled ul1(cy5)-labeled CD 10_1101-2020_12_31_424971 831 28 70S 70S NNP 10_1101-2020_12_31_424971 831 29 IC IC NNP 10_1101-2020_12_31_424971 831 30 ; ; : 10_1101-2020_12_31_424971 831 31 a a NN 10_1101-2020_12_31_424971 831 32 in in IN 10_1101-2020_12_31_424971 831 33 the the DT 10_1101-2020_12_31_424971 831 34 absence absence NN 10_1101-2020_12_31_424971 831 35 , , , 10_1101-2020_12_31_424971 831 36 or or CC 10_1101-2020_12_31_424971 831 37 b b NN 10_1101-2020_12_31_424971 831 38 in in IN 10_1101-2020_12_31_424971 831 39 the the DT 10_1101-2020_12_31_424971 831 40 1311 1311 CD 10_1101-2020_12_31_424971 831 41 presence presence NN 10_1101-2020_12_31_424971 831 42 of of IN 10_1101-2020_12_31_424971 831 43 EF EF NNP 10_1101-2020_12_31_424971 831 44 - - HYPH 10_1101-2020_12_31_424971 831 45 G G NNP 10_1101-2020_12_31_424971 831 46 , , , 10_1101-2020_12_31_424971 831 47 or or CC 10_1101-2020_12_31_424971 831 48 c c NN 10_1101-2020_12_31_424971 831 49 upon upon IN 10_1101-2020_12_31_424971 831 50 using use VBG 10_1101-2020_12_31_424971 831 51 puromycin puromycin NNS 10_1101-2020_12_31_424971 831 52 ( ( -LRB- 10_1101-2020_12_31_424971 831 53 Pmn pmn NN 10_1101-2020_12_31_424971 831 54 ) ) -RRB- 10_1101-2020_12_31_424971 831 55 to to TO 10_1101-2020_12_31_424971 831 56 deacylate deacylate VB 10_1101-2020_12_31_424971 831 57 the the DT 10_1101-2020_12_31_424971 831 58 P p NN 10_1101-2020_12_31_424971 831 59 site site NN 10_1101-2020_12_31_424971 831 60 - - HYPH 10_1101-2020_12_31_424971 831 61 bound bind VBN 10_1101-2020_12_31_424971 831 62 G37-state G37-state NNP 10_1101-2020_12_31_424971 831 63 1312 1312 CD 10_1101-2020_12_31_424971 831 64 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 831 65 or or CC 10_1101-2020_12_31_424971 831 66 ProL ProL NNP 10_1101-2020_12_31_424971 831 67 and and CC 10_1101-2020_12_31_424971 831 68 generate generate VB 10_1101-2020_12_31_424971 831 69 the the DT 10_1101-2020_12_31_424971 831 70 corresponding correspond VBG 10_1101-2020_12_31_424971 831 71 PRE PRE NNP 10_1101-2020_12_31_424971 831 72 – – : 10_1101-2020_12_31_424971 831 73 A a DT 10_1101-2020_12_31_424971 831 74 complex complex NN 10_1101-2020_12_31_424971 831 75 . . . 10_1101-2020_12_31_424971 832 1 The the DT 10_1101-2020_12_31_424971 832 2 30S 30S NNP 10_1101-2020_12_31_424971 832 3 and and CC 10_1101-2020_12_31_424971 832 4 50S 50S NNP 10_1101-2020_12_31_424971 832 5 subunits subunit NNS 10_1101-2020_12_31_424971 832 6 are be VBP 10_1101-2020_12_31_424971 832 7 1313 1313 CD 10_1101-2020_12_31_424971 832 8 tan tan NN 10_1101-2020_12_31_424971 832 9 and and CC 10_1101-2020_12_31_424971 832 10 light light NN 10_1101-2020_12_31_424971 832 11 blue blue NN 10_1101-2020_12_31_424971 832 12 , , , 10_1101-2020_12_31_424971 832 13 respectively respectively RB 10_1101-2020_12_31_424971 832 14 ; ; : 10_1101-2020_12_31_424971 832 15 the the DT 10_1101-2020_12_31_424971 832 16 L1 L1 NNP 10_1101-2020_12_31_424971 832 17 stalk stalk NN 10_1101-2020_12_31_424971 832 18 is be VBZ 10_1101-2020_12_31_424971 832 19 dark dark JJ 10_1101-2020_12_31_424971 832 20 blue blue JJ 10_1101-2020_12_31_424971 832 21 ; ; : 10_1101-2020_12_31_424971 832 22 Cy3 cy3 NN 10_1101-2020_12_31_424971 832 23 and and CC 10_1101-2020_12_31_424971 832 24 Cy5 cy5 NN 10_1101-2020_12_31_424971 832 25 are be VBP 10_1101-2020_12_31_424971 832 26 bright bright JJ 10_1101-2020_12_31_424971 832 27 green green JJ 10_1101-2020_12_31_424971 832 28 and and CC 10_1101-2020_12_31_424971 832 29 red red JJ 10_1101-2020_12_31_424971 832 30 1314 1314 CD 10_1101-2020_12_31_424971 832 31 spheres sphere NNS 10_1101-2020_12_31_424971 832 32 , , , 10_1101-2020_12_31_424971 832 33 respectively respectively RB 10_1101-2020_12_31_424971 832 34 ; ; : 10_1101-2020_12_31_424971 832 35 EF EF NNP 10_1101-2020_12_31_424971 832 36 - - HYPH 10_1101-2020_12_31_424971 832 37 Tu Tu NNP 10_1101-2020_12_31_424971 832 38 is be VBZ 10_1101-2020_12_31_424971 832 39 pink pink JJ 10_1101-2020_12_31_424971 832 40 ; ; : 10_1101-2020_12_31_424971 832 41 EF EF NNP 10_1101-2020_12_31_424971 832 42 - - HYPH 10_1101-2020_12_31_424971 832 43 G G NNP 10_1101-2020_12_31_424971 832 44 is be VBZ 10_1101-2020_12_31_424971 832 45 purple purple JJ 10_1101-2020_12_31_424971 832 46 ; ; : 10_1101-2020_12_31_424971 832 47 fMet fMet NNP 10_1101-2020_12_31_424971 832 48 - - HYPH 10_1101-2020_12_31_424971 832 49 tRNAfMet tRNAfMet NNP 10_1101-2020_12_31_424971 832 50 is be VBZ 10_1101-2020_12_31_424971 832 51 dark dark JJ 10_1101-2020_12_31_424971 832 52 green green NN 10_1101-2020_12_31_424971 832 53 ; ; : 10_1101-2020_12_31_424971 832 54 and and CC 10_1101-2020_12_31_424971 832 55 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 832 56 or or CC 10_1101-2020_12_31_424971 832 57 1315 1315 CD 10_1101-2020_12_31_424971 832 58 ProL ProL NNP 10_1101-2020_12_31_424971 832 59 is be VBZ 10_1101-2020_12_31_424971 832 60 dark dark JJ 10_1101-2020_12_31_424971 832 61 red red NN 10_1101-2020_12_31_424971 832 62 . . . 10_1101-2020_12_31_424971 833 1 d d NNP 10_1101-2020_12_31_424971 833 2 , , , 10_1101-2020_12_31_424971 833 3 e e NNP 10_1101-2020_12_31_424971 833 4 Hypothetical Hypothetical NNP 10_1101-2020_12_31_424971 833 5 ( ( -LRB- 10_1101-2020_12_31_424971 833 6 top top NNP 10_1101-2020_12_31_424971 833 7 ) ) -RRB- 10_1101-2020_12_31_424971 833 8 and and CC 10_1101-2020_12_31_424971 833 9 representative representative NN 10_1101-2020_12_31_424971 833 10 experimentally experimentally RB 10_1101-2020_12_31_424971 833 11 observed observe VBN 10_1101-2020_12_31_424971 833 12 ( ( -LRB- 10_1101-2020_12_31_424971 833 13 bottom bottom NN 10_1101-2020_12_31_424971 833 14 ) ) -RRB- 10_1101-2020_12_31_424971 833 15 1316 1316 CD 10_1101-2020_12_31_424971 833 16 EFRET EFRET NNP 10_1101-2020_12_31_424971 833 17 vs. vs. IN 10_1101-2020_12_31_424971 833 18 time time NN 10_1101-2020_12_31_424971 833 19 trajectories trajectory NNS 10_1101-2020_12_31_424971 833 20 recorded record VBN 10_1101-2020_12_31_424971 833 21 as as IN 10_1101-2020_12_31_424971 833 22 ProL ProL NNP 10_1101-2020_12_31_424971 833 23 - - HYPH 10_1101-2020_12_31_424971 833 24 TC TC NNP 10_1101-2020_12_31_424971 833 25 is be VBZ 10_1101-2020_12_31_424971 833 26 delivered deliver VBN 10_1101-2020_12_31_424971 833 27 to to IN 10_1101-2020_12_31_424971 833 28 a a DT 10_1101-2020_12_31_424971 833 29 70S 70S NNP 10_1101-2020_12_31_424971 833 30 IC IC NNP 10_1101-2020_12_31_424971 833 31 , , , 10_1101-2020_12_31_424971 833 32 d d NN 10_1101-2020_12_31_424971 833 33 in in IN 10_1101-2020_12_31_424971 833 34 the the DT 10_1101-2020_12_31_424971 833 35 absence absence NN 10_1101-2020_12_31_424971 833 36 and and CC 10_1101-2020_12_31_424971 833 37 e e NNP 10_1101-2020_12_31_424971 833 38 1317 1317 CD 10_1101-2020_12_31_424971 833 39 in in IN 10_1101-2020_12_31_424971 833 40 the the DT 10_1101-2020_12_31_424971 833 41 presence presence NN 10_1101-2020_12_31_424971 833 42 of of IN 10_1101-2020_12_31_424971 833 43 EF EF NNP 10_1101-2020_12_31_424971 833 44 - - HYPH 10_1101-2020_12_31_424971 833 45 G G NNP 10_1101-2020_12_31_424971 833 46 as as IN 10_1101-2020_12_31_424971 833 47 depicted depict VBN 10_1101-2020_12_31_424971 833 48 in in IN 10_1101-2020_12_31_424971 833 49 a a DT 10_1101-2020_12_31_424971 833 50 , , , 10_1101-2020_12_31_424971 833 51 b. b. NN 10_1101-2020_12_31_424971 834 1 The the DT 10_1101-2020_12_31_424971 834 2 waiting wait VBG 10_1101-2020_12_31_424971 834 3 times time NNS 10_1101-2020_12_31_424971 834 4 associated associate VBN 10_1101-2020_12_31_424971 834 5 with with IN 10_1101-2020_12_31_424971 834 6 k70S k70S NNP 10_1101-2020_12_31_424971 834 7 IC→GS2 IC→GS2 NNP 10_1101-2020_12_31_424971 834 8 , , , 10_1101-2020_12_31_424971 834 9 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 834 10 , , , 10_1101-2020_12_31_424971 834 11 1318 1318 CD 10_1101-2020_12_31_424971 834 12 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 834 13 , , , 10_1101-2020_12_31_424971 834 14 and and CC 10_1101-2020_12_31_424971 834 15 kGS2→POST kGS2→POST NNP 10_1101-2020_12_31_424971 834 16 are be VBP 10_1101-2020_12_31_424971 834 17 indicated indicate VBN 10_1101-2020_12_31_424971 834 18 in in IN 10_1101-2020_12_31_424971 834 19 each each DT 10_1101-2020_12_31_424971 834 20 hypothetical hypothetical JJ 10_1101-2020_12_31_424971 834 21 trajectory trajectory NN 10_1101-2020_12_31_424971 834 22 . . . 10_1101-2020_12_31_424971 835 1 f f NNP 10_1101-2020_12_31_424971 835 2 , , , 10_1101-2020_12_31_424971 835 3 g g NNP 10_1101-2020_12_31_424971 835 4 , , , 10_1101-2020_12_31_424971 835 5 and and CC 10_1101-2020_12_31_424971 835 6 h h NNP 10_1101-2020_12_31_424971 835 7 Surface Surface NNP 10_1101-2020_12_31_424971 835 8 contour contour NN 10_1101-2020_12_31_424971 835 9 1319 1319 CD 10_1101-2020_12_31_424971 835 10 .CC .CC , 10_1101-2020_12_31_424971 835 11 - - : 10_1101-2020_12_31_424971 835 12 BY by IN 10_1101-2020_12_31_424971 835 13 - - HYPH 10_1101-2020_12_31_424971 835 14 NC NC NNP 10_1101-2020_12_31_424971 835 15 - - HYPH 10_1101-2020_12_31_424971 835 16 ND ND NNP 10_1101-2020_12_31_424971 835 17 4.0 4.0 CD 10_1101-2020_12_31_424971 835 18 International International NNP 10_1101-2020_12_31_424971 835 19 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 835 20 under under IN 10_1101-2020_12_31_424971 835 21 a a DT 10_1101-2020_12_31_424971 835 22 ( ( -LRB- 10_1101-2020_12_31_424971 835 23 which which WDT 10_1101-2020_12_31_424971 835 24 was be VBD 10_1101-2020_12_31_424971 835 25 not not RB 10_1101-2020_12_31_424971 835 26 certified certify VBN 10_1101-2020_12_31_424971 835 27 by by IN 10_1101-2020_12_31_424971 835 28 peer peer NN 10_1101-2020_12_31_424971 835 29 review review NN 10_1101-2020_12_31_424971 835 30 ) ) -RRB- 10_1101-2020_12_31_424971 835 31 is be VBZ 10_1101-2020_12_31_424971 835 32 the the DT 10_1101-2020_12_31_424971 835 33 author author NN 10_1101-2020_12_31_424971 835 34 / / SYM 10_1101-2020_12_31_424971 835 35 funder funder NN 10_1101-2020_12_31_424971 835 36 , , , 10_1101-2020_12_31_424971 835 37 who who WP 10_1101-2020_12_31_424971 835 38 has have VBZ 10_1101-2020_12_31_424971 835 39 granted grant VBN 10_1101-2020_12_31_424971 835 40 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 835 41 a a DT 10_1101-2020_12_31_424971 835 42 license license NN 10_1101-2020_12_31_424971 835 43 to to TO 10_1101-2020_12_31_424971 835 44 display display VB 10_1101-2020_12_31_424971 835 45 the the DT 10_1101-2020_12_31_424971 835 46 preprint preprint NN 10_1101-2020_12_31_424971 835 47 in in IN 10_1101-2020_12_31_424971 835 48 perpetuity perpetuity NN 10_1101-2020_12_31_424971 835 49 . . . 