THE UNIVERSITY OF ILLINOIS LIBRARY £3 0.7 M TO b* cop. Z. ,o NON CIRCULATING CHECK FOR UNBOUND CIRCULATING CORY Digitized by the Internet Archive in 2016 https://archive.org/details/charactersconnec4656ethe UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 46 Characters Connected With the Yield of the Corn Plant (Publication Authorized August 6, 1921.) COLUMBIA, MISSOURI AUGUST, 1921 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY P. E. BURTON H. J. BLANTON St. Louis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION J. C. JONES, PH. D., LL. D., ACTING PRESIDENT OF THE UNIVERSITY F. B. MUMFORD, M. S., DIRECTOR STATION August, AGRICULTURAL CHEMISTRY C. R. Moui/ton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, A. M. E. E. Vanatta, M. S. R. M. Smith, A. M. A. R. Hall, B. S. in Agr. E. G. SiEvEicing, B. S. in Agr. C. F. Ahmann, A. B. AGRICULTURAL ENGINEERING J. C. Wooley, B. S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumeord, M. S. D. W. Chittenden, B. S. in Agr. Paul M. Bernard, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. E. F. Hopkins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale, B. S. in Agr. W. W. Swett, A. M. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. C. McBride, B. S. in A. FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, A. M. O. W. Letson, A. M. B. B. Branstetter, B. S. in Agr. B. M. King, B. S. in Agr. Alva C. Hill STAFF 1921 RURAL LIFE O. R. Johnson, A. M. S. D. GromEr, A. M. Ben H. Frame, B. S. in Agr. FORESTRY Frederick Dunlap, F. E. HORTICULTURE V. R. Gardner, M. S. A. H. D. Hooker, Jr., Ph. D. J. T. Rosa, Jr., M. S. F. C. Bradford, M. S. H. G. Swartwout, B. S. in Agr. POULTRY HUSBANDRY H. L. Kempster, B. S. Earl W. Henderson SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M W. A. Albrecht, Ph. D. F. L. Duley, A. M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr Richard Bradfield, A. B. O. B. Price, B. S. in Agr. veterinary SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. ZOOLOGY George Lefevre, Ph. D. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Secretary S. B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Bertha Hite , 1 Seed Analyst. Un service of U. S. Department of Agriculture. CHARACTERS CONNECTED WITH THE YIELD OF THE CORN PLANT W. C. Etheridge In 1909 the Department of Agronomy of the Missouri Ex- periment Station began a study of the factors influencing the de- velopment of the corn plant. In 1914 this department was divid- ed into the Departments of Soils and Field Crops, which there- after separately carried on those phases of the study most appro- priate for their respective attention. The study as a whole end- ed in 1920. That part of it directly concerned with the effect of nutrition upon growth has been reported by Duley and Miller. 1 That part concerned with the correlations between structure and function will be reported in this paper.* I. — A Study of the Correlation Between Yield and Certain Characters of the Corn Plant. The essential purpose of this study was to contribute to the solution of a problem then (1914) receiving much attention in the field of plant genetics — the correlation between yield and measurable variations in the visible characters of the corn plant. It is a familiar problem to all who have read closely the agro- nomic literature of the past 12 years. Likewise its conclusion, though never a real solution, is nearly conventional, for almost without exception its investigators have reported (1) that the correlations did not exist or (2) that those observed were not significant. The brief results reported in this paper are not ex- ceptional to the ensemble of evidence from similar studies by other investigators. They are reported because (1) though brief, they contribute a clear case and (2) the great weight of concor- dant evidence now existing would seem almost to preclude a pos- 1 This and subsequent numerical references are to the Bibliography. *The writer had no connection with this project. He is merely a reporter of results se- cured in 1910, 1911 and 1914, from studies by C. B. Hutchison, C. E. Neff, S. B. Nuck- ols and others. His presentations and interpretations are therefore critical. Possibly the original investigators would have presented their data more accurately; possibly they would have interpreted it differently. 4 Missouri Agr. Exp. Sta. Research Bulletin 46 sibility that further study of the problem by the present con- ventional methods would prove fruitful. REVIEW OF RELATED LITERATURE To review in detail the evidence contributed by previous inves- tigators would show to many readers a familiar picture. It seems unnecessary to do that. Therefore the following brief summary presents only the essential developments. In 1909, Montgomery 2 reported that a long (large) ear, medi um depth of the kernel and stockiness of the stalk, were corre lated with relatively high yield. Variation in other characters of the plant and ear showed no relation to yielding ability. The correlation between size of the ear and yield is of course obvious — one is an expression of the other. In the same year Hartley 3 reached this very pointed conclusion — “No visible characters of apparently good seed ears are indicative of high yielding power.” He had made more than 1,000 ear-rows tests of 4 varieties, over a period of 6 years. In 1910, Pearl