Corneil University Library SB 191.W5W65 Hybridizing wheat and HNN TA 924 003 mann ALBERT R. MANN LIBRARY NEw YorK STATE COLLEGES OF AGRICULTURE AND HomE ECONOMICS AT CORNELL UNIVERSITY PLANT BREEDING LIBRARY Hybridizing Wheat and Emmer by Roy G. Wiggans Sept. 1915 Ce Contents Introduction--------------------------«--------- Object of the investigation--------------------- Resumé of previous work------------------------- Material used~----------------------------------- Conditions of experiment------------------- Description of wheats---------------------- ' Planting and harvesting-------------------- Method of crossing------------- w2--------------- Emasculation------------------------------- Pollination-------------------------------- Description of first generation crosses--------- Results of second generation-------------------- Color of glumes---------------------------- Beardedness~-------------------------------- Velveting---------------------------------- Various types of heads--------------------- Segregation into types--------------------- Cone lusions------------------------------------- Reference s--~-------------------------------------+ Oo nA OF DH EF &F FW WN KF FH mo 16 16 16 Ly 17 26 27 Fig. Fig. Fig. Pig. Fig. Fig. Fig. Fig. Pig. Fig. 6 ” 8 2 Illustrations Heads of F, and parent plants of Jones Paris Pridé x Black Winter Emmer cross.----- Heads of B and parent plants of Harly Arca- dian’x Black Winter Hmmer cross.------ Heads of F_ and parent plants of Pride of Genesee x Black Winter Hmmer cross.--- Emmer typeSer---- 2 enn ne nn nn nnn Spelt types8.-------------~----------- 2 ----- Emmer-spelt types.-------------------------- Spelt-wheat types.-------------------------- Emmer wheat types.-------------------------- Heads of unclassified plants.--------------- 10 Heads of unclassified plants.--------------- Page 13 14 15 18 ao 20 21 22 £35 24 8 HYBRIDIZING WHEAT AND EMMER Introduction In the crossing of any kind of plents where uniform results are desired it is absolutely necessary to start with uniform parents. Even in pure lines there is always considerable variation. For the work reported in this paper, pure lines of wheat, so far as it is possible to secure them, were used for one parent and for the other e pure line of Black Winter Hmmer. Object The object of the investigation wes: (1) to determine, if possible, any characters that are inherited in Mendelian fashion; (2) to secure new types which might be of value. Review of Previous Work The work done with the wheat emmer cross has been in general very unsatisfactory and, like the work reported inthis paper, has not been carried to a successful end. C. H. Saunders! msde reciprocal crosses of Colorado wheat and winter emmer which gave him in BF, fifteen distinct types where winter emmer was used as the female parent and twelve types where winter emmer was the male parent. The poorest sorts were eliminated end no definite mathematical Le study was made. He reports in the same paper a cross of Red Fife Wheat and Polish Wheat which gave many types in Fg, one of which was very similar to spelt. He thinks thet few if any of the types can be regarded as fixed in the third generation. Another cross/1l of Polish and common wheats gave many types but very few of Fy seed grew to maturity. Be C. Buffumltt secured what he called a most remark- able series of mules and monstrositiesfrom a supposed mute- ting type of wheet and a supposed mutating emmer. Inter- mediate types were ‘absent while almost every known type heretofore produced was present es well as decidedly new types of grain. Types which seemed to indicate reversion were numerous, one of which was the single-seeded spikelet of the monococcum type; another gave the dicoccum type; and another, true Triticum spelta. Buffum thinks that the results indicate a common origin for all forms of wheat. M.i. Carleton IV states that emmer readily crosses with wheat and that the department at Washington, as well as ex- periment stations, have used spring emmer for the purpose of adding rust resistance to ordinary wheat. Winter emmer should have even e greater value in this phase of wheat in- provement. Carleton?” gives the possible lines of improvement by the wheat emmer cross as : (1) Resistance to fungous et- tacks, (&) drought resistance, (3) increased fertility of the