10_1101-2020_12_31_424971 836 1 It -PRON- PRP 10_1101-2020_12_31_424971 836 2 is be VBZ 10_1101-2020_12_31_424971 836 3 made make VBN 10_1101-2020_12_31_424971 836 4 The the DT 10_1101-2020_12_31_424971 836 5 copyright copyright NN 10_1101-2020_12_31_424971 836 6 holder holder NN 10_1101-2020_12_31_424971 836 7 for for IN 10_1101-2020_12_31_424971 836 8 this this DT 10_1101-2020_12_31_424971 836 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 836 10 version version NN 10_1101-2020_12_31_424971 836 11 posted post VBD 10_1101-2020_12_31_424971 836 12 January January NNP 10_1101-2020_12_31_424971 836 13 2 2 CD 10_1101-2020_12_31_424971 836 14 , , , 10_1101-2020_12_31_424971 836 15 2021 2021 CD 10_1101-2020_12_31_424971 836 16 . . . 10_1101-2020_12_31_424971 836 17 ; ; : 10_1101-2020_12_31_424971 836 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 836 19 : : : 10_1101-2020_12_31_424971 836 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 836 21 preprint preprint NN 10_1101-2020_12_31_424971 836 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 836 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 836 24 51 51 CD 10_1101-2020_12_31_424971 836 25 plots plot NNS 10_1101-2020_12_31_424971 836 26 of of IN 10_1101-2020_12_31_424971 836 27 the the DT 10_1101-2020_12_31_424971 836 28 time time NN 10_1101-2020_12_31_424971 836 29 evolution evolution NN 10_1101-2020_12_31_424971 836 30 of of IN 10_1101-2020_12_31_424971 836 31 population population NN 10_1101-2020_12_31_424971 836 32 FRET FRET NNS 10_1101-2020_12_31_424971 836 33 obtained obtain VBN 10_1101-2020_12_31_424971 836 34 by by IN 10_1101-2020_12_31_424971 836 35 superimposing superimpose VBG 10_1101-2020_12_31_424971 836 36 individual individual JJ 10_1101-2020_12_31_424971 836 37 EFRET EFRET NNP 10_1101-2020_12_31_424971 836 38 vs. vs. IN 10_1101-2020_12_31_424971 836 39 time time NN 10_1101-2020_12_31_424971 836 40 1320 1320 CD 10_1101-2020_12_31_424971 836 41 trajectories trajectory NNS 10_1101-2020_12_31_424971 836 42 in in IN 10_1101-2020_12_31_424971 836 43 the the DT 10_1101-2020_12_31_424971 836 44 experiments experiment NNS 10_1101-2020_12_31_424971 836 45 in in IN 10_1101-2020_12_31_424971 836 46 a a DT 10_1101-2020_12_31_424971 836 47 , , , 10_1101-2020_12_31_424971 836 48 b b NN 10_1101-2020_12_31_424971 836 49 , , , 10_1101-2020_12_31_424971 836 50 and and CC 10_1101-2020_12_31_424971 836 51 c c NN 10_1101-2020_12_31_424971 836 52 , , , 10_1101-2020_12_31_424971 836 53 respectively respectively RB 10_1101-2020_12_31_424971 836 54 , , , 10_1101-2020_12_31_424971 836 55 for for IN 10_1101-2020_12_31_424971 836 56 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 836 57 ( ( -LRB- 10_1101-2020_12_31_424971 836 58 top top NN 10_1101-2020_12_31_424971 836 59 ) ) -RRB- 10_1101-2020_12_31_424971 836 60 and and CC 10_1101-2020_12_31_424971 836 61 ProL ProL NNP 10_1101-2020_12_31_424971 836 62 ( ( -LRB- 10_1101-2020_12_31_424971 836 63 bottom bottom NN 10_1101-2020_12_31_424971 836 64 ) ) -RRB- 10_1101-2020_12_31_424971 836 65 . . . 10_1101-2020_12_31_424971 837 1 N n CD 10_1101-2020_12_31_424971 837 2 : : : 10_1101-2020_12_31_424971 837 3 1321 1321 CD 10_1101-2020_12_31_424971 837 4 the the DT 10_1101-2020_12_31_424971 837 5 number number NN 10_1101-2020_12_31_424971 837 6 of of IN 10_1101-2020_12_31_424971 837 7 trajectories trajectory NNS 10_1101-2020_12_31_424971 837 8 used use VBN 10_1101-2020_12_31_424971 837 9 to to TO 10_1101-2020_12_31_424971 837 10 construct construct VB 10_1101-2020_12_31_424971 837 11 each each DT 10_1101-2020_12_31_424971 837 12 contour contour NN 10_1101-2020_12_31_424971 837 13 plot plot NN 10_1101-2020_12_31_424971 837 14 . . . 10_1101-2020_12_31_424971 838 1 Surface surface NN 10_1101-2020_12_31_424971 838 2 contours contour NNS 10_1101-2020_12_31_424971 838 3 are be VBP 10_1101-2020_12_31_424971 838 4 colored color VBN 10_1101-2020_12_31_424971 838 5 as as IN 10_1101-2020_12_31_424971 838 6 1322 1322 CD 10_1101-2020_12_31_424971 838 7 denoted denote VBN 10_1101-2020_12_31_424971 838 8 in in IN 10_1101-2020_12_31_424971 838 9 the the DT 10_1101-2020_12_31_424971 838 10 population population NN 10_1101-2020_12_31_424971 838 11 color color NN 10_1101-2020_12_31_424971 838 12 bars bar NNS 10_1101-2020_12_31_424971 838 13 . . . 10_1101-2020_12_31_424971 839 1 For for IN 10_1101-2020_12_31_424971 839 2 pre pre JJ 10_1101-2020_12_31_424971 839 3 - - JJ 10_1101-2020_12_31_424971 839 4 steady steady JJ 10_1101-2020_12_31_424971 839 5 - - HYPH 10_1101-2020_12_31_424971 839 6 state state NN 10_1101-2020_12_31_424971 839 7 experiments experiment NNS 10_1101-2020_12_31_424971 839 8 , , , 10_1101-2020_12_31_424971 839 9 the the DT 10_1101-2020_12_31_424971 839 10 black black JJ 10_1101-2020_12_31_424971 839 11 dashed dash VBN 10_1101-2020_12_31_424971 839 12 lines line NNS 10_1101-2020_12_31_424971 839 13 1323 1323 CD 10_1101-2020_12_31_424971 839 14 indicate indicate VBP 10_1101-2020_12_31_424971 839 15 the the DT 10_1101-2020_12_31_424971 839 16 time time NN 10_1101-2020_12_31_424971 839 17 at at IN 10_1101-2020_12_31_424971 839 18 which which WDT 10_1101-2020_12_31_424971 839 19 the the DT 10_1101-2020_12_31_424971 839 20 TC TC NNP 10_1101-2020_12_31_424971 839 21 was be VBD 10_1101-2020_12_31_424971 839 22 delivered deliver VBN 10_1101-2020_12_31_424971 839 23 and and CC 10_1101-2020_12_31_424971 839 24 the the DT 10_1101-2020_12_31_424971 839 25 gray gray JJ 10_1101-2020_12_31_424971 839 26 shaded shaded JJ 10_1101-2020_12_31_424971 839 27 areas area NNS 10_1101-2020_12_31_424971 839 28 denote denote VBP 10_1101-2020_12_31_424971 839 29 the the DT 10_1101-2020_12_31_424971 839 30 time time NN 10_1101-2020_12_31_424971 839 31 1324 1324 CD 10_1101-2020_12_31_424971 839 32 required require VBN 10_1101-2020_12_31_424971 839 33 for for IN 10_1101-2020_12_31_424971 839 34 the the DT 10_1101-2020_12_31_424971 839 35 majority majority NN 10_1101-2020_12_31_424971 839 36 ( ( -LRB- 10_1101-2020_12_31_424971 839 37 54 54 CD 10_1101-2020_12_31_424971 839 38 - - SYM 10_1101-2020_12_31_424971 839 39 68 68 CD 10_1101-2020_12_31_424971 839 40 % % NN 10_1101-2020_12_31_424971 839 41 ) ) -RRB- 10_1101-2020_12_31_424971 839 42 of of IN 10_1101-2020_12_31_424971 839 43 the the DT 10_1101-2020_12_31_424971 839 44 70S 70S NNP 10_1101-2020_12_31_424971 839 45 ICs ic NNS 10_1101-2020_12_31_424971 839 46 to to TO 10_1101-2020_12_31_424971 839 47 transition transition NN 10_1101-2020_12_31_424971 839 48 to to IN 10_1101-2020_12_31_424971 839 49 GS2 GS2 NNP 10_1101-2020_12_31_424971 839 50 . . . 10_1101-2020_12_31_424971 840 1 Note note VB 10_1101-2020_12_31_424971 840 2 that that IN 10_1101-2020_12_31_424971 840 3 the the DT 10_1101-2020_12_31_424971 840 4 rate rate NN 10_1101-2020_12_31_424971 840 5 of of IN 10_1101-2020_12_31_424971 840 6 1325 1325 CD 10_1101-2020_12_31_424971 840 7 deacylated deacylated JJ 10_1101-2020_12_31_424971 840 8 SufB2 sufb2 JJ 10_1101-2020_12_31_424971 840 9 dissociation dissociation NN 10_1101-2020_12_31_424971 840 10 from from IN 10_1101-2020_12_31_424971 840 11 the the DT 10_1101-2020_12_31_424971 840 12 A A NNP 10_1101-2020_12_31_424971 840 13 site site NN 10_1101-2020_12_31_424971 840 14 under under IN 10_1101-2020_12_31_424971 840 15 our -PRON- PRP$ 10_1101-2020_12_31_424971 840 16 conditions condition NNS 10_1101-2020_12_31_424971 840 17 is be VBZ 10_1101-2020_12_31_424971 840 18 similar similar JJ 10_1101-2020_12_31_424971 840 19 to to IN 10_1101-2020_12_31_424971 840 20 that that DT 10_1101-2020_12_31_424971 840 21 of of IN 10_1101-2020_12_31_424971 840 22 EF EF NNP 10_1101-2020_12_31_424971 840 23 - - HYPH 10_1101-2020_12_31_424971 840 24 G-1326 G-1326 NNP 10_1101-2020_12_31_424971 840 25 catalyzed catalyzed JJ 10_1101-2020_12_31_424971 840 26 translocation translocation NN 10_1101-2020_12_31_424971 840 27 , , , 10_1101-2020_12_31_424971 840 28 thereby thereby RB 10_1101-2020_12_31_424971 840 29 resulting result VBG 10_1101-2020_12_31_424971 840 30 in in IN 10_1101-2020_12_31_424971 840 31 the the DT 10_1101-2020_12_31_424971 840 32 buildup buildup NN 10_1101-2020_12_31_424971 840 33 of of IN 10_1101-2020_12_31_424971 840 34 a a DT 10_1101-2020_12_31_424971 840 35 PRE PRE NNP 10_1101-2020_12_31_424971 840 36 complex complex JJ 10_1101-2020_12_31_424971 840 37 sub sub NN 10_1101-2020_12_31_424971 840 38 - - NN 10_1101-2020_12_31_424971 840 39 population population NN 10_1101-2020_12_31_424971 840 40 over over IN 10_1101-2020_12_31_424971 840 41 3 3 CD 10_1101-2020_12_31_424971 840 42 - - SYM 10_1101-2020_12_31_424971 840 43 1327 1327 CD 10_1101-2020_12_31_424971 840 44 20 20 CD 10_1101-2020_12_31_424971 840 45 min min NN 10_1101-2020_12_31_424971 840 46 post post JJ 10_1101-2020_12_31_424971 840 47 - - JJ 10_1101-2020_12_31_424971 840 48 delivery delivery NN 10_1101-2020_12_31_424971 840 49 that that WDT 10_1101-2020_12_31_424971 840 50 lacks lack VBZ 10_1101-2020_12_31_424971 840 51 an an DT 10_1101-2020_12_31_424971 840 52 A a NN 10_1101-2020_12_31_424971 840 53 site site NN 10_1101-2020_12_31_424971 840 54 tRNA trna NN 10_1101-2020_12_31_424971 840 55 and and CC 10_1101-2020_12_31_424971 840 56 is be VBZ 10_1101-2020_12_31_424971 840 57 incapable incapable JJ 10_1101-2020_12_31_424971 840 58 of of IN 10_1101-2020_12_31_424971 840 59 translocation translocation NN 10_1101-2020_12_31_424971 840 60 . . . 10_1101-2020_12_31_424971 841 1 This this DT 10_1101-2020_12_31_424971 841 2 sub-1328 sub-1328 JJ 10_1101-2020_12_31_424971 841 3 population population NN 10_1101-2020_12_31_424971 841 4 exhibits exhibit VBZ 10_1101-2020_12_31_424971 841 5 kGS1→GS2 kgs1→gs2 CD 10_1101-2020_12_31_424971 841 6 , , , 10_1101-2020_12_31_424971 841 7 kGS2→GS1 kgs2→gs1 CD 10_1101-2020_12_31_424971 841 8 , , , 10_1101-2020_12_31_424971 841 9 and and CC 10_1101-2020_12_31_424971 841 10 Keq Keq NNP 10_1101-2020_12_31_424971 841 11 values value NNS 10_1101-2020_12_31_424971 841 12 similar similar JJ 10_1101-2020_12_31_424971 841 13 to to IN 10_1101-2020_12_31_424971 841 14 those those DT 10_1101-2020_12_31_424971 841 15 observed observe VBN 10_1101-2020_12_31_424971 841 16 in in IN 10_1101-2020_12_31_424971 841 17 experiments experiment NNS 10_1101-2020_12_31_424971 841 18 1329 1329 CD 10_1101-2020_12_31_424971 841 19 recorded record VBN 10_1101-2020_12_31_424971 841 20 in in IN 10_1101-2020_12_31_424971 841 21 the the DT 10_1101-2020_12_31_424971 841 22 absence absence NN 10_1101-2020_12_31_424971 841 23 of of IN 10_1101-2020_12_31_424971 841 24 EF EF NNP 10_1101-2020_12_31_424971 841 25 - - HYPH 10_1101-2020_12_31_424971 841 26 G G NNP 10_1101-2020_12_31_424971 841 27 ( ( -LRB- 10_1101-2020_12_31_424971 841 28 Supplementary Supplementary NNP 10_1101-2020_12_31_424971 841 29 Table Table NNP 10_1101-2020_12_31_424971 841 30 6 6 CD 10_1101-2020_12_31_424971 841 31 ) ) -RRB- 10_1101-2020_12_31_424971 841 32 . . . 