and Surface 4 said they found no evidence of close association between the conformation of the seed ear and the yield that it produced. They had studied two very different types of sweet corn, giving particular attention to the shape of the ear and the covering of the tip. Ewing 5 after a very thorough study of the variation in several dimensional characters, such as height, and leaf breadth, concluded that “No single character among those studied has shown itself so closely connected with yield as to stand out as a safe guide to the breeder.” In 1911, Love 6 concluded that no characters of the ear were highly correlated with earliness and that none could serve as an index of earliness. Sconce 7 an Illinois seed corn breeder, after a study of 6 years, stated his belief that the number of kernel-rows, the form of the kernel and the size of the germ were correlated with the yield of grain. Funk 8 , another Illinois seedsman, while not denying the existence of correlations, concluded that the conventional corn score card does not emphasize the points that affect yield. When he maintained by selection the type which made the highest yields, he gradually produced an ear very diff- erent from that idealized by the score card. In 1913 McCall and Wheeler 9 presented their interpretations of various statistical data of other investigators. They concluded Characters Connected with the Yield of the Corn Plant 5 that significant correlations between yield and length, weight, circumference and density of the ear, had not been shown. In 1914 Williams and Welton 10 made an exhaustive report of studies through a period of 10 years. As an average, long ears showed an advantage in acre yield of 1.39 bushels over short ears; but tapering ears showed an advantage of 1.65 bushels over cylin- drical ears; bare-tipped ears 0.34 bushels over full-tipped ears; smooth-dented kernels 1.76 bushels over rough-dented kernels; and ears of a high shelling percentage (88.16) were 0.52 bushels lower in acre yield than ears with a low shelling percentage (76.07). In 1916, Cunningham 11 reported that smooth and medium smooth kernels outyielded rough kernels by a considerable mar- gin. Variation in several other characters showed no correla- tion with yield. He concluded that correlations were variable with the environment. In 1917, Cove and Wentz 6 found that “The characters of length, ratio of butt to tip, average circumference of cob, weight of ear, average weight of kernels, number of rows of kernels, and average length and width of kernels in seed ears do not show correlations significant enough to be of value in judging seed corn.” They reached this conclusion after five years of study with one variety. Hughes 12 believed the first year’s results of his experi- ment with seed corn indicated a close correlation between yield and the ear characters idealized by the score card. In 1918 Hutcheson and Wolfe 13 believed they had found sig- nificant correlations between yield and the size and trueness to type of the ears. Many other characters, such as shelling per- centage, number of rows, space between rows, and the filling of the butt, were not related to yield in a significant way. Olson, Bull and Hayes 14 from apparently the soundest and most com- prehensive study yet conducted, failed to find a significant cor- relation between yield and any of a broad range of characters ob- served. They made the very practical statement that “Close se- lection for high scoring ears is of no practical value in increas- ing yield.” 6 Missouri Agr. Exp. Sta. Research Bulletin 46 MATERIAL AND METHODS Ten ears of each type in the following groups were selected from the variety Boone County White, as seed for the crop in which the correlation study was to be made. A. Long, slim, smooth ears B. Long, thick, smooth ears C. Short, thick, smooth ears D. Medium long, medium thick, medium rough E. Long, slim, rough ears F. Long, thick, rough ears G. Short, thick, rough ears Each of the 70 ears was planted in an ear-row, each type in a separate series. Thus there were 10 ear-rows of long, slim, smooth ears ; 10 of long, thick, smooth ears ; and so on. The series were contiguous and each fifth row was a check, planted with seed of one type. The hills were spaced 44 inches apart, each way, and two plants -were left in each hill as the final stand. The crop re- ceived ordinary cultivation. Just before the tasseling stage 40 normal plants in each row were labeled, each plant standing among similar normal plants. There were labeled a total of 2800 plants and for each the following data were recorded : 1. Date of tasseling. 2. Date of silking. 3. Number of tillers at full growth. 4. Leaf area above the ear at full growth. 5. Leaf area below the ear at full growth. 6. Total leaf area at full growth. 7. Height of ear at full growth. 8. Relative position of the ear-shank. 9. Height of stalk at full growth. 10. Number of nodes in stalk. 11. Circumference of first internode above ground, at full growth. 12. Circumference of first internode above ear, at full growth. 