head, (4) non-shattering, (5) stiffness of straw, and (6) increase of gluten content of the grain. Material Used All the work was done in & greenhouse under con- trolled conditions. The temperature was regulated for the best growth of cereals and water was added when needed. It was necessary at timesto fumigate for aphids. The plants used, as stated above, were from pure lines of the wheats and emmer. The particular wheats used in the crosses were: (1) Red Hussar; (2) Pride of Genesee; (3) Jones Paris Pride; and (4) Early Arcadian. The crosses were made reciprocslly so far as the material permitted. The period of flowering varied somewhat, mak- ing it impossible at times to get the desired pollen for some crosses. Table I gives the description of winter emmer and the various wheats used in the crosses. 4. Table I. Characters |Winter Red Pride of | Early Jones Emmer Hussar Genesee Arcadian| Paris Pride Awned or Awned Awnless Awned Awnless | Awnless Awnlass Color of |Black at Light Awn base to x yellow =x x gray at top No. spike- Clubbed lets per ed. 8 10 wheat 8 5 cm. Color of |Grayish | Light Light Similar | A little Glumes black yellow yellow to Jones} darker to white| Paris than Pride Red Hussar Glumes smooth or | Velvet Smooth Smooth Smooth Smooth velvet Glumes loose or | Adhering | Loose Loose Loose Loose adhering tightly closely De The original crosses were made in the summer of 1913. The seeds thus secured were grown together with the parents the following year. In the fall of 1914 the seeds of Fy plants were sown in the greenhouse and parent plants were grown under the same conditions to eliminate any chance of change due to environment. In every case a sample head wes saved from the parent plants and where possible seeds of the Fy plents were saved. If this was not practicable photographs were taken so that the record would be complete. In June of the summer of 1915 the plants of Bo were harvested, each one separately in order that a study of the various characters might be made on the individual plants. Method of Crossing The actual method of making the crosses is very simple. Only a fair knowledge of the floral structures is necessary to make the work @ success. After working with the flowers for some time, it was found that it was easier to get the pollen from the wheat to pollinate the emmer flowers than to make the reciprocal cross. How- ever, the crosses were made both ways. The head is first prepared for emasculating by removing the lower spikelets and clipping out the top of the head until only eight or ten spikelets remain. The center flower or flowers of the spikelet are then removed with a pair of forceps. In be wheat there ere sometimes two and even more fertile cen- tral flowers, but in emmer these are generally sterile. This leaves sixteen to twenty lower flowers on each head. The anthers (three in number) are removed and the flower is left in this condition as a rule for two days. The time varies with the maturity of the flower when emasculated. Care must always be exercised not to injure the flower parts and especially the two-parted pistil. The heads after emasculation are covered with a semi-transparent bag two inches wide and eight inches long to prevent any possible crocs pollination. In the green- house it is not necesssry to tie the bags on the heads. When the pistils are mature, that is, receptive to pollen, they have developed a two-parted pistil. Both divisions are very much branched which gives e feathery appearance to the opened flower. &t this stage the stig- mas have usually begun to show between the palea and the flowering glume. If the flower is not pollinated, the glumes begin to spread apart exposing the pistil not only to its own pollen but to any pollen that might be cerried to it. This is the most advantageous time to pollinate as the stigmas are most receptive at this stage and a greater percentage of grain will be secured. The pollen to be effective should be ripe. Anthers that are opening are the best, but if they cannot be found, anthers which are very yellow must be used. Some of these will soon open and shed their pollen. In this way ell the anthers 7. that are gathered are covered with ripe pollen. It is advisable to place a whole anther in each flower thet is to be pollinated. Pollen grains are #11 that is necessery if properly placed on the stigma of the