10_1101-2020_12_31_424971 842 1 1330 1330 CD 10_1101-2020_12_31_424971 842 2 1331 1331 CD 10_1101-2020_12_31_424971 842 3 Figure Figure NNP 10_1101-2020_12_31_424971 842 4 7 7 CD 10_1101-2020_12_31_424971 842 5 . . . 10_1101-2020_12_31_424971 843 1 Structure structure NN 10_1101-2020_12_31_424971 843 2 - - HYPH 10_1101-2020_12_31_424971 843 3 based base VBN 10_1101-2020_12_31_424971 843 4 mechanistic mechanistic JJ 10_1101-2020_12_31_424971 843 5 model model NN 10_1101-2020_12_31_424971 843 6 for for IN 10_1101-2020_12_31_424971 843 7 SufB2-induced SufB2-induced NNP 10_1101-2020_12_31_424971 843 8 +1 +1 NNS 10_1101-2020_12_31_424971 843 9 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 843 10 . . . 10_1101-2020_12_31_424971 844 1 A a DT 10_1101-2020_12_31_424971 844 2 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 844 3 - - HYPH 10_1101-2020_12_31_424971 844 4 1332 1332 CD 10_1101-2020_12_31_424971 844 5 TC TC NNP 10_1101-2020_12_31_424971 844 6 uses use VBZ 10_1101-2020_12_31_424971 844 7 triplet triplet NN 10_1101-2020_12_31_424971 844 8 anticodon anticodon NN 10_1101-2020_12_31_424971 844 9 - - HYPH 10_1101-2020_12_31_424971 844 10 codon codon NNP 10_1101-2020_12_31_424971 844 11 pairing pair VBG 10_1101-2020_12_31_424971 844 12 in in IN 10_1101-2020_12_31_424971 844 13 the the DT 10_1101-2020_12_31_424971 844 14 0-frame 0-frame CD 10_1101-2020_12_31_424971 844 15 at at IN 10_1101-2020_12_31_424971 844 16 a a DT 10_1101-2020_12_31_424971 844 17 CCC CCC NNP 10_1101-2020_12_31_424971 844 18 - - HYPH 10_1101-2020_12_31_424971 844 19 C C NNP 10_1101-2020_12_31_424971 844 20 motif motif NN 10_1101-2020_12_31_424971 844 21 , , , 10_1101-2020_12_31_424971 844 22 undergoes undergo VBZ 10_1101-2020_12_31_424971 844 23 peptide peptide NN 10_1101-2020_12_31_424971 844 24 - - HYPH 10_1101-2020_12_31_424971 844 25 bond bond NN 10_1101-2020_12_31_424971 844 26 1333 1333 CD 10_1101-2020_12_31_424971 844 27 formation formation NN 10_1101-2020_12_31_424971 844 28 , , , 10_1101-2020_12_31_424971 844 29 and and CC 10_1101-2020_12_31_424971 844 30 enables enable VBZ 10_1101-2020_12_31_424971 844 31 the the DT 10_1101-2020_12_31_424971 844 32 resulting result VBG 10_1101-2020_12_31_424971 844 33 PRE PRE NNP 10_1101-2020_12_31_424971 844 34 complex complex JJ 10_1101-2020_12_31_424971 844 35 to to TO 10_1101-2020_12_31_424971 844 36 undergo undergo VB 10_1101-2020_12_31_424971 844 37 a a DT 10_1101-2020_12_31_424971 844 38 GS1→GS2 GS1→GS2 NNP 10_1101-2020_12_31_424971 844 39 transition transition NN 10_1101-2020_12_31_424971 844 40 , , , 10_1101-2020_12_31_424971 844 41 all all DT 10_1101-2020_12_31_424971 844 42 with with IN 10_1101-2020_12_31_424971 844 43 1334 1334 CD 10_1101-2020_12_31_424971 844 44 rates rate NNS 10_1101-2020_12_31_424971 844 45 similar similar JJ 10_1101-2020_12_31_424971 844 46 to to IN 10_1101-2020_12_31_424971 844 47 those those DT 10_1101-2020_12_31_424971 844 48 of of IN 10_1101-2020_12_31_424971 844 49 ProL ProL NNP 10_1101-2020_12_31_424971 844 50 - - HYPH 10_1101-2020_12_31_424971 844 51 TC TC NNP 10_1101-2020_12_31_424971 844 52 . . . 10_1101-2020_12_31_424971 845 1 During during IN 10_1101-2020_12_31_424971 845 2 the the DT 10_1101-2020_12_31_424971 845 3 GS1→GS2 GS1→GS2 NNP 10_1101-2020_12_31_424971 845 4 transition transition NN 10_1101-2020_12_31_424971 845 5 , , , 10_1101-2020_12_31_424971 845 6 the the DT 10_1101-2020_12_31_424971 845 7 30S 30s CD 10_1101-2020_12_31_424971 845 8 subunit subunit NN 10_1101-2020_12_31_424971 845 9 rotates rotate VBZ 10_1101-2020_12_31_424971 845 10 1335 1335 CD 10_1101-2020_12_31_424971 845 11 relative relative JJ 10_1101-2020_12_31_424971 845 12 to to IN 10_1101-2020_12_31_424971 845 13 the the DT 10_1101-2020_12_31_424971 845 14 50S 50S NNP 10_1101-2020_12_31_424971 845 15 subunit subunit NN 10_1101-2020_12_31_424971 845 16 by by IN 10_1101-2020_12_31_424971 845 17 8º 8º NNS 10_1101-2020_12_31_424971 845 18 in in IN 10_1101-2020_12_31_424971 845 19 the the DT 10_1101-2020_12_31_424971 845 20 counter counter NN 10_1101-2020_12_31_424971 845 21 - - NN 10_1101-2020_12_31_424971 845 22 clockwise clockwise NN 10_1101-2020_12_31_424971 845 23 ( ( -LRB- 10_1101-2020_12_31_424971 845 24 + + SYM 10_1101-2020_12_31_424971 845 25 ) ) -RRB- 10_1101-2020_12_31_424971 845 26 direction direction NN 10_1101-2020_12_31_424971 845 27 along along IN 10_1101-2020_12_31_424971 845 28 the the DT 10_1101-2020_12_31_424971 845 29 black black JJ 10_1101-2020_12_31_424971 845 30 curved curved JJ 10_1101-2020_12_31_424971 845 31 1336 1336 CD 10_1101-2020_12_31_424971 845 32 arrow arrow NN 10_1101-2020_12_31_424971 845 33 ; ; : 10_1101-2020_12_31_424971 845 34 the the DT 10_1101-2020_12_31_424971 845 35 30S 30s CD 10_1101-2020_12_31_424971 845 36 subunit subunit NN 10_1101-2020_12_31_424971 845 37 head head NN 10_1101-2020_12_31_424971 845 38 swivels swivel VBZ 10_1101-2020_12_31_424971 845 39 relative relative JJ 10_1101-2020_12_31_424971 845 40 to to IN 10_1101-2020_12_31_424971 845 41 the the DT 10_1101-2020_12_31_424971 845 42 30S 30s CD 10_1101-2020_12_31_424971 845 43 subunit subunit NN 10_1101-2020_12_31_424971 845 44 body body NN 10_1101-2020_12_31_424971 845 45 by by IN 10_1101-2020_12_31_424971 845 46 5º 5º CD 10_1101-2020_12_31_424971 845 47 in in IN 10_1101-2020_12_31_424971 845 48 the the DT 10_1101-2020_12_31_424971 845 49 clockwise clockwise NN 10_1101-2020_12_31_424971 845 50 ( ( -LRB- 10_1101-2020_12_31_424971 845 51 – – : 10_1101-2020_12_31_424971 845 52 ) ) -RRB- 10_1101-2020_12_31_424971 845 53 1337 1337 CD 10_1101-2020_12_31_424971 845 54 direction direction NN 10_1101-2020_12_31_424971 845 55 against against IN 10_1101-2020_12_31_424971 845 56 the the DT 10_1101-2020_12_31_424971 845 57 black black JJ 10_1101-2020_12_31_424971 845 58 curved curved NNP 10_1101-2020_12_31_424971 845 59 arrow arrow NN 10_1101-2020_12_31_424971 845 60 ; ; : 10_1101-2020_12_31_424971 845 61 the the DT 10_1101-2020_12_31_424971 845 62 L1 L1 NNP 10_1101-2020_12_31_424971 845 63 stalk stalk NN 10_1101-2020_12_31_424971 845 64 closes close VBZ 10_1101-2020_12_31_424971 845 65 by by IN 10_1101-2020_12_31_424971 845 66 ~60 ~60 NFP 10_1101-2020_12_31_424971 845 67 Å å NN 10_1101-2020_12_31_424971 845 68 ; ; : 10_1101-2020_12_31_424971 845 69 and and CC 10_1101-2020_12_31_424971 845 70 the the DT 10_1101-2020_12_31_424971 845 71 tRNAs trnas CD 10_1101-2020_12_31_424971 845 72 are be VBP 10_1101-2020_12_31_424971 845 73 1338 1338 CD 10_1101-2020_12_31_424971 845 74 reconfigured reconfigure VBN 10_1101-2020_12_31_424971 845 75 from from IN 10_1101-2020_12_31_424971 845 76 their -PRON- PRP$ 10_1101-2020_12_31_424971 845 77 P p NN 10_1101-2020_12_31_424971 845 78 / / SYM 10_1101-2020_12_31_424971 845 79 P p NN 10_1101-2020_12_31_424971 845 80 and and CC 10_1101-2020_12_31_424971 845 81 A a NN 10_1101-2020_12_31_424971 845 82 / / , 10_1101-2020_12_31_424971 845 83 A a NN 10_1101-2020_12_31_424971 845 84 to to IN 10_1101-2020_12_31_424971 845 85 their -PRON- PRP$ 10_1101-2020_12_31_424971 845 86 P p NN 10_1101-2020_12_31_424971 845 87 / / SYM 10_1101-2020_12_31_424971 845 88 E e NN 10_1101-2020_12_31_424971 845 89 and and CC 10_1101-2020_12_31_424971 845 90 A a NN 10_1101-2020_12_31_424971 845 91 / / SYM 10_1101-2020_12_31_424971 845 92 P p NN 10_1101-2020_12_31_424971 845 93 configurations configuration NNS 10_1101-2020_12_31_424971 845 94 . . . 10_1101-2020_12_31_424971 846 1 EF EF NNP 10_1101-2020_12_31_424971 846 2 - - HYPH 10_1101-2020_12_31_424971 846 3 G G NNP 10_1101-2020_12_31_424971 846 4 then then RB 10_1101-2020_12_31_424971 846 5 binds bind VBZ 10_1101-2020_12_31_424971 846 6 to to IN 10_1101-2020_12_31_424971 846 7 the the DT 10_1101-2020_12_31_424971 846 8 1339 1339 CD 10_1101-2020_12_31_424971 846 9 PRE PRE NNP 10_1101-2020_12_31_424971 846 10 complex complex JJ 10_1101-2020_12_31_424971 846 11 to to TO 10_1101-2020_12_31_424971 846 12 form form VB 10_1101-2020_12_31_424971 846 13 PRE PRE NNP 10_1101-2020_12_31_424971 846 14 - - HYPH 10_1101-2020_12_31_424971 846 15 G1 G1 NNP 10_1101-2020_12_31_424971 846 16 and and CC 10_1101-2020_12_31_424971 846 17 subsequently subsequently RB 10_1101-2020_12_31_424971 846 18 catalyzes catalyze VBZ 10_1101-2020_12_31_424971 846 19 a a DT 10_1101-2020_12_31_424971 846 20 series series NN 10_1101-2020_12_31_424971 846 21 of of IN 10_1101-2020_12_31_424971 846 22 conformational conformational JJ 10_1101-2020_12_31_424971 846 23 1340 1340 CD 10_1101-2020_12_31_424971 846 24 rearrangements rearrangement NNS 10_1101-2020_12_31_424971 846 25 of of IN 10_1101-2020_12_31_424971 846 26 the the DT 10_1101-2020_12_31_424971 846 27 complex complex NN 10_1101-2020_12_31_424971 846 28 ( ( -LRB- 10_1101-2020_12_31_424971 846 29 PRE PRE NNP 10_1101-2020_12_31_424971 846 30 - - HYPH 10_1101-2020_12_31_424971 846 31 G1 G1 NNP 10_1101-2020_12_31_424971 846 32 to to IN 10_1101-2020_12_31_424971 846 33 PRE PRE NNP 10_1101-2020_12_31_424971 846 34 - - HYPH 10_1101-2020_12_31_424971 846 35 G4 G4 NNP 10_1101-2020_12_31_424971 846 36 ) ) -RRB- 10_1101-2020_12_31_424971 846 37 that that WDT 10_1101-2020_12_31_424971 846 38 encompass encompass VBP 10_1101-2020_12_31_424971 846 39 further further JJ 10_1101-2020_12_31_424971 846 40 counter counter NN 10_1101-2020_12_31_424971 846 41 - - NN 10_1101-2020_12_31_424971 846 42 clockwise clockwise JJ 10_1101-2020_12_31_424971 846 43 1341 1341 CD 10_1101-2020_12_31_424971 846 44 and and CC 10_1101-2020_12_31_424971 846 45 clockwise clockwise NN 10_1101-2020_12_31_424971 846 46 rotations rotation NNS 10_1101-2020_12_31_424971 846 47 of of IN 10_1101-2020_12_31_424971 846 48 the the DT 10_1101-2020_12_31_424971 846 49 subunits subunit NNS 10_1101-2020_12_31_424971 846 50 ; ; , 10_1101-2020_12_31_424971 846 51 severing sever VBG 10_1101-2020_12_31_424971 846 52 of of IN 10_1101-2020_12_31_424971 846 53 decoding decode VBG 10_1101-2020_12_31_424971 846 54 center center NN 10_1101-2020_12_31_424971 846 55 interactions interaction NNS 10_1101-2020_12_31_424971 846 56 with with IN 10_1101-2020_12_31_424971 846 57 the the DT 10_1101-2020_12_31_424971 846 58 1342 1342 CD 10_1101-2020_12_31_424971 846 59 anticodon anticodon NNP 10_1101-2020_12_31_424971 846 60 - - HYPH 10_1101-2020_12_31_424971 846 61 codon codon NNP 10_1101-2020_12_31_424971 846 62 duplex duplex NN 10_1101-2020_12_31_424971 846 63 in in IN 10_1101-2020_12_31_424971 846 64 the the DT 10_1101-2020_12_31_424971 846 65 A A NNP 10_1101-2020_12_31_424971 846 66 site site NN 10_1101-2020_12_31_424971 846 67 ; ; : 10_1101-2020_12_31_424971 846 68 counter counter NN 10_1101-2020_12_31_424971 846 69 - - NN 10_1101-2020_12_31_424971 846 70 clockwise clockwise NN 10_1101-2020_12_31_424971 846 71 and and CC 10_1101-2020_12_31_424971 846 72 clockwise clockwise NN 10_1101-2020_12_31_424971 846 73 swiveling swiveling NN 10_1101-2020_12_31_424971 846 74 of of IN 10_1101-2020_12_31_424971 