13. Tassel length. When two ears were borne by a plant, all measurements with reference to the ear were made by the upper ear only, although the total yield of both ears was determined. The leaf area was the Characters Connected with the Yield of the Corn Plant 7 sum of the areas of individual leaves measured by Montgomery’s formula — Area=12x^4 (breadth x length). The tassel length was measured by finding the sum of the length of five average lateral branches, dividing this sum by five and multiplying the quotient by the number of laterals, then adding to the product the length of the central spike. Sound ears were gathered from 1,761 of the 2,800 plants measured, and stored under good drying conditions for six weeks. The weight of shelled grain produced by each plant was computed on the basis of a uniform content of moisture. THE RESULTS The mean yields of shelled grain produced by plants from the various types of seed-ears are shown here. Table 1. — The Relative Productivity oe Seed From DieeerEnt Types oe Ears. Series Type of original seed ear No. of ears harvested for yield test Mean yield in ounces of shelled grain per plant Probable error (±) A Long, slim, smooth 195 7.7400 .1409 B Long, thick, smooth 228 7.7150 .1169 C Short, thick, smooth 256 8.1836 .1209 D Medium long, medium thick, medium rough 264 7.9924 .1100 E Long, slim, rough 256 8.7740 .1182 F Long, thick, rough 270 8.2741 .1190 G Short, thick, rough 292 8.2363 .1111 Composite 1761 8.1525 .0452 The mean yields range highest in Series E, F, and G, and lowest in Series A, B, C, and D. But between the highest yield, Series E, and the lowest, Series B, there is a difference of only 1.06 ounces of shelled grain per plant. This difference, though small, might be significant did not the yields of the check rows (Figure 1) show that Series E was favored by a variation in the fertility of the soil. Doubtless Series F and G were likewise fav- ored. There were then no significant differences between the yields of plants produced from the various seed-ears representing an extremely wide range of form and identation, in the variety Boone County White. Missouri Agr. Exp. Sta. Research Bulletin 46 •Showing the comparative location of Series A to G, and the location and yield in pounds of grain per row of the check rows marked O. Characters Connected with the Yield of the Corn Plant 9 The correlation coefficients determined for the weight of shell- ed grain as the subject and various plant characters as the rela- tives are now shown. Table II. — Correlations Between Variations in Plant Characters and Weight or Shelled Grain per Plant. Character Coefficient of correlation Probable error (±) Days from planting to silking -.4181 .0133 Leaf area above ear .0885 .0167 Leaf area below ear .0565 .0167 Total leaf area .0702 .0161 Height of stalk .1109 .0160 Number of nodes in stalk .0843 .0161 Height of ear -.0006 .0161 Relative position of the ear-node Circumference of internode .0340 .0161 above ground Circumference of internode .1846 .0155 above ear .0893 .0160 Length of tassel -.1251 .0170 Number of tillers -.0160 .0160 Although some of these correlations are statistically signifi- cant, none of them are high enough to be valuable as an index of yield. The negative correlation between yield and the age of the plant at silking, the highest correlation found, would doubtless vary greatly with the season. DISCUSSION The results of this brief study are concordant with those of other studies herein cited, in finding no significant correlations be- tween the yield of the corn plant and variations in its visible struc- tures and characters. But these and all similar results make no proof that such correlations do not exist, although the total evi- dence has come from a very exhaustive analysis. To accept fully the negation of correlations would lead to the conclusion that the corn plant is exhibiting the phenomenon of no relationship be- tween external structure and function. Of course the correlations exist. Why then are they not found in a measure that would justify 10 Missouri Agr. Exp. Sta. Research Bulletin 46 their use as an index of the relative ability of the progeny to yield? A very simple explanation may be suggested. In all studies of such correlations yield has been treated, con- sciously or not, as a single character of the plant. Obviously, this conception of yield is fundamentally wrong. Yield is a per- formance, not a character. It is the ultimate performance of the whole complex relationship of functions and structures that make up the plant. No doubt each function and structure varies with the environment. No doubt each variation influences yield ; but only as it contributes to the final complex result of all variations. And so the influence of a given variation upon yield cannot be finally measured, simply because it cannot be identified and sep- arated from the combined influence of innumerable other variations. But is there no visible index of yielding ability that may serve as a guide in the practical operation of selecting seed corn? It was to answer this question that all correlation studies of corn were begun. Certainly there is such an index. It is yield itself — al- most the old and simple idea of selecting the biggest ears. Taking as an example any common one-eared variety of the Middle West, the yield of grain from plant to plant must vary with the size of the ear, excluding of course the slight variation in shelling percentage and the losses from unsoundness. So far then as yield can be improved by seed selection, the most exhaus- tive studies have discovered no better method than field selection of