flower. Polline- tion is accomplished by carefully opening the flower either with forceps or by gently pressing the upper end of the glumes with the left forefinger while holding the base firmly and placing ripe anther on the stigma. There are many circumstances that mitigate against . success in this work, but with sufficient care crosses can be secured in either direction with ell the varieties re- ported in this paper. Crosses with wheat and emmer are considerably more difficult than wheat crossed with wheat. This is probebly explained by the fact that emmer is some distance from wheat speaking from the standpoint of evolu- tion, but we would assume that it is closer than rye as crosses with rye are very difficult to secure and all that heve been reported are sterile. The success of the crosses herein reported may be given briefly es follows: Cross No. of kernels Winter emmer x Early Arcadian 37 = : x Pride of Genesee 55 " u x Jones Paris Pride 9 WT " x Red Hussar 33 8. The reciprocal crosses are given together es there was no difference in the results whichever way the cross was made. The heads pollinated varied from a complete failure to sixty-five per cent set. In the care of wheat x wheat a large percentage was very common, fifty to sixty per cent being the average with an occasional percentage as high as ninety. The seeds secured from the crosses were planted together with parent plants as before stated. The characteristics of the i plants produced from the above seed are given in Table II. Description of First Generation Crosses Table II. 9. ones Red Marly Arca-|Red Hussar | Pride of Gene- Characters Mane dian see x x x x Black Winter| Black Win-|Black Win- | plack Winter _Emmer ter Hmmer | ter Emmer Emmer Awned or Intermediate Inter- Int er- Awned awnless mediate mediate. Color of |Dark gray Gray Light gray | Light gray awn No. spike- lets over 10 Clubbed 12 iz 5 cm. Color of From al- Little (Inter- Varying from glumes most as darker mediate) almost as dark as than dark dark aS emmer emmer to Red gray to much Lighter very light Hussar (smooth Glumes( or Smooth Smooth Smooth Smooth (velvet (loose Glumes ( Interme- Inter- Inter- Intermediate (tight diate mediate |mediate 10. Table II gives the average results of F,, which indicate in some way what has previously been demonstreted in wheat crosses. Beardedness in F, is an intermediate character. The results of Biffen’! lead one to conclude this although he classes beardedness aS recessive. He fails to count in the bearded class the heads which are only slightly bearded but places them in the class with the beardless. It is probable all the results that have been obtained are similar in regard to the beardedness being recessive put the interpretation has been different. WilsonV2! like Biffen reports that awnlessness is dominant, giving 664 awnless and 207 awned. BatesonVill e1iso reports the same thing but upon observation of the illustrations given, it can be seen that some of those which he calls beardless possess beards, varing in number and length. A like statement can be found which was made by Punnet®. He fails to give any data to support what he assumes as a fact, probably in part because he uses it only as an argument for a theory which he was presenting. The work of Alvin Keyser shows very conclusively that the result when bearded andbeardless types are crosses is an inter- mediate product. This is not based upon one cross but upon many crosses and as his observations were upon the one chsrac---- ter they probably represent very accurately what really ex- isted. In F, he found practically 25 berrdeds 50 inter- z mediate; 25 beardless, the actual per cents being 21 bearded, Les 53 intermediete and 26 beardless. Both the awned and ewnless bred true in Bae The inheritance of beardedness is an unsettled question and offers a very good opportunity for an accurate and detailed statisticel study from a Mendelian standpoint. The color of awns is © cheracter which is probably intermediate in F,. There is, however, scarcely sufficient evidence to support this statement. The record of the inheritance of lsexness or denseness in the ears seems much more conclusive. Fy hes practically the same head, so far as lamess is concerned, as does the perent with the lexer head. Biffen’? reports that loose heads are dominant to compact heads. W. He