846 75 the the DT 10_1101-2020_12_31_424971 846 76 head head NN 10_1101-2020_12_31_424971 846 77 and and CC 10_1101-2020_12_31_424971 846 78 1343 1343 CD 10_1101-2020_12_31_424971 846 79 the the DT 10_1101-2020_12_31_424971 846 80 associated associate VBN 10_1101-2020_12_31_424971 846 81 opening opening NN 10_1101-2020_12_31_424971 846 82 and and CC 10_1101-2020_12_31_424971 846 83 closing closing NN 10_1101-2020_12_31_424971 846 84 of of IN 10_1101-2020_12_31_424971 846 85 the the DT 10_1101-2020_12_31_424971 846 86 E e NN 10_1101-2020_12_31_424971 846 87 - - NN 10_1101-2020_12_31_424971 846 88 site site NN 10_1101-2020_12_31_424971 846 89 gate gate NN 10_1101-2020_12_31_424971 846 90 ; ; , 10_1101-2020_12_31_424971 846 91 opening opening NN 10_1101-2020_12_31_424971 846 92 of of IN 10_1101-2020_12_31_424971 846 93 the the DT 10_1101-2020_12_31_424971 846 94 L1 L1 NNP 10_1101-2020_12_31_424971 846 95 stalk stalk NN 10_1101-2020_12_31_424971 846 96 ; ; : 10_1101-2020_12_31_424971 846 97 and and CC 10_1101-2020_12_31_424971 846 98 1344 1344 CD 10_1101-2020_12_31_424971 846 99 .CC .CC NFP 10_1101-2020_12_31_424971 846 100 - - HYPH 10_1101-2020_12_31_424971 846 101 BY by IN 10_1101-2020_12_31_424971 846 102 - - HYPH 10_1101-2020_12_31_424971 846 103 NC NC NNP 10_1101-2020_12_31_424971 846 104 - - HYPH 10_1101-2020_12_31_424971 846 105 ND ND NNP 10_1101-2020_12_31_424971 846 106 4.0 4.0 CD 10_1101-2020_12_31_424971 846 107 International International NNP 10_1101-2020_12_31_424971 846 108 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 846 109 under under IN 10_1101-2020_12_31_424971 846 110 a a DT 10_1101-2020_12_31_424971 846 111 ( ( -LRB- 10_1101-2020_12_31_424971 846 112 which which WDT 10_1101-2020_12_31_424971 846 113 was be VBD 10_1101-2020_12_31_424971 846 114 not not RB 10_1101-2020_12_31_424971 846 115 certified certify VBN 10_1101-2020_12_31_424971 846 116 by by IN 10_1101-2020_12_31_424971 846 117 peer peer NN 10_1101-2020_12_31_424971 846 118 review review NN 10_1101-2020_12_31_424971 846 119 ) ) -RRB- 10_1101-2020_12_31_424971 846 120 is be VBZ 10_1101-2020_12_31_424971 846 121 the the DT 10_1101-2020_12_31_424971 846 122 author author NN 10_1101-2020_12_31_424971 846 123 / / SYM 10_1101-2020_12_31_424971 846 124 funder funder NN 10_1101-2020_12_31_424971 846 125 , , , 10_1101-2020_12_31_424971 846 126 who who WP 10_1101-2020_12_31_424971 846 127 has have VBZ 10_1101-2020_12_31_424971 846 128 granted grant VBN 10_1101-2020_12_31_424971 846 129 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 846 130 a a DT 10_1101-2020_12_31_424971 846 131 license license NN 10_1101-2020_12_31_424971 846 132 to to TO 10_1101-2020_12_31_424971 846 133 display display VB 10_1101-2020_12_31_424971 846 134 the the DT 10_1101-2020_12_31_424971 846 135 preprint preprint NN 10_1101-2020_12_31_424971 846 136 in in IN 10_1101-2020_12_31_424971 846 137 perpetuity perpetuity NN 10_1101-2020_12_31_424971 846 138 . . . 10_1101-2020_12_31_424971 847 1 It -PRON- PRP 10_1101-2020_12_31_424971 847 2 is be VBZ 10_1101-2020_12_31_424971 847 3 made make VBN 10_1101-2020_12_31_424971 847 4 The the DT 10_1101-2020_12_31_424971 847 5 copyright copyright NN 10_1101-2020_12_31_424971 847 6 holder holder NN 10_1101-2020_12_31_424971 847 7 for for IN 10_1101-2020_12_31_424971 847 8 this this DT 10_1101-2020_12_31_424971 847 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 847 10 version version NN 10_1101-2020_12_31_424971 847 11 posted post VBD 10_1101-2020_12_31_424971 847 12 January January NNP 10_1101-2020_12_31_424971 847 13 2 2 CD 10_1101-2020_12_31_424971 847 14 , , , 10_1101-2020_12_31_424971 847 15 2021 2021 CD 10_1101-2020_12_31_424971 847 16 . . . 10_1101-2020_12_31_424971 847 17 ; ; : 10_1101-2020_12_31_424971 847 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 847 19 : : : 10_1101-2020_12_31_424971 847 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 847 21 preprint preprint NN 10_1101-2020_12_31_424971 847 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 NN 10_1101-2020_12_31_424971 847 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 847 24 52 52 CD 10_1101-2020_12_31_424971 847 25 reconfigurations reconfiguration NNS 10_1101-2020_12_31_424971 847 26 of of IN 10_1101-2020_12_31_424971 847 27 the the DT 10_1101-2020_12_31_424971 847 28 tRNAs trnas CD 10_1101-2020_12_31_424971 847 29 as as IN 10_1101-2020_12_31_424971 847 30 they -PRON- PRP 10_1101-2020_12_31_424971 847 31 move move VBP 10_1101-2020_12_31_424971 847 32 from from IN 10_1101-2020_12_31_424971 847 33 the the DT 10_1101-2020_12_31_424971 847 34 P p NN 10_1101-2020_12_31_424971 847 35 and and CC 10_1101-2020_12_31_424971 847 36 A a DT 10_1101-2020_12_31_424971 847 37 sites site NNS 10_1101-2020_12_31_424971 847 38 to to IN 10_1101-2020_12_31_424971 847 39 the the DT 10_1101-2020_12_31_424971 847 40 E e NN 10_1101-2020_12_31_424971 847 41 and and CC 10_1101-2020_12_31_424971 847 42 P p NN 10_1101-2020_12_31_424971 847 43 sites site NNS 10_1101-2020_12_31_424971 847 44 . . . 10_1101-2020_12_31_424971 848 1 It -PRON- PRP 10_1101-2020_12_31_424971 848 2 is be VBZ 10_1101-2020_12_31_424971 848 3 during during IN 10_1101-2020_12_31_424971 848 4 1345 1345 CD 10_1101-2020_12_31_424971 848 5 these these DT 10_1101-2020_12_31_424971 848 6 steps step NNS 10_1101-2020_12_31_424971 848 7 , , , 10_1101-2020_12_31_424971 848 8 shown show VBN 10_1101-2020_12_31_424971 848 9 in in IN 10_1101-2020_12_31_424971 848 10 red red JJ 10_1101-2020_12_31_424971 848 11 arrows arrow NNS 10_1101-2020_12_31_424971 848 12 within within IN 10_1101-2020_12_31_424971 848 13 the the DT 10_1101-2020_12_31_424971 848 14 gray gray JJ 10_1101-2020_12_31_424971 848 15 shaded shaded JJ 10_1101-2020_12_31_424971 848 16 box box NN 10_1101-2020_12_31_424971 848 17 , , , 10_1101-2020_12_31_424971 848 18 that that IN 10_1101-2020_12_31_424971 848 19 SufB2 SufB2 NNP 10_1101-2020_12_31_424971 848 20 impedes impede NNS 10_1101-2020_12_31_424971 848 21 forward forward RB 10_1101-2020_12_31_424971 848 22 and/or and/or CC 10_1101-2020_12_31_424971 848 23 1346 1346 CD 10_1101-2020_12_31_424971 848 24 reverse reverse JJ 10_1101-2020_12_31_424971 848 25 swiveling swiveling NN 10_1101-2020_12_31_424971 848 26 of of IN 10_1101-2020_12_31_424971 848 27 the the DT 10_1101-2020_12_31_424971 848 28 head head NN 10_1101-2020_12_31_424971 848 29 and and CC 10_1101-2020_12_31_424971 848 30 the the DT 10_1101-2020_12_31_424971 848 31 associated associate VBN 10_1101-2020_12_31_424971 848 32 opening open VBG 10_1101-2020_12_31_424971 848 33 and/or and/or CC 10_1101-2020_12_31_424971 848 34 closing closing NN 10_1101-2020_12_31_424971 848 35 of of IN 10_1101-2020_12_31_424971 848 36 the the DT 10_1101-2020_12_31_424971 848 37 E e NN 10_1101-2020_12_31_424971 848 38 - - NN 10_1101-2020_12_31_424971 848 39 site site NN 10_1101-2020_12_31_424971 848 40 gate gate NN 10_1101-2020_12_31_424971 848 41 , , , 10_1101-2020_12_31_424971 848 42 1347 1347 CD 10_1101-2020_12_31_424971 848 43 facilitating facilitate VBG 10_1101-2020_12_31_424971 848 44 +1 +1 NNS 10_1101-2020_12_31_424971 848 45 frameshifting frameshifte VBG 10_1101-2020_12_31_424971 848 46 . . . 10_1101-2020_12_31_424971 849 1 Next next RB 10_1101-2020_12_31_424971 849 2 , , , 10_1101-2020_12_31_424971 849 3 EF EF NNP 10_1101-2020_12_31_424971 849 4 - - HYPH 10_1101-2020_12_31_424971 849 5 G G NNP 10_1101-2020_12_31_424971 849 6 and and CC 10_1101-2020_12_31_424971 849 7 the the DT 10_1101-2020_12_31_424971 849 8 deacylated deacylated JJ 10_1101-2020_12_31_424971 849 9 tRNA trna NN 10_1101-2020_12_31_424971 849 10 dissociate dissociate NN 10_1101-2020_12_31_424971 849 11 from from IN 10_1101-2020_12_31_424971 849 12 PRE PRE NNP 10_1101-2020_12_31_424971 849 13 - - HYPH 10_1101-2020_12_31_424971 849 14 G4 G4 NNP 10_1101-2020_12_31_424971 849 15 , , , 10_1101-2020_12_31_424971 849 16 1348 1348 CD 10_1101-2020_12_31_424971 849 17 leaving leave VBG 10_1101-2020_12_31_424971 849 18 a a DT 10_1101-2020_12_31_424971 849 19 POST POST NNP 10_1101-2020_12_31_424971 849 20 complex complex NN 10_1101-2020_12_31_424971 849 21 ready ready JJ 10_1101-2020_12_31_424971 849 22 to to TO 10_1101-2020_12_31_424971 849 23 enter enter VB 10_1101-2020_12_31_424971 849 24 the the DT 10_1101-2020_12_31_424971 849 25 next next JJ 10_1101-2020_12_31_424971 849 26 elongation elongation NN 10_1101-2020_12_31_424971 849 27 cycle cycle NN 10_1101-2020_12_31_424971 849 28 . . . 10_1101-2020_12_31_424971 850 1 The the DT 10_1101-2020_12_31_424971 850 2 cartoons cartoon NNS 10_1101-2020_12_31_424971 850 3 depicting depict VBG 10_1101-2020_12_31_424971 850 4 PRE-1349 PRE-1349 NNP 10_1101-2020_12_31_424971 850 5 G1(GS1 G1(GS1 NNP 10_1101-2020_12_31_424971 850 6 ) ) -RRB- 10_1101-2020_12_31_424971 850 7 and and CC 10_1101-2020_12_31_424971 850 8 PRE PRE NNP 10_1101-2020_12_31_424971 850 9 - - HYPH 10_1101-2020_12_31_424971 850 10 G1(GS2 G1(GS2 NNP 10_1101-2020_12_31_424971 850 11 ) ) -RRB- 10_1101-2020_12_31_424971 850 12 were be VBD 10_1101-2020_12_31_424971 850 13 generated generate VBN 10_1101-2020_12_31_424971 850 14 using use VBG 10_1101-2020_12_31_424971 850 15 Biological Biological NNP 10_1101-2020_12_31_424971 850 16 Assemblies Assemblies NNPS 10_1101-2020_12_31_424971 850 17 2 2 CD 10_1101-2020_12_31_424971 850 18 and and CC 10_1101-2020_12_31_424971 850 19 1 1 CD 10_1101-2020_12_31_424971 850 20 , , , 10_1101-2020_12_31_424971 850 21 respectively respectively RB 10_1101-2020_12_31_424971 850 22 , , , 10_1101-2020_12_31_424971 850 23 1350 1350 CD 10_1101-2020_12_31_424971 850 24 of of IN 10_1101-2020_12_31_424971 850 25 PDB PDB NNP 10_1101-2020_12_31_424971 850 26 entry entry NN 10_1101-2020_12_31_424971 850 27 4V9D. 4v9d. CD 10_1101-2020_12_31_424971 851 1 Due due IN 10_1101-2020_12_31_424971 851 2 to to IN 10_1101-2020_12_31_424971 851 3 the the DT 10_1101-2020_12_31_424971 851 4 lack lack NN 10_1101-2020_12_31_424971 851 5 of of IN 10_1101-2020_12_31_424971 851 6 an an DT 10_1101-2020_12_31_424971 851 7 A A NNP 10_1101-2020_12_31_424971 851 8 - - HYPH 10_1101-2020_12_31_424971 851 9 site site NN 10_1101-2020_12_31_424971 851 10 tRNA trna NN 10_1101-2020_12_31_424971 851 11 or or CC 10_1101-2020_12_31_424971 851 12 EF EF NNP 10_1101-2020_12_31_424971 851 13 - - HYPH 10_1101-2020_12_31_424971 851 14 G G NNP 10_1101-2020_12_31_424971 851 15 in in IN 10_1101-2020_12_31_424971 851 16 4V9D 4v9d CD 10_1101-2020_12_31_424971 851 17 , , , 10_1101-2020_12_31_424971 851 18 cartoons cartoon NNS 10_1101-2020_12_31_424971 851 19 of of IN 10_1101-2020_12_31_424971 851 20 the the DT 10_1101-2020_12_31_424971 851 21 A- a- XX 10_1101-2020_12_31_424971 851 22 and and CC 10_1101-2020_12_31_424971 851 23 1351 1351 CD 10_1101-2020_12_31_424971 851 24 P p NN 10_1101-2020_12_31_424971 851 25 - - HYPH 