the biggest, soundest ears, well matured and unfavored by ap- parent differences in their local environment — stand, fertility, and so forth. Or if the plant bears more than one ear, of course its total yield, rather than the size of the individual ear, should be considered. In a given environment the best adapted and best yielding strain will of course show some distinguishing character- istic. For example, under certain conditions the highest yielding strain may have smooth kernels. But it does not follow that con- tinued close selection of smooth seed ears will increase or even maintain the yield of the strain. For by that operation a specialized strain not so well balanced with the environment might be pro- duced. CONCLUSION Within the conventional limits of a variety of corn, no varia- tion in the visible structures or characters of a normal, healthy plant is a reliable index of the relative ability of its progeny to Characters Connected with the Yield of the Corn Plant 11 yield. The relative yield of the mother plant is the only indica- tion, uncertain as it may be, of the relative yield of the progeny. This conclusion is based not wholly upon the brief evidence presented in this paper, but upon the total evidence contributed by all investigators of the problem. BIBLIOGRAPHY 1. Duley, F. L. and Miller, M. F. The Effect of a Varying Supply of Nutrients Upon the Character and Composition of the Maize Plant at Different Periods of Growth. Mo. Agr. Expt. Sta. Res. Bui. 42. 1921. 2. Montgomery, E. G. Experiments with Corn. Neb. Agr. Expt. Sta. Bui. 112. 1909. 3. Hartley, C. P. Producing Higher Yielding Strains of Corn. U. S. Dept. Agr. Yearbook, 1909: 309-320. 4. Pearl, R. and Surface, F. M. Experiments in Breeding Sweet Corn. Me. Agr. Expt. Sta. Bui. 183. 1910. 5. Ewing, E. C. Correlation of Characters in Corn. Cornell Univ. Agr. Expt. Sta. Bui. 287. 1910. 6. Love, H. H. The Relation of Seed Ear Characters To Earliness in Corn. Amer. Breeders Ossoc. Rpt. 8: 330-334. 1911. 7 and Wentz, J. B. Correlations Between Ear Char- acters and Yield in Corn. Jour. Amer. Soc. Agron., 8, 7: 315-322. 1917. 8. Sconce, J. H. Scientific Corn Breeding. Amer. Breeders Assoc. Rpt. 7: 43-50. 1911. 9. Funk, E. Ten Years of Corn Breeding. Amer. Breeders Mag. 3, 4: 295. 1911. 10. McCall, A. G. and Wheeler, C. S. Ear Characters Not Correlated with Yield in Corn. Jour. Amer. Soc. Agron., 5, 2: 117. 1913. 11. Williams, C. G. and Welton, F. A. Corn Experiments. Ohio Agr. Expt. Sta. Bui. 282. 1915. 12. Cunningham, C. C. The Relation of Ear Characters of Corn to Yield. Jour. Amer. Soc. Agron., 8, 3: 188-196. 1916. 13. Hughes, H. D. An Interesting Experiment with Seed Corn. Iowa Agr., 17, 9: 424, 425, 428. 1917. 14. Hutcheson, T. B. and Wolfe, T. K. Relation Between Yield and Ear Characters in Corn. Jour. Amer. Soc. Agron., 10, 6: 250-225. 1918. 15. Olson, P. J., Bull, C. P. and Hayes, H. K. Ear Type Selection and Yield in Corn. Minn. Agr. Expt. Sta. Bui. 174. 1918. 12 Missouri Agr. Exp. Sta. Research Bulletin 46 II. — A Study of the Relation of Certain Ear Characters to Shelling Percentage Shrinkage and Viability. This study was made in 1910 and 1911. Its purpose was to find whether variations in certain characters commonly used in judging seed ears were indicative of the relative shelling per- centage, shrinkage and germination. MATERIAL AND GENERAL METHODS In 1910, 660 sound ears of a rough, large-eared strain of Boone County White, grown on rich alluvial soil, harvested in December of 1909 and stored in a tightly boarded crib until March 1, were used as experimental material. They will be designated as Lot A. In 1911, 500 sound ears of the same variety, but of a more variable strain, were used. They too had been grown on rich alluvial soil, but had been harvested early in October and air- dried on racks in a mouse-proof seed room for a period of 12 weeks. They will be designated as Lot B. Both lots were selected at random, except with reference to soundness. In both lots the individual ears were described in the details hereafter stated in Tables I — IV. All descriptions were recorded by the same person. No mathematical correlations were determined, but all comparisons were made between two classes showing extreme variation of the character in question, each class constituting about 15 percent of the total number of ears in the lot. For example, in studying the relation of length of ear to shelling percentage in Lot A, the average shelling percentage of the 100 longest ears was compared with that of the 100 shortest ears in the same lot. THE RELATION OF EAR CHARACTERS TO THE SHELL- ING PERCENTAGE OF THE EAR In Table I is shown the relation of various ear characters to shelling percentage, as determined by this method. The differ- ences are in most cases slight and inconsistent. Except the differ- ence between light and heavy ears, which may be attributed to the higher moisture content of the latter (see Table II), the size Characters Connected with the Yield of the Corn Plant 13 and shape of the ear show no significant relation to shelling per- centage; but ears marked by deep kernels, narrow kernels, and starchy kernels produced a slightly higher proportion of grain than ears marked by shallow, wide, and horny kernels. Table I. — Ear Characters and Shelling Percentage. Lot A — 1910 Lot B - —1911 Character of the ears Shelling percentage Ave. weight of grain (grams) Shelling percentage Ave. weight of grain (grams) Long 84.1 451.4 84.4 393.3 Short 86.1 367.5 84.5 347.1 Large circumference 84.4 450.7 84.7 408.3 Small circumference 84.8 371.1 86.3 357.0 Heavy 83.8 464.0 86.0 422.0 Light 89.9 358.5 85.7 327.2 Many rows of kernels 85.0 441.2 85.5 411.6 Few rows of kernels 83.3 384.4 84.5 369.4 Cylindrical 84.1 412.4 85.2 326.5 Tapering 84.4 411.8 84.6 381.0 Rough indentation 84.1 421.7 83.9 396 . 