Parker*! in a rather extensive report of "Lax and Dense-esared Wheat" finds the problem of inheritance of denseness and laxness a very complex and entirely unsettled one. His results are conflicting and definite conclusions cannot be drawn from them. Prof. Spilimen*t1 reports 14 crosses of dense and lax wheats with the result of 1 lax; 2 intermediate; 1 dense. His actual percentages were 27.2; 47.2: 25.6. The different varieties gave very different results but the above percentages are the average of all his crosses. Nilsson-Bhle“!!! ginas 2 1:2:1 ratio in Fp with dense and lax-eared wheats. cufficient evidence is not given in oy of the crosses reported in this paper to indicate sccurately how the color of glume is inherited. 12. All the heads in FP. were smooth but it is possible iL _ that a microscopic examination would have shown velveting of the glumes. Birfen’? reports velveting to be dominant. The studies in Fp indicate that this is the case. In F, the tightness with which the glumes adhered to the kernels wes intermediate in character. A detailed study of this was not made on Bo plants. A very profitable study could be made on this character as the shattering of wheat is a serious problem in some districts where the grain is permitted to become very ripe before harvesting. A wheat with a glume that adhered more closely would be much more desirable than the ones now grown if this character could be combined with the desirable characters already possessed by the present wheats. The following photographs show chsracteristic heads of the wheats and emmer used. The heads of the parent stocks used for the crosses are shown together with a head of the F, plants. Some of the characters described above are apparent in the photographs. 13. Fig.l. Heads of parent plants with the F. cross. 1256 X Jones Paris Pride. 1 1256 A F_ cross. 1256 Y lBlack Winter Emmer. 14. L260 a.| G2CIY 0 Fig. 2. Heads of parent plents with the K cross. 1260 X Black Winter Emmer. 1260 A F. cross. 1260 Y Barly Arcadian. 15. Fig. 3. Heads of parent plants with Py cross. 1252 X Pride of Genesee. 1262 A By cross. lebe Black Winter Emmer. 16. Results of second Generation The grain of the F_ plants was sown and the Fo 2 plants grown and harvested separately. The results as they come out in the individual plants are very confusing but some characters could be studied very accurately. The inheritance of the color in the glumes comes out in a very good Mendelian ratio of 3:1, the actual numbers be- ing 179 black; 53 white. In the blacks were included all shades and degrees of blackness from the color of emmer to the slightest indication of the presence of a color factor. All shades could be found in every cross where there were sufficient numbers. The bearded character gave 87 bearded; 90 intermediate :::::: 64 beardless. If the bearded and intermediates sre grouped together 177 bearded : 64 intermediates or a 3:1 meek no indication as to dominance of beardlessness over bearded- nes8. The velveting character wes not studied on all the crosses but particular care was given it in the cross be- tween Red Hussar x Black Winter Emmer. In this cross there seems to be two factors which control the inheritance /of the hairs. The Red Hussar type has hairs at the base (of the spikelet but the glumes are free while the Winter | Emmer used had heirs on the glumes put the base was free. It was often necessary to use a strong lens to determine the presence of the hairs. The ratio secured was: Les 35 with hairs at base and on the glumes. 26 with beirs at bese only. 23 with hairs on the glumes only. 4 with no hairs. The same thing studied with a cross of Mediah and Minnesota Wheat #169 gave the ratio 56:29:25:2. Both made very poor 9:3:3:1 ratios but they give sufficient evidence to show that there is a segregation of two charac- ters and that they are inherited independently. The shape of kernel, shape of glume, and texture of glume were in- herited independently but were not studied especially for segregation. The emmer textured glume seemed to be dominant. By texture is meant the stiffness end coarseness of the glumes. Various Types of Heads in Bos So far as a segregation of types is concerned it is very difficult to get any ratios or any definite classes. In order to make a study of the Fy plants they were arranged in groups with characteristic