10_1101-2020_12_31_424971 851 26 site site NN 10_1101-2020_12_31_424971 851 27 tRNAs trnas CD 10_1101-2020_12_31_424971 851 28 from from IN 10_1101-2020_12_31_424971 851 29 previous previous JJ 10_1101-2020_12_31_424971 851 30 structures1 structures1 NN 10_1101-2020_12_31_424971 851 31 were be VBD 10_1101-2020_12_31_424971 851 32 positioned position VBN 10_1101-2020_12_31_424971 851 33 into into IN 10_1101-2020_12_31_424971 851 34 the the DT 10_1101-2020_12_31_424971 851 35 two two CD 10_1101-2020_12_31_424971 851 36 assemblies assembly NNS 10_1101-2020_12_31_424971 851 37 using use VBG 10_1101-2020_12_31_424971 851 38 the the DT 10_1101-2020_12_31_424971 851 39 P p NN 10_1101-2020_12_31_424971 851 40 - - HYPH 10_1101-2020_12_31_424971 851 41 site site NN 10_1101-2020_12_31_424971 851 42 1352 1352 CD 10_1101-2020_12_31_424971 851 43 tRNAs trnas CD 10_1101-2020_12_31_424971 851 44 in in IN 10_1101-2020_12_31_424971 851 45 4V9D 4v9d CD 10_1101-2020_12_31_424971 851 46 as as IN 10_1101-2020_12_31_424971 851 47 guides guide NNS 10_1101-2020_12_31_424971 851 48 and and CC 10_1101-2020_12_31_424971 851 49 a a DT 10_1101-2020_12_31_424971 851 50 cartoon cartoon NN 10_1101-2020_12_31_424971 851 51 of of IN 10_1101-2020_12_31_424971 851 52 EF EF NNP 10_1101-2020_12_31_424971 851 53 - - HYPH 10_1101-2020_12_31_424971 851 54 G G NNP 10_1101-2020_12_31_424971 851 55 generated generate VBD 10_1101-2020_12_31_424971 851 56 from from IN 10_1101-2020_12_31_424971 851 57 4V7D 4v7d CD 10_1101-2020_12_31_424971 851 58 was be VBD 10_1101-2020_12_31_424971 851 59 manually manually RB 10_1101-2020_12_31_424971 851 60 positioned position VBN 10_1101-2020_12_31_424971 851 61 1353 1353 CD 10_1101-2020_12_31_424971 851 62 near near IN 10_1101-2020_12_31_424971 851 63 the the DT 10_1101-2020_12_31_424971 851 64 factor factor NN 10_1101-2020_12_31_424971 851 65 binding bind VBG 10_1101-2020_12_31_424971 851 66 site site NN 10_1101-2020_12_31_424971 851 67 of of IN 10_1101-2020_12_31_424971 851 68 the the DT 10_1101-2020_12_31_424971 851 69 ribosomes ribosome NNS 10_1101-2020_12_31_424971 851 70 . . . 10_1101-2020_12_31_424971 852 1 The the DT 10_1101-2020_12_31_424971 852 2 cartoons cartoon NNS 10_1101-2020_12_31_424971 852 3 depicting depict VBG 10_1101-2020_12_31_424971 852 4 PRE PRE NNP 10_1101-2020_12_31_424971 852 5 - - HYPH 10_1101-2020_12_31_424971 852 6 G2 G2 NNP 10_1101-2020_12_31_424971 852 7 , , , 10_1101-2020_12_31_424971 852 8 PRE PRE NNP 10_1101-2020_12_31_424971 852 9 - - HYPH 10_1101-2020_12_31_424971 852 10 G3 G3 NNP 10_1101-2020_12_31_424971 852 11 , , , 10_1101-2020_12_31_424971 852 12 and and CC 10_1101-2020_12_31_424971 852 13 PRE-1354 PRE-1354 NNP 10_1101-2020_12_31_424971 852 14 G4 G4 NNP 10_1101-2020_12_31_424971 852 15 were be VBD 10_1101-2020_12_31_424971 852 16 generated generate VBN 10_1101-2020_12_31_424971 852 17 from from IN 10_1101-2020_12_31_424971 852 18 4V7D 4v7d CD 10_1101-2020_12_31_424971 852 19 , , , 10_1101-2020_12_31_424971 852 20 4W29 4w29 CD 10_1101-2020_12_31_424971 852 21 , , , 10_1101-2020_12_31_424971 852 22 and and CC 10_1101-2020_12_31_424971 852 23 4V5F 4v5f CD 10_1101-2020_12_31_424971 852 24 , , , 10_1101-2020_12_31_424971 852 25 respectively respectively RB 10_1101-2020_12_31_424971 852 26 , , , 10_1101-2020_12_31_424971 852 27 and and CC 10_1101-2020_12_31_424971 852 28 colored color VBD 10_1101-2020_12_31_424971 852 29 as as IN 10_1101-2020_12_31_424971 852 30 in in IN 10_1101-2020_12_31_424971 852 31 Figure Figure NNP 10_1101-2020_12_31_424971 852 32 6 6 CD 10_1101-2020_12_31_424971 852 33 , , , 10_1101-2020_12_31_424971 852 34 with with IN 10_1101-2020_12_31_424971 852 35 1355 1355 CD 10_1101-2020_12_31_424971 852 36 the the DT 10_1101-2020_12_31_424971 852 37 head head NN 10_1101-2020_12_31_424971 852 38 domain domain NN 10_1101-2020_12_31_424971 852 39 shown show VBN 10_1101-2020_12_31_424971 852 40 in in IN 10_1101-2020_12_31_424971 852 41 orange orange NN 10_1101-2020_12_31_424971 852 42 . . . 10_1101-2020_12_31_424971 853 1 1356 1356 CD 10_1101-2020_12_31_424971 853 2 1357 1357 CD 10_1101-2020_12_31_424971 853 3 1358 1358 CD 10_1101-2020_12_31_424971 853 4 1359 1359 CD 10_1101-2020_12_31_424971 853 5 .CC .cc SYM 10_1101-2020_12_31_424971 853 6 - - HYPH 10_1101-2020_12_31_424971 853 7 BY by IN 10_1101-2020_12_31_424971 853 8 - - HYPH 10_1101-2020_12_31_424971 853 9 NC NC NNP 10_1101-2020_12_31_424971 853 10 - - HYPH 10_1101-2020_12_31_424971 853 11 ND ND NNP 10_1101-2020_12_31_424971 853 12 4.0 4.0 CD 10_1101-2020_12_31_424971 853 13 International International NNP 10_1101-2020_12_31_424971 853 14 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 853 15 under under IN 10_1101-2020_12_31_424971 853 16 a a DT 10_1101-2020_12_31_424971 853 17 ( ( -LRB- 10_1101-2020_12_31_424971 853 18 which which WDT 10_1101-2020_12_31_424971 853 19 was be VBD 10_1101-2020_12_31_424971 853 20 not not RB 10_1101-2020_12_31_424971 853 21 certified certify VBN 10_1101-2020_12_31_424971 853 22 by by IN 10_1101-2020_12_31_424971 853 23 peer peer NN 10_1101-2020_12_31_424971 853 24 review review NN 10_1101-2020_12_31_424971 853 25 ) ) -RRB- 10_1101-2020_12_31_424971 853 26 is be VBZ 10_1101-2020_12_31_424971 853 27 the the DT 10_1101-2020_12_31_424971 853 28 author author NN 10_1101-2020_12_31_424971 853 29 / / SYM 10_1101-2020_12_31_424971 853 30 funder funder NN 10_1101-2020_12_31_424971 853 31 , , , 10_1101-2020_12_31_424971 853 32 who who WP 10_1101-2020_12_31_424971 853 33 has have VBZ 10_1101-2020_12_31_424971 853 34 granted grant VBN 10_1101-2020_12_31_424971 853 35 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 853 36 a a DT 10_1101-2020_12_31_424971 853 37 license license NN 10_1101-2020_12_31_424971 853 38 to to TO 10_1101-2020_12_31_424971 853 39 display display VB 10_1101-2020_12_31_424971 853 40 the the DT 10_1101-2020_12_31_424971 853 41 preprint preprint NN 10_1101-2020_12_31_424971 853 42 in in IN 10_1101-2020_12_31_424971 853 43 perpetuity perpetuity NN 10_1101-2020_12_31_424971 853 44 . . . 10_1101-2020_12_31_424971 854 1 It -PRON- PRP 10_1101-2020_12_31_424971 854 2 is be VBZ 10_1101-2020_12_31_424971 854 3 made make VBN 10_1101-2020_12_31_424971 854 4 The the DT 10_1101-2020_12_31_424971 854 5 copyright copyright NN 10_1101-2020_12_31_424971 854 6 holder holder NN 10_1101-2020_12_31_424971 854 7 for for IN 10_1101-2020_12_31_424971 854 8 this this DT 10_1101-2020_12_31_424971 854 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 854 10 version version NN 10_1101-2020_12_31_424971 854 11 posted post VBD 10_1101-2020_12_31_424971 854 12 January January NNP 10_1101-2020_12_31_424971 854 13 2 2 CD 10_1101-2020_12_31_424971 854 14 , , , 10_1101-2020_12_31_424971 854 15 2021 2021 CD 10_1101-2020_12_31_424971 854 16 . . . 10_1101-2020_12_31_424971 854 17 ; ; : 10_1101-2020_12_31_424971 854 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 854 19 : : : 10_1101-2020_12_31_424971 854 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 854 21 preprint preprint NN 10_1101-2020_12_31_424971 854 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 854 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 854 24 53 53 CD 10_1101-2020_12_31_424971 854 25 Figure figure NN 10_1101-2020_12_31_424971 854 26 1 1 CD 10_1101-2020_12_31_424971 854 27 1360 1360 CD 10_1101-2020_12_31_424971 854 28 1361 1361 CD 10_1101-2020_12_31_424971 854 29 1362 1362 CD 10_1101-2020_12_31_424971 854 30 .CC .cc SYM 10_1101-2020_12_31_424971 854 31 - - : 10_1101-2020_12_31_424971 854 32 BY by IN 10_1101-2020_12_31_424971 854 33 - - HYPH 10_1101-2020_12_31_424971 854 34 NC NC NNP 10_1101-2020_12_31_424971 854 35 - - HYPH 10_1101-2020_12_31_424971 854 36 ND ND NNP 10_1101-2020_12_31_424971 854 37 4.0 4.0 CD 10_1101-2020_12_31_424971 854 38 International International NNP 10_1101-2020_12_31_424971 854 39 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 854 40 under under IN 10_1101-2020_12_31_424971 854 41 a a DT 10_1101-2020_12_31_424971 854 42 ( ( -LRB- 10_1101-2020_12_31_424971 854 43 which which WDT 10_1101-2020_12_31_424971 854 44 was be VBD 10_1101-2020_12_31_424971 854 45 not not RB 10_1101-2020_12_31_424971 854 46 certified certify VBN 10_1101-2020_12_31_424971 854 47 by by IN 10_1101-2020_12_31_424971 854 48 peer peer NN 10_1101-2020_12_31_424971 854 49 review review NN 10_1101-2020_12_31_424971 854 50 ) ) -RRB- 10_1101-2020_12_31_424971 854 51 is be VBZ 10_1101-2020_12_31_424971 854 52 the the DT 10_1101-2020_12_31_424971 854 53 author author NN 10_1101-2020_12_31_424971 854 54 / / SYM 10_1101-2020_12_31_424971 854 55 funder funder NN 10_1101-2020_12_31_424971 854 56 , , , 10_1101-2020_12_31_424971 854 57 who who WP 10_1101-2020_12_31_424971 854 58 has have VBZ 10_1101-2020_12_31_424971 854 59 granted grant VBN 10_1101-2020_12_31_424971 854 60 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 854 61 a a DT 10_1101-2020_12_31_424971 854 62 license license NN 10_1101-2020_12_31_424971 854 63 to to TO 10_1101-2020_12_31_424971 854 64 display display VB 10_1101-2020_12_31_424971 854 65 the the DT 10_1101-2020_12_31_424971 854 66 preprint preprint NN 10_1101-2020_12_31_424971 854 67 in in IN 10_1101-2020_12_31_424971 854 68 perpetuity perpetuity NN 10_1101-2020_12_31_424971 854 69 . . . 10_1101-2020_12_31_424971 855 1 It -PRON- PRP 10_1101-2020_12_31_424971 855 2 is be VBZ 10_1101-2020_12_31_424971 855 3 made make VBN 10_1101-2020_12_31_424971 855 4 The the DT 10_1101-2020_12_31_424971 855 5 copyright copyright NN 10_1101-2020_12_31_424971 855 6 holder holder NN 10_1101-2020_12_31_424971 855 7 for for IN 10_1101-2020_12_31_424971 855 8 this this DT 10_1101-2020_12_31_424971 855 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 855 10 version version NN 10_1101-2020_12_31_424971 855 11 posted post VBD 10_1101-2020_12_31_424971 855 12 January January NNP 10_1101-2020_12_31_424971 855 13 2 2 CD 10_1101-2020_12_31_424971 855 14 , , , 10_1101-2020_12_31_424971 855 15 2021 2021 CD 10_1101-2020_12_31_424971 855 16 . . . 