8 . Smooth indentation 84.2 411.8 84.2 337 . 6 . Deep kernels 85.7 450.1 86.4 409.2 Shallow kernels 83.1 383.2 83.2 334.7 Wide kernels 83.8 415.8 84.1 382.2 Narrow kernels 85.2 419.0 85.7 393.0 Horny kernels 82.0 391.2 83.8 346.3 Starchy kernels 85.5 415.8 85.6 379.8 THE RELATION OF EAR CHARACTERS TO THE SHRINK- AGE OF THE EAR The relation of ear characters to shrinkage was studied in Lot B by comparing the length, circumference, and weight of the ears as first stored, with their length, circumference and weight at the close of the total drying period of 6 weeks. The results of this study are shown in Table II. Little relation is shown between shrinkage and indentation or between shrinkage and the length and shape of the ear. Ap- parently, however, heavy ears, thick ears, deep-kerneled ears, and ears with a large number of rows, lost considerably more weight than ears of the opposite types. 14 Missouri Agr. Exp. Sta. Research Bulletin 46 Table II. — Ear Characters and the Average Shrinkage in Length, Cir- cumference and Weight of Ears of Lot B. Character Loss in Length Loss in Circumference Loss in Weight of the ears Inches Percent Inches Percent Grams Percent Long .4572 4.4 .3825 5.3 69.4 15.6 Short .3520 4.0 .3843 5.2 61.6 16.5 Large circumference .4250 4.6 .4624 5.9 75.4 17.0 Small circumference .3354 3.4 .3329 4.9 49.9 12.6 Many rows .3790 4.0 .3930 5.1 77.3 17.1 Eew rows .3612 3.7 .2295 3.4 59.8 14.5 Heavy .5140 5.2 .4183 5.6 92.4 19.1 Light .3412 3.3 .3300 4.7 44.5 12.8 Cylindrical .4362 4.5 .4000 5.8 63.9 17.3 Tapering .3710 3.9 .4000 5.5 65.8 15.6 Rough indentation .3651 3.8 .3475 4.6 58.2 13.7 Smooth indentation .4222 4.5 .4030 5.9 67.8 14.2 Deep kernels .3849 4.0 .3520 4.6 89.7 18.3 Shallow kernels .3108 3.2 .3700 5.5 59.8 15.7 In the same lot of ears the rapidity of shrinkage was deter- mined by weighing at intervals of two weeks, 300 ears grouped in extreme classes as previously described. The results are shown in Table III. Table III. — Ear Characters and the Progressive Rate of Shrinkage. Character Percentage of Loss in weight of the ears 2-wks 4-wks 6-wks 8-wks 10-wks 12-wks Total Large circumerence 8.6 5.6 1.7 1.1 0.7 0.5 18.2 Small circumference 6.6 5.3 1.6 1.2 0.4 0.4 15.5 Heavy 9.6 6.2 1.6 1.1 0.5 0.5 19.5 Light 6.9 4.5 1.8 0.8 0.3 0.6 14.9 Many rows 8.6 6.2 1.5 1.0 0.6 0.3 18.2 Few rows 7.8 5.2 1.9 1.0 0.8 0.4 17.1 Rough indentation 7.8 5.5 1.7 0.9 0.4 0.5 16.8 Smooth indentation 8.2 5.9 1.8 0.9 0.8 0.6 18.2 Deep kernels 10.5 5.9 1.9 0.9 0.6 0.2 20.0 Shallow kernels 7.9 5.3 1.5 1.2 0.5 0.2 16.6 Horny kernels 8.4 5.7 1.6 1.1 0.2 0.4 17.4 Starchy kernels 8.9 5.7 1.8 1.0 0.4 0.5 18.3 Characters Connected with the Yield of the Corn Plant 15 It may be noted first that in all classes of ears more than 75 percent of the total shrinkage occurred during the first four weeks, and that thereafter the shrinkage in all classes of ears was very slight from one two-week interval to another. Weather con- ditions were about the average for October, November and De- cember in this section. These results then may indicate the probable time required to air-dry seed corn under good condi- tions of farm storage. Apparently it would not be necessary to keep the seed ears on racks or various other drying devices for longer than a month. They could then be stored in a more con- venient bulk without damage because of the moisture they con- tained. Their remaining moisture would be given off very slowly and uniformly over a long period. There seems little significance in the relative rates of shink- age by ears of the different types. Large ears and heavy ears lost moisture more rapidly during the first two weeks than ears of the opposite types, due probably to their large, heavy cobs. The comparatively rapid drying of deep-kerneled ears may indi- cate the desirability of this type for seed, provided they are also large, sound, and well matured. THE RELATIONS OF EAR CHARACTERS TO VIABILITY At the time of this study the ears (Lot A) were two years old. They had been harvested in December 1909 and stored for nearly 3 months under rather poor conditions before they were sent to the Experiment Station. Their shelling percentage had been determined (Table I) and the grain of individual ears, stored separately in bottles, had been fumigated several times with hydro- cyanic acid gas. In November, 1911 this seed was tested for germ- ination. To make the tests, kernels were planted at a depth of 1 inch in sand which was kept at a temperature of about 80°F during the day and about 60°F during the night, and in a fairly uniform condition of moisture. A composite hundred kernels from each ear of the GGO-ear lot — a total of 66,000 kernels — were planted in two equal series, one 12 days later than the other. Ten days after planting, the numbers of strong sprouts, weak sprouts, and sprouts not appearing above the ground, were counted. The results are given in the following table. 