differences. The groups were named (1) Emmer type, (2) Spelt type, (3) Emmer spelt, (4) Spelt wheat, and (5) Emmer wheat. Those which did not fall into these arbitrary groups were placed in a sixth group which was made up of all kinds and shapes of heads most of which were clubbed. The names of the groups explain more or less the characters of the heads of the plants placed in them. Likewise the photographs illustrate their general cherac- teristics. In each group were found both bleck and 18. white, and beardless and bearded except in groups (1) and (3) where no beardless types appeared. Fig. 4 Emmer types : I White bearded, II Black bearded, III Intermediate black beard- ed, IV Black beardless. Fig. 5. Spelt types: I Black bearded; II Intermediate bearded black; III Black beardless; IV White bearded; V White intermediate bearded; VI White beardless. 206 Fig. 6. Emmer spelt types: I Black bearded; II Black intermediate bearded; III White intermediate bearded. Fig. 7 Seplt wheat types: I Black bearded; II Black intermediate bearded; IiI White bearded; IV White inter- mediate bearded. an 226 Fig. 8. Eqmer wheat types: I Bleck bearded; II Black intermediate bearded; III Black beardless; IV White bearded; V White intermediate bearded. Heads of unclassified plants; e; Il Einkorn type; , and VI Club types. 256 Heads of unclassified plants con- tinued: VII-X all club types most of which are sterile. 24. 256 Table III. Jones | #arly Red Pride Paris | Arcadian |Hussar of Total Pride Genesee Black chaff 56 Ac: 61 10 179 White chaff 6 3 ne 3 53 Bearded 15 5 20 15 87 intermediate Bearded 16 5 o7 ae 90 Beardless AL a 24 C 64 (Bearded Black c 5 Clo Oy Emmer (Bearded White i G Git, codecs Type (Reardless Black [ ¢ |. Ct ee (Rearijless White 0 0 Xx 0 (Bearded Black [| 3 | J 2 i 2 Spelt (Bearded White 0 spe ___4 Type (Beardless Black 6 eh [ie ea a (Beardless White 0 a x 3 (Bearded Black 4 2B 5 e LF Emmer (Bearded White i a o |. 2 Spelt (Beardless Bleck 0 0 x 0 (Beardless White 0 0 x 0 (Bearded Bleck 1 a: Ei ee Spelt (Bearded White {; 0 0 0 2 Wheat (Beardless Black’ 0 2 x 2 (Beardless White; 0 0 x 1 (Bearded Black | 6 4 16 3 56 Emmer (Bearded White — & 2 13 2 a Wheat (Beardless Black 2 2 or x 14 (Beardless White’ 6 iE S xX 4 Wheat )-Black bearded ze min a a Type ) Velvet ) 29 Glabrous ) i be 7 Not classified only as | 8 0 a2 14 41 clubs H)-B,,)/Both clubbed H)-A, )but clacsed as emmer wheats. £6. TableIII gives the distribution in the various classes of the Fo plants of the four crosses studied. The unclassified group illustreted by Figures 9 and 10 represent a great variety of forms varying from einkorn to true wheat as well as the vaious clubbed forms. Conclusions ile That in any cross of wheat and emmer there is obtained in Fe & series of forms which resemble einkorn spelt, emmer, common wheats, and club wheats. Be That no definite Mendelian segregations were found as to form of head. De That the characters beardedness, color of glume, and velveting are inherited in Mendelian fashion. 4. That large numbers would have to be studied through a series of years before any accurate conclusions could be drawn. ACKNOWLEDGMENT The writer desires to thank Dr. H. H. Love for his directions and kindly assistance in the study re- ported above and for the material which made it possi- ble for the work to be done. ils Ad din IV. Vile VEIL Ix. Xe XI. ail. XIII. VA References Seunders, C. H. 1905 Some observations on heredity in wheat. AeBeAe Vol.l, pPe 77=—78 1905 A natural hybrid in wheat. A.3.A. Vol. Ls ppe 137-138. Buffum, 3. C. 1909 Effect of environment on plant breeding. ABA. Vol. VI4 ppe 212-225. Carleton, Mark Alfred. ei Winter Emmer. U. S. Dept. of Agr. Farmers Bul. 4¢66. 1SC01l Emmer: A grain for the semiarid regions. U.S.Dept. of Agr. Farmers Bul. 139. Biffen, R. H. 1905 Mendel's laws of inheritance and wheat breeding. Jour. of Agr. Se. Vol. I, pp. 4-48. Wilson, John H. 1907-08 The hybridization of cereals. Jour. Of Agrs Ses Vols Il, pps 68-88. Bateson, W. 1909 Mendel's principles of heredity. Punnet, He. C. 1911 Mendelisme Keyser, Alvin. 1906 Variation in wheat hybrids. ABA. Vol. By pp,84-89, Parker, W. H. 1914 Lax and dense-eared wheats. Jour. of Agr. Sc. Vol. VI, pp. 371-386. Spillman, IgGs Journ. Royal Hort. Soc. XXVII, p. 876. Nilsson-Ehle Lunds Universitets Arsskrift, N.F. a.f.d. 2, BA.V, Nre&, ps 104 et seg.