10_1101-2020_12_31_424971 855 17 ; ; : 10_1101-2020_12_31_424971 855 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 855 19 : : : 10_1101-2020_12_31_424971 855 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 855 21 preprint preprint NN 10_1101-2020_12_31_424971 855 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 855 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 855 24 54 54 CD 10_1101-2020_12_31_424971 855 25 Figure figure NN 10_1101-2020_12_31_424971 855 26 2 2 CD 10_1101-2020_12_31_424971 855 27 1363 1363 CD 10_1101-2020_12_31_424971 855 28 1364 1364 CD 10_1101-2020_12_31_424971 855 29 1365 1365 CD 10_1101-2020_12_31_424971 855 30 1366 1366 CD 10_1101-2020_12_31_424971 855 31 .CC .cc SYM 10_1101-2020_12_31_424971 855 32 - - HYPH 10_1101-2020_12_31_424971 855 33 BY by IN 10_1101-2020_12_31_424971 855 34 - - HYPH 10_1101-2020_12_31_424971 855 35 NC NC NNP 10_1101-2020_12_31_424971 855 36 - - HYPH 10_1101-2020_12_31_424971 855 37 ND ND NNP 10_1101-2020_12_31_424971 855 38 4.0 4.0 CD 10_1101-2020_12_31_424971 855 39 International International NNP 10_1101-2020_12_31_424971 855 40 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 855 41 under under IN 10_1101-2020_12_31_424971 855 42 a a DT 10_1101-2020_12_31_424971 855 43 ( ( -LRB- 10_1101-2020_12_31_424971 855 44 which which WDT 10_1101-2020_12_31_424971 855 45 was be VBD 10_1101-2020_12_31_424971 855 46 not not RB 10_1101-2020_12_31_424971 855 47 certified certify VBN 10_1101-2020_12_31_424971 855 48 by by IN 10_1101-2020_12_31_424971 855 49 peer peer NN 10_1101-2020_12_31_424971 855 50 review review NN 10_1101-2020_12_31_424971 855 51 ) ) -RRB- 10_1101-2020_12_31_424971 855 52 is be VBZ 10_1101-2020_12_31_424971 855 53 the the DT 10_1101-2020_12_31_424971 855 54 author author NN 10_1101-2020_12_31_424971 855 55 / / SYM 10_1101-2020_12_31_424971 855 56 funder funder NN 10_1101-2020_12_31_424971 855 57 , , , 10_1101-2020_12_31_424971 855 58 who who WP 10_1101-2020_12_31_424971 855 59 has have VBZ 10_1101-2020_12_31_424971 855 60 granted grant VBN 10_1101-2020_12_31_424971 855 61 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 855 62 a a DT 10_1101-2020_12_31_424971 855 63 license license NN 10_1101-2020_12_31_424971 855 64 to to TO 10_1101-2020_12_31_424971 855 65 display display VB 10_1101-2020_12_31_424971 855 66 the the DT 10_1101-2020_12_31_424971 855 67 preprint preprint NN 10_1101-2020_12_31_424971 855 68 in in IN 10_1101-2020_12_31_424971 855 69 perpetuity perpetuity NN 10_1101-2020_12_31_424971 855 70 . . . 10_1101-2020_12_31_424971 856 1 It -PRON- PRP 10_1101-2020_12_31_424971 856 2 is be VBZ 10_1101-2020_12_31_424971 856 3 made make VBN 10_1101-2020_12_31_424971 856 4 The the DT 10_1101-2020_12_31_424971 856 5 copyright copyright NN 10_1101-2020_12_31_424971 856 6 holder holder NN 10_1101-2020_12_31_424971 856 7 for for IN 10_1101-2020_12_31_424971 856 8 this this DT 10_1101-2020_12_31_424971 856 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 856 10 version version NN 10_1101-2020_12_31_424971 856 11 posted post VBD 10_1101-2020_12_31_424971 856 12 January January NNP 10_1101-2020_12_31_424971 856 13 2 2 CD 10_1101-2020_12_31_424971 856 14 , , , 10_1101-2020_12_31_424971 856 15 2021 2021 CD 10_1101-2020_12_31_424971 856 16 . . . 10_1101-2020_12_31_424971 856 17 ; ; : 10_1101-2020_12_31_424971 856 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 856 19 : : : 10_1101-2020_12_31_424971 856 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 856 21 preprint preprint NN 10_1101-2020_12_31_424971 856 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 856 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 856 24 55 55 CD 10_1101-2020_12_31_424971 856 25 Figure figure NN 10_1101-2020_12_31_424971 856 26 3 3 CD 10_1101-2020_12_31_424971 856 27 1367 1367 CD 10_1101-2020_12_31_424971 856 28 1368 1368 CD 10_1101-2020_12_31_424971 856 29 1369 1369 CD 10_1101-2020_12_31_424971 856 30 .CC .cc SYM 10_1101-2020_12_31_424971 856 31 - - : 10_1101-2020_12_31_424971 856 32 BY by IN 10_1101-2020_12_31_424971 856 33 - - HYPH 10_1101-2020_12_31_424971 856 34 NC NC NNP 10_1101-2020_12_31_424971 856 35 - - HYPH 10_1101-2020_12_31_424971 856 36 ND ND NNP 10_1101-2020_12_31_424971 856 37 4.0 4.0 CD 10_1101-2020_12_31_424971 856 38 International International NNP 10_1101-2020_12_31_424971 856 39 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 856 40 under under IN 10_1101-2020_12_31_424971 856 41 a a DT 10_1101-2020_12_31_424971 856 42 ( ( -LRB- 10_1101-2020_12_31_424971 856 43 which which WDT 10_1101-2020_12_31_424971 856 44 was be VBD 10_1101-2020_12_31_424971 856 45 not not RB 10_1101-2020_12_31_424971 856 46 certified certify VBN 10_1101-2020_12_31_424971 856 47 by by IN 10_1101-2020_12_31_424971 856 48 peer peer NN 10_1101-2020_12_31_424971 856 49 review review NN 10_1101-2020_12_31_424971 856 50 ) ) -RRB- 10_1101-2020_12_31_424971 856 51 is be VBZ 10_1101-2020_12_31_424971 856 52 the the DT 10_1101-2020_12_31_424971 856 53 author author NN 10_1101-2020_12_31_424971 856 54 / / SYM 10_1101-2020_12_31_424971 856 55 funder funder NN 10_1101-2020_12_31_424971 856 56 , , , 10_1101-2020_12_31_424971 856 57 who who WP 10_1101-2020_12_31_424971 856 58 has have VBZ 10_1101-2020_12_31_424971 856 59 granted grant VBN 10_1101-2020_12_31_424971 856 60 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 856 61 a a DT 10_1101-2020_12_31_424971 856 62 license license NN 10_1101-2020_12_31_424971 856 63 to to TO 10_1101-2020_12_31_424971 856 64 display display VB 10_1101-2020_12_31_424971 856 65 the the DT 10_1101-2020_12_31_424971 856 66 preprint preprint NN 10_1101-2020_12_31_424971 856 67 in in IN 10_1101-2020_12_31_424971 856 68 perpetuity perpetuity NN 10_1101-2020_12_31_424971 856 69 . . . 10_1101-2020_12_31_424971 857 1 It -PRON- PRP 10_1101-2020_12_31_424971 857 2 is be VBZ 10_1101-2020_12_31_424971 857 3 made make VBN 10_1101-2020_12_31_424971 857 4 The the DT 10_1101-2020_12_31_424971 857 5 copyright copyright NN 10_1101-2020_12_31_424971 857 6 holder holder NN 10_1101-2020_12_31_424971 857 7 for for IN 10_1101-2020_12_31_424971 857 8 this this DT 10_1101-2020_12_31_424971 857 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 857 10 version version NN 10_1101-2020_12_31_424971 857 11 posted post VBD 10_1101-2020_12_31_424971 857 12 January January NNP 10_1101-2020_12_31_424971 857 13 2 2 CD 10_1101-2020_12_31_424971 857 14 , , , 10_1101-2020_12_31_424971 857 15 2021 2021 CD 10_1101-2020_12_31_424971 857 16 . . . 10_1101-2020_12_31_424971 857 17 ; ; : 10_1101-2020_12_31_424971 857 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 857 19 : : : 10_1101-2020_12_31_424971 857 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 857 21 preprint preprint NN 10_1101-2020_12_31_424971 857 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 857 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 857 24 56 56 CD 10_1101-2020_12_31_424971 857 25 Figure figure NN 10_1101-2020_12_31_424971 857 26 4 4 CD 10_1101-2020_12_31_424971 857 27 1370 1370 CD 10_1101-2020_12_31_424971 857 28 1371 1371 CD 10_1101-2020_12_31_424971 857 29 .CC .cc SYM 10_1101-2020_12_31_424971 857 30 - - HYPH 10_1101-2020_12_31_424971 857 31 BY by IN 10_1101-2020_12_31_424971 857 32 - - HYPH 10_1101-2020_12_31_424971 857 33 NC NC NNP 10_1101-2020_12_31_424971 857 34 - - HYPH 10_1101-2020_12_31_424971 857 35 ND ND NNP 10_1101-2020_12_31_424971 857 36 4.0 4.0 CD 10_1101-2020_12_31_424971 857 37 International International NNP 10_1101-2020_12_31_424971 857 38 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 857 39 under under IN 10_1101-2020_12_31_424971 857 40 a a DT 10_1101-2020_12_31_424971 857 41 ( ( -LRB- 10_1101-2020_12_31_424971 857 42 which which WDT 10_1101-2020_12_31_424971 857 43 was be VBD 10_1101-2020_12_31_424971 857 44 not not RB 10_1101-2020_12_31_424971 857 45 certified certify VBN 10_1101-2020_12_31_424971 857 46 by by IN 10_1101-2020_12_31_424971 857 47 peer peer NN 10_1101-2020_12_31_424971 857 48 review review NN 10_1101-2020_12_31_424971 857 49 ) ) -RRB- 10_1101-2020_12_31_424971 857 50 is be VBZ 10_1101-2020_12_31_424971 857 51 the the DT 10_1101-2020_12_31_424971 857 52 author author NN 10_1101-2020_12_31_424971 857 53 / / SYM 10_1101-2020_12_31_424971 857 54 funder funder NN 10_1101-2020_12_31_424971 857 55 , , , 10_1101-2020_12_31_424971 857 56 who who WP 10_1101-2020_12_31_424971 857 57 has have VBZ 10_1101-2020_12_31_424971 857 58 granted grant VBN 10_1101-2020_12_31_424971 857 59 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 857 60 a a DT 10_1101-2020_12_31_424971 857 61 license license NN 10_1101-2020_12_31_424971 857 62 to to TO 10_1101-2020_12_31_424971 857 63 display display VB 10_1101-2020_12_31_424971 857 64 the the DT 10_1101-2020_12_31_424971 857 65 preprint preprint NN 10_1101-2020_12_31_424971 857 66 in in IN 10_1101-2020_12_31_424971 857 67 perpetuity perpetuity NN 10_1101-2020_12_31_424971 857 68 . . . 10_1101-2020_12_31_424971 858 1 It -PRON- PRP 10_1101-2020_12_31_424971 858 2 is be VBZ 10_1101-2020_12_31_424971 858 3 made make VBN 10_1101-2020_12_31_424971 858 4 The the DT 10_1101-2020_12_31_424971 858 5 copyright copyright NN 10_1101-2020_12_31_424971 858 6 holder holder NN 10_1101-2020_12_31_424971 858 7 for for IN 10_1101-2020_12_31_424971 858 8 this this DT 10_1101-2020_12_31_424971 858 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 858 10 version version NN 10_1101-2020_12_31_424971 858 11 posted post VBD 10_1101-2020_12_31_424971 858 12 January January NNP 10_1101-2020_12_31_424971 858 13 2 2 CD 10_1101-2020_12_31_424971 858 14 , , , 10_1101-2020_12_31_424971 858 15 2021 2021 CD 10_1101-2020_12_31_424971 858 16 . . . 10_1101-2020_12_31_424971 858 17 ; ; : 10_1101-2020_12_31_424971 858 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 858 19 : : : 10_1101-2020_12_31_424971 858 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 858 21 preprint preprint NN 10_1101-2020_12_31_424971 858 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 858 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 858 24 57 57 CD 10_1101-2020_12_31_424971 858 25 Figure figure NN 10_1101-2020_12_31_424971 858 26 5 5 CD 10_1101-2020_12_31_424971 858 27 1372 1372 CD 10_1101-2020_12_31_424971 858 28 1373 1373 CD 10_1101-2020_12_31_424971 858 29 1374 1374 CD 10_1101-2020_12_31_424971 858 30 .CC .CC NFP 10_1101-2020_12_31_424971 858 31 - - : 10_1101-2020_12_31_424971 858 32 BY by IN 10_1101-2020_12_31_424971 858 33 - - HYPH 10_1101-2020_12_31_424971 858 34 NC NC NNP 10_1101-2020_12_31_424971 858 35 - - HYPH 10_1101-2020_12_31_424971 858 36 ND ND NNP 10_1101-2020_12_31_424971 858 37 4.0 4.0 CD 10_1101-2020_12_31_424971 858 38 International International NNP 10_1101-2020_12_31_424971 858 39 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 858 40 under under IN 10_1101-2020_12_31_424971 858 41 a a DT 10_1101-2020_12_31_424971 858 42 ( ( -LRB- 10_1101-2020_12_31_424971 858 43 which which WDT 10_1101-2020_12_31_424971 858 44 was be VBD 10_1101-2020_12_31_424971 858 45 not not RB 10_1101-2020_12_31_424971 858 46 certified certify VBN 10_1101-2020_12_31_424971 858 47 by by IN 10_1101-2020_12_31_424971 858 48 peer peer NN 10_1101-2020_12_31_424971 858 49 review review NN 10_1101-2020_12_31_424971 858 50 ) ) -RRB- 10_1101-2020_12_31_424971 858 51 is be VBZ 10_1101-2020_12_31_424971 858 52 the the DT 10_1101-2020_12_31_424971 858 53 author author NN 10_1101-2020_12_31_424971 858 54 / / SYM 10_1101-2020_12_31_424971 858 55 funder funder NN 10_1101-2020_12_31_424971 858 56 , , , 10_1101-2020_12_31_424971 858 57 who who WP 10_1101-2020_12_31_424971 858 58 has have VBZ 10_1101-2020_12_31_424971 858 59 granted grant VBN 10_1101-2020_12_31_424971 858 60 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 858 61 a a DT 10_1101-2020_12_31_424971 858 62 license license NN 10_1101-2020_12_31_424971 858 63 to to TO 10_1101-2020_12_31_424971 858 64 display display VB 10_1101-2020_12_31_424971 858 65 the the DT 10_1101-2020_12_31_424971 858 66 preprint preprint NN 10_1101-2020_12_31_424971 858 67 in in IN 10_1101-2020_12_31_424971 858 68 perpetuity perpetuity NN 10_1101-2020_12_31_424971 858 69 . . . 