16 Missouri Agr. Exp. Sta. Research Bulletin 46 Table IV. — Ear Characters and Germination. (Percentage of Germination in 10 Days) Character of the ears Strong plants Weak Plants not plants above ground Total germination Long 34.8 12.1 7.2 54.1 Short 37.3 13.3 9.6 60.1 Large circumference 30.5 11.0 7.3 48.8 Small circumference 45.4 12.9 6.8 65.1 Heavy 28.3 11.8 7.1 47.2 Light 42.2 12.6 7.0 61.8 Many rows (22 and more) 30.9 11.5 6.4 48.8 Few rows (16 and less) 44.7 13.2 7.0 64.9 Twisted rows 39.6 12.6 6.8 59.0 Straight rows 38.1 12.0 6.6 56.7 Cylindrical 38.2 11.8 6.2 56.2 Tapering 39.0 12.4 7.2 58.6 Close spaced rows 42.4 13.7 7.5 63.6 Open spaced rows 37.2 11.6 8.3 57.1 Rough indentation 35.3 12.6 7.5 55.4 Smooth indentation 46.4 11.8 5.4 63.6 Wide kernels 33.4 12.5 7.8 53.7 Narrow kernels 35.5 12.1 5.8 53.4 Deep kernels 27.0 11.4 6.7 46.6 Shallow kernels 44.2 12.9 5.5 62.0 Horny kernels 54.4 8.0 4.4 66.8 Medium horny kernels 39.4 11.8 6.1 57.3 Starchy kernels 36.0 10.1 7.2 53.3 Large germs 34.1 10.9 6.4 51.4 Small germs 41.6 12.4 6.1 60.1 High shelling percentage 30.2 11.5 6.5 48.2 Low shelling percentage 43.6 11.3 5.8 60.7 Heavy grains 32.3 12.6 7.4 52.3 Light grains 37.1 11.9 5.5 54.5 Heavy cobs 33.4 12.2 7.0 52.5 Light cobs 39.3 12.5 6.3 58.1 A brief inspection of the data will show that seed from short ears, light ears, ears with few rows, and ears of small circumfer- ence, germinated better than seed from ears of the opposite ex- treme types. However, it can hardly be assumed that these var- ious characteristics of size bear a direct relation to the viability of the seed. Each of them is in some degree merely an expression of the circumference or weight of the cob ; and one might expect a comparatively low germination in seed borne on a large, sappy Characters Connected with the Yield of the Corn Plant 17 cob, because of the unfavorable effect of a higher moisture con- tent. Some verification of this is found in the fact that seed from light cobs germinated 58 percent, while seed from heavy cobs germinated 52 percent. The data do not show a material difference in the germina- tion of seed from ears extremely variable in shape and in the form and spacing of the kernel rows. However, smooth, shallow, horny kernels, germinated better than rough, deep, and starchy kernels, respectively. Small germs sprouted better than large germs. It is possible that the treatment of the seed previous to the germination tests — late harvesting, 3 months storage in a crib, and several fumigations with hydrocyanic acid gas — may have affect- ed differently the viability of the various types. Certainly the viability of all types was very low as a result of this treatment. SUMMARY 1. Ears extremely characterized by deep kernels, narrow kernels or starchy kernels, had a slightly higher shelling percentage than ears of the opposite extremes. No other characteristics of the ear showed a significant relation to the proportion of grain. 2. Heavy ears, thick ears, deep-kerneled ears, and ears with a large number of rows, lost considerably more weight than ears of the opposite extremes, during a total drying period of 6 weeks. These characteristics are of course closely related to the size of the cob. Other characteristics of the ear showed no relation to the total loss of moisture. 3. In all types of ears more than 75 percent of the total shrinkage occurred during the first 4 weeks of a drying period of 12 weeks. Additional shrinkage was very slow over the following 8 weeks period. This indicates that when seed corn has been air- dried on racks or other devices for about a month, under climatic conditions similar to those of this experiment, it may safely be stored in a more convenient bulk. 4. Smooth kernels, shallow kernels, horny kernels, and ker- nels with small germs, showed a higher viability than kernels of the opposite extremes. No characteristic of the ear as a whole showed a relation to viability which may not be traced to the mois- ure content of the cob. Possibly the previous treatment of the seed influenced the relative viability of the different types. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 47 Localization of the Factors Determining Fruit Bud Formation (Publication Authorized August 26 , 1921 ) COLUMBIA, MISSOURI SEPTEMBER, 1921 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. EANSING RAY, P. E. BURTON, H. J. BEANTON, St. Eouis Joplin Paris ADVISORY COUNCIL, THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION F. B. MUMFORD, M. S., DIRECTOR J. C. JONES, Ph. D., LL. D„ ACTING PRESIDENT OF THE UNIVERSITY STATION STAFF SEPTEMBER, 1921 AGRICULTURAL CHEMISTRY C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. VV. S. Ritchie, A. M. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. Sieveking, B. S. in Agr. AGRICULTURAL ENGINEERING J. C. WoolEy, B .S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. E. F. Hopkins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale. B. S. in As* - - W. W. Swett, A. M. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. C. McBride, FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, A. M. O. W. Letson, B. S. in Agr. B. M. King, B. S. in Agr. A. C. Hill Miss Bertha C. Hite, A. B. 1 Miss Pearl Drummond, A. A . 1 *In service of U. S. Department of Agr 2 On leave of absence. RURAL LIFE O. R. Johnson, A. M. S. D. Gromer, A. M. E. L. Morgan, A. M. Ben H. Frame, B. S. in Agr. HORTICULTURE V. R. Gardner, M. S. A. H. D. Hooker, Jr., Ph. D. J. T. Rosa, Jr.. M. S. F. C. Bradford, M. S. H. G. Swartwout, B. S. in Agr. POULTRY HUSBANDRY' H. L. Kempster, B. S. Earl W. Henderson SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M. W. A. Albrecht, Ph. D. F. L. DulEy, A. M . 2 R. R. Hudelson. A. M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr. Richard Bradfield, A. B. O. B. Price, B. S. in Agr. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Sercretary S. B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsham, Librarian E. E. Brown, Business Manager culture, Seed Testing Laboratory. LOCALIZATION OF THE FACTORS DE- TERMINING FRUIT BUD FORMATION H. D. Hooker, Jr. and F. C. Bradford Belief in a relationship between slow growth and a fruitful condition in apple and pear trees has come down to the present with the approval of many generations of growers. Said John Lawrence 1 , in 1717, concerning the pear: “*****but yet for the sake of that noble Fruit which some Kinds produce by the Help of a Wall, it is worth while to humble him and keep him in Order. For which purpose****I sometimes plash the most vigorous Branches, cutting them near the place from whence they shoot, more than half through, which effectually checks its Vigour, and consequently renders it more disposed to make weaker Shoots, and form bearing Buds.” The chief concern of the older writers on fruit bud formation seems to have been the prevention of excessive growth. This was natural, since they were dealing chiefly with fruit gardens, manured and cultivated and consequently with trees growing luxuriantly. When fruit growing spread to the orchard the literary heritage from the garden survived and though there was an undoubted realization of the unfruitfulness of greatly weakened trees it is but recently that there has been a crystallization into definite phrases of this feeling that a certain amount of growth is necessary for fruit bud formation and that, within limits, fruitfulness and vegetative development are associated phenomena. SOME OF THE FACTORS INVOLVED The work of Klebs 2 , of Fisher 3 , of Kraus and Kraybill 4 , and of Hooker 5 has given some conception of the internal chemical factors connected with the initiation of fruit bud differentiation. Briefly stated, this seems to be associated primarily with carbohy- drate accumulation and in apple spurs, with starch storage in particular. However, even though carbohydrate accumulation oc- cur, fruit and differentiation does not take place if there be a lim- iting factor which seriously retards or altogether stops vegetative growth. The inference seems warranted, therefore, that the supply of water, of heat 6 , of nitrates or of any other essential nutrient 4 Missouri Agr. Exp. Sta. Research Bulletin 47 may so check growth and carbohydrate utilization that carbohy- drate accumulation results, if conditions be favorable for carbohy- drate manufacture ; any one of these factors may become limiting and prevent fruitfulness, though under field conditions the nitrogen supply seems to be the factor most frequently operative in this direction. When the nitrogen supply is plentiful, carbohydrate is usually found in the plant in small amounts, because it has been utilized in growth ; when the nitrogen supply is low, carbohydrate is usually found accumulated in relatively large amounts. In the two year cycle involving fruit bud differentiation one year and fruit formation the next, through which most apple spurs on fruitful trees usually pass, starch accumulation is associated with a rela- tively low nitrogen content during the period of fruit bud differ- entiation the one year and a practical absence of starch is associ- ated with an exceptionally high nitrogen content during the period of fruit setting the other year. ^ It should be pointed out that this inverse correlation between nitrogen and carbohydrate (particularly starch) content does not represent a relationship of fundamental importance for fruit bud differentiation. It is, in a sense, accidental, though it is common because nitrogen supply is most often the limiting factor determin- ing carbohydrate accumulation. In case some other factor, for ex- ample water supply, were operative in checking growth, it is clear that carbohydrate accumulation might take place even in the pres- ence of abundant nitrogen. In fact some such situation must ob- tain in those spurs on certain apple varieties which form fruit buds regularly every year. If fruit setting depend on the presence of a relatively large amount of nitrogen in the spur as Harvey and Murneek 7 suggest and if fruit bud differentiation depend on starch accumulation, then large amounts of nitrogen and of carbohydrates must be present almost simultaneously in these spurs. This situ- ation has been observed in spurs of Payne’s Late Keeper, a local variety in which a large percentage of the spurs are characterized by successive fruit bud formation. A sample of bearing spurs collected July 3, 1920, had 1.236 per cent nitrogen and 3.16 per cent starch. Comparison of these figures with the data published in Research Bulletin 40 of this Station shows that this nitrogen content is of the order found in spurs of other varieties during