10_1101-2020_12_31_424971 859 1 It -PRON- PRP 10_1101-2020_12_31_424971 859 2 is be VBZ 10_1101-2020_12_31_424971 859 3 made make VBN 10_1101-2020_12_31_424971 859 4 The the DT 10_1101-2020_12_31_424971 859 5 copyright copyright NN 10_1101-2020_12_31_424971 859 6 holder holder NN 10_1101-2020_12_31_424971 859 7 for for IN 10_1101-2020_12_31_424971 859 8 this this DT 10_1101-2020_12_31_424971 859 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 859 10 version version NN 10_1101-2020_12_31_424971 859 11 posted post VBD 10_1101-2020_12_31_424971 859 12 January January NNP 10_1101-2020_12_31_424971 859 13 2 2 CD 10_1101-2020_12_31_424971 859 14 , , , 10_1101-2020_12_31_424971 859 15 2021 2021 CD 10_1101-2020_12_31_424971 859 16 . . . 10_1101-2020_12_31_424971 859 17 ; ; : 10_1101-2020_12_31_424971 859 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 859 19 : : : 10_1101-2020_12_31_424971 859 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 859 21 preprint preprint NN 10_1101-2020_12_31_424971 859 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 859 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN 10_1101-2020_12_31_424971 859 24 58 58 CD 10_1101-2020_12_31_424971 859 25 Figure figure NN 10_1101-2020_12_31_424971 859 26 6 6 CD 10_1101-2020_12_31_424971 859 27 1375 1375 CD 10_1101-2020_12_31_424971 859 28 1376 1376 CD 10_1101-2020_12_31_424971 859 29 1377 1377 CD 10_1101-2020_12_31_424971 859 30 P p NN 10_1101-2020_12_31_424971 859 31 A a DT 10_1101-2020_12_31_424971 859 32 1.00 1.00 CD 10_1101-2020_12_31_424971 859 33 0.50 0.50 CD 10_1101-2020_12_31_424971 859 34 0.00 0.00 CD 10_1101-2020_12_31_424971 859 35 E e NN 10_1101-2020_12_31_424971 859 36 F f NN 10_1101-2020_12_31_424971 859 37 R r NN 10_1101-2020_12_31_424971 859 38 E e NN 10_1101-2020_12_31_424971 859 39 T t NN 10_1101-2020_12_31_424971 859 40 0 0 CD 10_1101-2020_12_31_424971 859 41 10 10 CD 10_1101-2020_12_31_424971 859 42 20 20 CD 10_1101-2020_12_31_424971 859 43 30 30 CD 10_1101-2020_12_31_424971 859 44 0 0 CD 10_1101-2020_12_31_424971 859 45 10 10 CD 10_1101-2020_12_31_424971 859 46 1 1 CD 10_1101-2020_12_31_424971 859 47 Time Time NNP 10_1101-2020_12_31_424971 859 48 ( ( -LRB- 10_1101-2020_12_31_424971 859 49 s s NNP 10_1101-2020_12_31_424971 859 50 ) ) -RRB- 10_1101-2020_12_31_424971 859 51 Time Time NNP 10_1101-2020_12_31_424971 859 52 ( ( -LRB- 10_1101-2020_12_31_424971 859 53 s s NNP 10_1101-2020_12_31_424971 859 54 ) ) -RRB- 10_1101-2020_12_31_424971 859 55 ProL ProL NNP 10_1101-2020_12_31_424971 859 56 - - HYPH 10_1101-2020_12_31_424971 859 57 TC TC NNP 10_1101-2020_12_31_424971 859 58 ProL ProL NNP 10_1101-2020_12_31_424971 859 59 - - HYPH 10_1101-2020_12_31_424971 859 60 TC TC NNP 10_1101-2020_12_31_424971 859 61 + + CC 10_1101-2020_12_31_424971 859 62 EF EF NNP 10_1101-2020_12_31_424971 859 63 - - HYPH 10_1101-2020_12_31_424971 859 64 G G NNP 10_1101-2020_12_31_424971 859 65 Deliver Deliver NNP 10_1101-2020_12_31_424971 859 66 TCsat TCsat NNP 10_1101-2020_12_31_424971 859 67 5 5 CD 10_1101-2020_12_31_424971 859 68 s s NN 10_1101-2020_12_31_424971 859 69 Deliver deliver VB 10_1101-2020_12_31_424971 859 70 TCs+EF tcs+ef NN 10_1101-2020_12_31_424971 859 71 - - HYPH 10_1101-2020_12_31_424971 859 72 Gat Gat NNP 10_1101-2020_12_31_424971 859 73 5 5 CD 10_1101-2020_12_31_424971 859 74 s s NNP 10_1101-2020_12_31_424971 859 75 N= N= NNP 10_1101-2020_12_31_424971 859 76 k70S k70S NNP 10_1101-2020_12_31_424971 859 77 IC→GS IC→GS NNP 10_1101-2020_12_31_424971 859 78 kGS1→GS kGS1→GS NNP 10_1101-2020_12_31_424971 859 79 a a DT 10_1101-2020_12_31_424971 859 80 kGS kg NNS 10_1101-2020_12_31_424971 859 81 →GS1 →gs1 CD 10_1101-2020_12_31_424971 859 82 A a NN 10_1101-2020_12_31_424971 859 83 P. p. NN 10_1101-2020_12_31_424971 859 84 . . . 10_1101-2020_12_31_424971 860 1 Time Time NNP 10_1101-2020_12_31_424971 860 2 e e NNP 10_1101-2020_12_31_424971 860 3 Time time NN 10_1101-2020_12_31_424971 860 4 0.55 0.55 CD 10_1101-2020_12_31_424971 860 5 ( ( -LRB- 10_1101-2020_12_31_424971 860 6 1 1 CD 10_1101-2020_12_31_424971 860 7 e e NN 10_1101-2020_12_31_424971 860 8 ) ) -RRB- 10_1101-2020_12_31_424971 860 9 0 0 NFP 10_1101-2020_12_31_424971 860 10 . . . 10_1101-2020_12_31_424971 861 1 1 1 CD 10_1101-2020_12_31_424971 861 2 ( ( -LRB- 10_1101-2020_12_31_424971 861 3 1 1 CD 10_1101-2020_12_31_424971 861 4 l l NN 10_1101-2020_12_31_424971 861 5 se se NN 10_1101-2020_12_31_424971 861 6 ) ) -RRB- 10_1101-2020_12_31_424971 861 7 0 0 NFP 10_1101-2020_12_31_424971 861 8 . . . 10_1101-2020_12_31_424971 862 1 0.7 0.7 CD 10_1101-2020_12_31_424971 862 2 15 15 CD 10_1101-2020_12_31_424971 862 3 15 15 CD 10_1101-2020_12_31_424971 862 4 10 10 CD 10_1101-2020_12_31_424971 862 5 10 10 CD 10_1101-2020_12_31_424971 862 6 N= n= NN 10_1101-2020_12_31_424971 862 7 5 5 CD 10_1101-2020_12_31_424971 862 8 1 1 CD 10_1101-2020_12_31_424971 862 9 ProL ProL NNP 10_1101-2020_12_31_424971 862 10 GS GS NNP 10_1101-2020_12_31_424971 862 11 P P NNP 10_1101-2020_12_31_424971 862 12 E1 e1 NN 10_1101-2020_12_31_424971 862 13 Stal Stal NNP 10_1101-2020_12_31_424971 862 14 P P NNP 10_1101-2020_12_31_424971 862 15 ST ST NNP 10_1101-2020_12_31_424971 862 16 GS1 GS1 NNP 10_1101-2020_12_31_424971 862 17 ( ( -LRB- 10_1101-2020_12_31_424971 862 18 EF EF NNP 10_1101-2020_12_31_424971 862 19 - - HYPH 10_1101-2020_12_31_424971 862 20 G G NNP 10_1101-2020_12_31_424971 862 21 ) ) -RRB- 10_1101-2020_12_31_424971 862 22 GS GS NNP 10_1101-2020_12_31_424971 862 23 ( ( -LRB- 10_1101-2020_12_31_424971 862 24 EF EF NNP 10_1101-2020_12_31_424971 862 25 - - HYPH 10_1101-2020_12_31_424971 862 26 G G NNP 10_1101-2020_12_31_424971 862 27 ) ) -RRB- 10_1101-2020_12_31_424971 862 28 E E NNP 10_1101-2020_12_31_424971 862 29 GS GS NNP 10_1101-2020_12_31_424971 862 30 GS1 GS1 NNP 10_1101-2020_12_31_424971 862 31 E E NNP 10_1101-2020_12_31_424971 862 32 F f NN 10_1101-2020_12_31_424971 862 33 R r NN 10_1101-2020_12_31_424971 862 34 E e NN 10_1101-2020_12_31_424971 862 35 T t NN 10_1101-2020_12_31_424971 862 36 k70S k70 NNS 10_1101-2020_12_31_424971 862 37 IC→GS ic→gs CC 10_1101-2020_12_31_424971 862 38 kGS kg NNS 10_1101-2020_12_31_424971 862 39 →P →P -LRB- 10_1101-2020_12_31_424971 862 40 ST ST NNP 10_1101-2020_12_31_424971 862 41 0 0 NNP 10_1101-2020_12_31_424971 862 42 5 5 CD 10_1101-2020_12_31_424971 862 43 Time Time NNP 10_1101-2020_12_31_424971 862 44 ( ( -LRB- 10_1101-2020_12_31_424971 862 45 s s NN 10_1101-2020_12_31_424971 862 46 ) ) -RRB- 10_1101-2020_12_31_424971 862 47 7.5.5 7.5.5 CD 10_1101-2020_12_31_424971 862 48 E e NN 10_1101-2020_12_31_424971 862 49 F f NN 10_1101-2020_12_31_424971 862 50 R r NN 10_1101-2020_12_31_424971 862 51 E e NN 10_1101-2020_12_31_424971 862 52 T t NN 10_1101-2020_12_31_424971 862 53 N n NN 10_1101-2020_12_31_424971 862 54 = = SYM 10_1101-2020_12_31_424971 862 55 N= N= NNP 10_1101-2020_12_31_424971 862 56 N= N= NNP 10_1101-2020_12_31_424971 862 57 N= N= NNP 10_1101-2020_12_31_424971 862 58 5 5 CD 10_1101-2020_12_31_424971 862 59 Suf Suf NNP 10_1101-2020_12_31_424971 862 60 N=1 N=1 NNP 10_1101-2020_12_31_424971 862 61 N= N= NNP 10_1101-2020_12_31_424971 862 62 7 7 CD 10_1101-2020_12_31_424971 862 63 N= n= NN 10_1101-2020_12_31_424971 862 64 1 1 CD 10_1101-2020_12_31_424971 862 65 ProL ProL NNP 10_1101-2020_12_31_424971 862 66 0 0 CD 10_1101-2020_12_31_424971 862 67 . . . 10_1101-2020_12_31_424971 863 1 0.0 0.0 CD 10_1101-2020_12_31_424971 863 2 0 0 CD 10_1101-2020_12_31_424971 863 3 . . . 10_1101-2020_12_31_424971 864 1 0 0 NFP 10_1101-2020_12_31_424971 864 2 . . . 10_1101-2020_12_31_424971 865 1 0 0 NFP 10_1101-2020_12_31_424971 865 2 . . . 10_1101-2020_12_31_424971 866 1 0 0 NFP 10_1101-2020_12_31_424971 866 2 . . . 10_1101-2020_12_31_424971 867 1 1.0 1.0 CD 10_1101-2020_12_31_424971 867 2 1 1 CD 10_1101-2020_12_31_424971 867 3 . . . 10_1101-2020_12_31_424971 868 1 E E NNP 10_1101-2020_12_31_424971 868 2 F F NNP 10_1101-2020_12_31_424971 868 3 R r NN 10_1101-2020_12_31_424971 868 4 E e NN 10_1101-2020_12_31_424971 868 5 T t NN 10_1101-2020_12_31_424971 868 6 Suf suf NN 10_1101-2020_12_31_424971 868 7 0 0 CD 10_1101-2020_12_31_424971 868 8 . . . 10_1101-2020_12_31_424971 869 1 0.0 0.0 CD 10_1101-2020_12_31_424971 869 2 0 0 CD 10_1101-2020_12_31_424971 869 3 . . . 10_1101-2020_12_31_424971 870 1 0 0 NFP 10_1101-2020_12_31_424971 870 2 . . . 10_1101-2020_12_31_424971 871 1 0 0 NFP 10_1101-2020_12_31_424971 871 2 . . . 10_1101-2020_12_31_424971 872 1 0 0 NFP 10_1101-2020_12_31_424971 872 2 . . . 10_1101-2020_12_31_424971 873 1 1.0 1.0 CD 10_1101-2020_12_31_424971 873 2 1 1 CD 10_1101-2020_12_31_424971 873 3 . . . 10_1101-2020_12_31_424971 874 1 N=1 N=1 NNP 10_1101-2020_12_31_424971 874 2 ProL ProL NNP 10_1101-2020_12_31_424971 874 3 70S 70S NNP 10_1101-2020_12_31_424971 874 4 IC IC NNP 10_1101-2020_12_31_424971 874 5 TC TC NNP 10_1101-2020_12_31_424971 874 6 EF EF NNP 10_1101-2020_12_31_424971 874 7 - - HYPH 10_1101-2020_12_31_424971 874 8 G G NNP 10_1101-2020_12_31_424971 874 9 Pm Pm NNP 10_1101-2020_12_31_424971 874 10 N=1 N=1 NNP 10_1101-2020_12_31_424971 874 11 Time Time NNP 10_1101-2020_12_31_424971 874 12 ( ( -LRB- 10_1101-2020_12_31_424971 874 13 mi mi NNP 10_1101-2020_12_31_424971 874 14 ) ) -RRB- 10_1101-2020_12_31_424971 874 15 Time Time NNP 10_1101-2020_12_31_424971 874 16 ( ( -LRB- 10_1101-2020_12_31_424971 874 17 mi mi NNP 10_1101-2020_12_31_424971 874 18 ) ) -RRB- 10_1101-2020_12_31_424971 874 19 .5 .5 . 10_1101-2020_12_31_424971 874 20 GS1 GS1 NNP 10_1101-2020_12_31_424971 874 21 0 0 CD 10_1101-2020_12_31_424971 874 22 . . . 10_1101-2020_12_31_424971 875 1 5 5 CD 10_1101-2020_12_31_424971 875 2 0 0 CD 10_1101-2020_12_31_424971 875 3 . . . 10_1101-2020_12_31_424971 876 1 0 0 CD 10_1101-2020_12_31_424971 876 2 5 5 CD 10_1101-2020_12_31_424971 876 3 107.5 107.5 CD 10_1101-2020_12_31_424971 876 4 .5 .5 NN 10_1101-2020_12_31_424971 876 5 0 0 CD 10_1101-2020_12_31_424971 876 6 5 5 CD 10_1101-2020_12_31_424971 876 7 Time Time NNP 10_1101-2020_12_31_424971 876 8 ( ( -LRB- 10_1101-2020_12_31_424971 876 9 s s NN 10_1101-2020_12_31_424971 876 10 ) ) -RRB- 10_1101-2020_12_31_424971 876 11 10 10 CD 10_1101-2020_12_31_424971 876 12 Time Time NNP 10_1101-2020_12_31_424971 876 13 ( ( -LRB- 10_1101-2020_12_31_424971 876 14 s s NNP 10_1101-2020_12_31_424971 876 15 ) ) -RRB- 10_1101-2020_12_31_424971 876 16 .5 .5 . 10_1101-2020_12_31_424971 876 17 0 0 NFP 10_1101-2020_12_31_424971 876 18 5 5 CD 10_1101-2020_12_31_424971 876 19 Time Time NNP 10_1101-2020_12_31_424971 876 20 ( ( -LRB- 10_1101-2020_12_31_424971 876 21 s s NN 10_1101-2020_12_31_424971 876 22 ) ) -RRB- 10_1101-2020_12_31_424971 876 23 7.5 7.5 CD 10_1101-2020_12_31_424971 876 24 a a DT 10_1101-2020_12_31_424971 876 25 0 0 CD 10_1101-2020_12_31_424971 876 26 1 1 CD 10_1101-2020_12_31_424971 876 27 0 0 CD 10_1101-2020_12_31_424971 876 28 1 1 CD 10_1101-2020_12_31_424971 876 29 10 10 CD 10_1101-2020_12_31_424971 876 30 N=17 n=17 NN 10_1101-2020_12_31_424971 876 31 0 0 CD 10_1101-2020_12_31_424971 876 32 0 0 CD 10_1101-2020_12_31_424971 876 33 N=55 N=55 NNP 10_1101-2020_12_31_424971 876 34 Suf Suf NNP 10_1101-2020_12_31_424971 876 35 0 0 CD 10_1101-2020_12_31_424971 876 36 . . . 10_1101-2020_12_31_424971 877 1 0 0 NFP 10_1101-2020_12_31_424971 877 2 . . . 10_1101-2020_12_31_424971 878 1 0 0 NFP 10_1101-2020_12_31_424971 878 2 5 5 CD 10_1101-2020_12_31_424971 878 3 Time Time NNP 10_1101-2020_12_31_424971 878 4 ( ( -LRB- 10_1101-2020_12_31_424971 878 5 s s NN 10_1101-2020_12_31_424971 878 6 ) ) -RRB- 10_1101-2020_12_31_424971 878 7 10 10 CD 10_1101-2020_12_31_424971 878 8 0 0 CD 10_1101-2020_12_31_424971 878 9 5 5 CD 10_1101-2020_12_31_424971 878 10 Time Time NNP 10_1101-2020_12_31_424971 878 11 ( ( -LRB- 10_1101-2020_12_31_424971 878 12 s s NN 10_1101-2020_12_31_424971 878 13 ) ) -RRB- 10_1101-2020_12_31_424971 878 14 7.5.5 7.5.5 CD 10_1101-2020_12_31_424971 878 15 0 0 CD 10_1101-2020_12_31_424971 878 16 5 5 CD 10_1101-2020_12_31_424971 878 17 Time Time NNP 10_1101-2020_12_31_424971 878 18 ( ( -LRB- 10_1101-2020_12_31_424971 878 19 s s NN 10_1101-2020_12_31_424971 878 20 ) ) -RRB- 10_1101-2020_12_31_424971 878 21 7.5.5 7.5.5 CD 10_1101-2020_12_31_424971 878 22 0 0 CD 10_1101-2020_12_31_424971 878 23 5 5 CD 10_1101-2020_12_31_424971 878 24 Time Time NNP 10_1101-2020_12_31_424971 878 25 ( ( -LRB- 10_1101-2020_12_31_424971 878 26 s s NN 10_1101-2020_12_31_424971 878 27 ) ) -RRB- 10_1101-2020_12_31_424971 878 28 7.5 7.5 CD 10_1101-2020_12_31_424971 878 29 .CC .CC NFP 10_1101-2020_12_31_424971 878 30 - - HYPH 10_1101-2020_12_31_424971 878 31 BY by IN 10_1101-2020_12_31_424971 878 32 - - HYPH 10_1101-2020_12_31_424971 878 33 NC NC NNP 10_1101-2020_12_31_424971 878 34 - - HYPH 10_1101-2020_12_31_424971 878 35 ND ND NNP 10_1101-2020_12_31_424971 878 36 4.0 4.0 CD 10_1101-2020_12_31_424971 878 37 International International NNP 10_1101-2020_12_31_424971 878 38 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 878 39 under under IN 10_1101-2020_12_31_424971 878 40 a a DT 10_1101-2020_12_31_424971 878 41 ( ( -LRB- 10_1101-2020_12_31_424971 878 42 which which WDT 10_1101-2020_12_31_424971 878 43 was be VBD 10_1101-2020_12_31_424971 878 44 not not RB 10_1101-2020_12_31_424971 878 45 certified certify VBN 10_1101-2020_12_31_424971 878 46 by by IN 10_1101-2020_12_31_424971 878 47 peer peer NN 10_1101-2020_12_31_424971 878 48 review review NN 10_1101-2020_12_31_424971 878 49 ) ) -RRB- 10_1101-2020_12_31_424971 878 50 is be VBZ 10_1101-2020_12_31_424971 878 51 the the DT 10_1101-2020_12_31_424971 878 52 author author NN 10_1101-2020_12_31_424971 878 53 / / SYM 10_1101-2020_12_31_424971 878 54 funder funder NN 10_1101-2020_12_31_424971 878 55 , , , 10_1101-2020_12_31_424971 878 56 who who WP 10_1101-2020_12_31_424971 878 57 has have VBZ 10_1101-2020_12_31_424971 878 58 granted grant VBN 10_1101-2020_12_31_424971 878 59 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 878 60 a a DT 10_1101-2020_12_31_424971 878 61 license license NN 10_1101-2020_12_31_424971 878 62 to to TO 10_1101-2020_12_31_424971 878 63 display display VB 10_1101-2020_12_31_424971 878 64 the the DT 10_1101-2020_12_31_424971 878 65 preprint preprint NN 10_1101-2020_12_31_424971 878 66 in in IN 10_1101-2020_12_31_424971 878 67 perpetuity perpetuity NN 10_1101-2020_12_31_424971 878 68 . . . 10_1101-2020_12_31_424971 879 1 It -PRON- PRP 10_1101-2020_12_31_424971 879 2 is be VBZ 10_1101-2020_12_31_424971 879 3 made make VBN 10_1101-2020_12_31_424971 879 4 The the DT 10_1101-2020_12_31_424971 879 5 copyright copyright NN 10_1101-2020_12_31_424971 879 6 holder holder NN 10_1101-2020_12_31_424971 879 7 for for IN 10_1101-2020_12_31_424971 879 8 this this DT 10_1101-2020_12_31_424971 879 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 879 10 version version NN 10_1101-2020_12_31_424971 879 11 posted post VBD 10_1101-2020_12_31_424971 879 12 January January NNP 10_1101-2020_12_31_424971 879 13 2 2 CD 10_1101-2020_12_31_424971 879 14 , , , 10_1101-2020_12_31_424971 879 15 2021 2021 CD 10_1101-2020_12_31_424971 879 16 . . . 10_1101-2020_12_31_424971 879 17 ; ; : 10_1101-2020_12_31_424971 879 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 879 19 : : : 10_1101-2020_12_31_424971 879 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 879 21 preprint preprint NN 10_1101-2020_12_31_424971 879 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 879 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ CD 10_1101-2020_12_31_424971 879 24 59 59 CD 10_1101-2020_12_31_424971 879 25 Figure figure NN 10_1101-2020_12_31_424971 879 26 7 7 CD 10_1101-2020_12_31_424971 879 27 1378 1378 CD 10_1101-2020_12_31_424971 879 28 1379 1379 CD 10_1101-2020_12_31_424971 879 29 1380 1380 CD 10_1101-2020_12_31_424971 879 30 1381 1381 CD 10_1101-2020_12_31_424971 879 31 60 60 CD 10_1101-2020_12_31_424971 879 32 Å Å NNP 10_1101-2020_12_31_424971 879 33 POST POST NNP 10_1101-2020_12_31_424971 879 34 PRE PRE NNP 10_1101-2020_12_31_424971 879 35 PRE PRE NNP 10_1101-2020_12_31_424971 879 36 - - HYPH 10_1101-2020_12_31_424971 879 37 G1 G1 NNP 10_1101-2020_12_31_424971 879 38 PRE PRE NNP 10_1101-2020_12_31_424971 879 39 - - HYPH 10_1101-2020_12_31_424971 879 40 G2 G2 NNP 10_1101-2020_12_31_424971 879 41 PRE PRE NNP 10_1101-2020_12_31_424971 879 42 - - HYPH 10_1101-2020_12_31_424971 879 43 G3 G3 NNP 10_1101-2020_12_31_424971 879 44 PRE PRE NNP 10_1101-2020_12_31_424971 879 45 - - HYPH 10_1101-2020_12_31_424971 879 46 G4 G4 NNP 10_1101-2020_12_31_424971 879 47 POST POST NNP 10_1101-2020_12_31_424971 879 48 GS1 GS1 NNP 10_1101-2020_12_31_424971 879 49 GS1 GS1 NNP 10_1101-2020_12_31_424971 879 50 GS2 GS2 NNP 10_1101-2020_12_31_424971 879 51 GS2 GS2 NNP 10_1101-2020_12_31_424971 879 52 60 60 CD 10_1101-2020_12_31_424971 879 53 Å Å NNP 10_1101-2020_12_31_424971 879 54 Intersubunit intersubunit JJ 10_1101-2020_12_31_424971 879 55 rotation rotation NN 10_1101-2020_12_31_424971 879 56 8 8 CD 10_1101-2020_12_31_424971 879 57 ° ° , 10_1101-2020_12_31_424971 879 58 8 8 CD 10_1101-2020_12_31_424971 879 59 ° ° , 10_1101-2020_12_31_424971 879 60 12 12 CD 10_1101-2020_12_31_424971 879 61 ° ° , 10_1101-2020_12_31_424971 879 62 2 2 CD 10_1101-2020_12_31_424971 879 63 ° ° , 10_1101-2020_12_31_424971 879 64 0 0 CD 10_1101-2020_12_31_424971 879 65 ° ° , 10_1101-2020_12_31_424971 879 66 Head Head NNP 10_1101-2020_12_31_424971 879 67 swive swive JJ 10_1101-2020_12_31_424971 879 68 in in IN 10_1101-2020_12_31_424971 879 69 - - HYPH 10_1101-2020_12_31_424971 879 70 ° ° NN 10_1101-2020_12_31_424971 879 71 - - HYPH 10_1101-2020_12_31_424971 879 72 ° ° NNP 10_1101-2020_12_31_424971 879 73 3 3 CD 10_1101-2020_12_31_424971 879 74 ° ° , 10_1101-2020_12_31_424971 879 75 21 21 CD 10_1101-2020_12_31_424971 879 76 ° ° NNS 10_1101-2020_12_31_424971 879 77 0 0 CD 10_1101-2020_12_31_424971 879 78 ° ° NNS 10_1101-2020_12_31_424971 879 79 .CC .CC NFP 10_1101-2020_12_31_424971 879 80 - - HYPH 10_1101-2020_12_31_424971 879 81 BY by IN 10_1101-2020_12_31_424971 879 82 - - HYPH 10_1101-2020_12_31_424971 879 83 NC NC NNP 10_1101-2020_12_31_424971 879 84 - - HYPH 10_1101-2020_12_31_424971 879 85 ND ND NNP 10_1101-2020_12_31_424971 879 86 4.0 4.0 CD 10_1101-2020_12_31_424971 879 87 International International NNP 10_1101-2020_12_31_424971 879 88 licenseavailable licenseavailable NN 10_1101-2020_12_31_424971 879 89 under under IN 10_1101-2020_12_31_424971 879 90 a a DT 10_1101-2020_12_31_424971 879 91 ( ( -LRB- 10_1101-2020_12_31_424971 879 92 which which WDT 10_1101-2020_12_31_424971 879 93 was be VBD 10_1101-2020_12_31_424971 879 94 not not RB 10_1101-2020_12_31_424971 879 95 certified certify VBN 10_1101-2020_12_31_424971 879 96 by by IN 10_1101-2020_12_31_424971 879 97 peer peer NN 10_1101-2020_12_31_424971 879 98 review review NN 10_1101-2020_12_31_424971 879 99 ) ) -RRB- 10_1101-2020_12_31_424971 879 100 is be VBZ 10_1101-2020_12_31_424971 879 101 the the DT 10_1101-2020_12_31_424971 879 102 author author NN 10_1101-2020_12_31_424971 879 103 / / SYM 10_1101-2020_12_31_424971 879 104 funder funder NN 10_1101-2020_12_31_424971 879 105 , , , 10_1101-2020_12_31_424971 879 106 who who WP 10_1101-2020_12_31_424971 879 107 has have VBZ 10_1101-2020_12_31_424971 879 108 granted grant VBN 10_1101-2020_12_31_424971 879 109 bioRxiv biorxiv IN 10_1101-2020_12_31_424971 879 110 a a DT 10_1101-2020_12_31_424971 879 111 license license NN 10_1101-2020_12_31_424971 879 112 to to TO 10_1101-2020_12_31_424971 879 113 display display VB 10_1101-2020_12_31_424971 879 114 the the DT 10_1101-2020_12_31_424971 879 115 preprint preprint NN 10_1101-2020_12_31_424971 879 116 in in IN 10_1101-2020_12_31_424971 879 117 perpetuity perpetuity NN 10_1101-2020_12_31_424971 879 118 . . . 10_1101-2020_12_31_424971 880 1 It -PRON- PRP 10_1101-2020_12_31_424971 880 2 is be VBZ 10_1101-2020_12_31_424971 880 3 made make VBN 10_1101-2020_12_31_424971 880 4 The the DT 10_1101-2020_12_31_424971 880 5 copyright copyright NN 10_1101-2020_12_31_424971 880 6 holder holder NN 10_1101-2020_12_31_424971 880 7 for for IN 10_1101-2020_12_31_424971 880 8 this this DT 10_1101-2020_12_31_424971 880 9 preprintthis preprintthis NN 10_1101-2020_12_31_424971 880 10 version version NN 10_1101-2020_12_31_424971 880 11 posted post VBD 10_1101-2020_12_31_424971 880 12 January January NNP 10_1101-2020_12_31_424971 880 13 2 2 CD 10_1101-2020_12_31_424971 880 14 , , , 10_1101-2020_12_31_424971 880 15 2021 2021 CD 10_1101-2020_12_31_424971 880 16 . . . 10_1101-2020_12_31_424971 880 17 ; ; : 10_1101-2020_12_31_424971 880 18 https://doi.org/10.1101/2020.12.31.424971doi https://doi.org/10.1101/2020.12.31.424971doi NFP 10_1101-2020_12_31_424971 880 19 : : : 10_1101-2020_12_31_424971 880 20 bioRxiv biorxiv VB 10_1101-2020_12_31_424971 880 21 preprint preprint NN 10_1101-2020_12_31_424971 880 22 https://doi.org/10.1101/2020.12.31.424971 https://doi.org/10.1101/2020.12.31.424971 UH 10_1101-2020_12_31_424971 880 23 http://creativecommons.org/licenses/by-nc-nd/4.0/ http://creativecommons.org/licenses/by-nc-nd/4.0/ NN