the spring of their bearing year and that the starch content is equal in amount to that found in those spurs of these varieties that are differentiating fruit buds. Localization of Factors Determining Fruit Bud Formation 5 THE RELATION OF GROWTH TO PERFORMANCE The fact that very little growth and very vigorous vegetative development are alike unfavorable for fruit bud differentiation suggests relationship between spur growth and spur performance in the apple. Roberts 8 found that in Wealthy and other apple varieties under certain conditions spurs of certain length growths showed the highest percentage of fruit bud differentiation and that both longer and shorter spurs showed lower percentages. To establish, for closer selection of samples for chemical study, the value of such an index to the probable performance of the in- dividual spurs under conditions obtaining in the trees growing in -- Man's Recreation, p. 49. 5th. ed. London, 1717. 2. Klebs, G. Proc. Roy. Soc. London 82 : 547-558. 1910. 3. Fisher, H. Gartenflora 65 : 232-237. 1916. 4. Kraus, E. J. and Kraybill, H. R. Oreg. Agr. Exp. Sta. Bui. 149. 1918. 5. Hooker, H. D. Jr. Mo. Agr. Exp. Sta. Res. Bui. 40. 1920. 6. Walster, H. L. Bot. Gaz. 69 : 97-125. 1920. 7. Harvey, E. M. and Murneek, A. E. Oreg. Agr. Exp. Sta. Bui. 176. 1921 . 8. Roberts, R. H. Wis. Agr. Exp. Sta. Bui. 317. 1920. 9. Hartig, R. Anatomie und Physiologie der Pflanzen, pp. 251-253. Ber- lin, 1891. 10. McCue, C. A. Del. Agr. Exp. Sta. Bui." 126, 1920. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 48 INVESTIGATIONS ON THE HARDENING PROCESS IN VEGETABLE PLANTS (Publication Authorized October 22 , 1921 ) COLUMBIA, MISSOURI DECEMBER, 1921 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL, THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY, P. E. BURTON, H. J. BLANTON, St. Louis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION F. B. MUMFORD, M. S., DIRECTOR J. C. JONES, Ph. D., LL. D., ACTING PRESIDENT OF THE UNIVERSITY STATION STAFF DECEMBER, 1921 AGRICULTURAL CHEMISTRY C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, A. M. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. Sieveking, B. S. in Agr. AGRICULTURAL ENGINEERING J. C. Wooley, B .S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edingf.r, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. E. F. Hopkins, Ph. D. DAIRY HUSBANDRY C. Ragsdale. B. S. in j\gr. . W. Swett, A. M. M . H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. C. McBride, FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, A. M. O. W. Letson, B. S. in Agr. B. M. King, B. S. in Agr. A. C. Hill, B. S. in Agr. Miss Bertha C. Hite, A. B. 1 Miss Pearl Drummond, A. A. 1 RURAL LIFE O. R. Johnson, A. M. S. D. Gromer, A. M. E. L. Morgan, A. M. Ben H. Frame, B. S. in Agr. HORTICULTURE V. R. Gardner, M. S. A. H. D. Hooker, Jr., Ph. D. J. T. Rosa, Jr., M. S. F. C. Bradford, M. S. H. G. Swartwout, B S. in Agr. POULTRY HUSBANDRY H. L. Kempster, B. S. Earl W. Henderson SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M. W. A. Albrecht, Ph. D. F. L. DulEy, A. M. 2 R. R. HudElson. A. M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr. Richard Bradfield, A. B. O. B. Price, B. S. in Agr. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crtsler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Sercretary S. B. Shirkey, A. M., Asst, to Director A. A. Teffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsham, Librarian E. E. Brown, Business Manager Hn service of U. S. Department of Agriculture, Seed Testing Laboratory. 2 On leave of absence. TABLE OF CONTENTS. Page Introduction 5 Review of Literature 5 The physical process of freezing in plants 5 Nature of the killing of plant tissue by cold 10 Result of water loss-desiccation 11 Injury to Plasma Membrane by water withdrawal 12 Protein precipitation through “salting out” 13 Protein precipitation by increase in acidity 13 Relation of water withdrawal from the cells to killing by cold 14 Factors influencing the water-retaining power of cells 15 Osmotic concentration and water-retaining power 15 Imbibition and water-retaining power 17 Relation of factors influencing water-loss by the plant as a whole, to hardiness 22 Statement of the problem 25 Experimental work 20 Materials used 20 Methods of hardening plants 26 Effect of hardening treatments on plants 28 Morphological differences in hardened plants 29 Effect of hardening treatments on rate of growth 30 Effect of hardening treatments on percentage of dry matter 31 Effect of hardening treatments on depression of freezing point 32 Effect of hardening treatment on ice formation in plants 37 Method of measuring the amount of water freezing in plant tissues . . 41 Effect of temperature on amount of water freezing in hardened and non-hardened cabbage leaves 42 Changes in amount of freezable water during the hardening process . 45 Influence of time of day on percentage of water frozen 47 Effect of watering plants with salt solutions on amount of easily frozen water in the leaves 48 Relation of amount of freezable water to percentage of dry matter and freezing point depression in garden plants 53 Rate of water-loss by transpiration in hardened and tender cabbage .... 55 Rate of dehydration in hardened and tender plants 58 Changes in Carbohydrates on hardening of plants 60 Formation of sugar by low temperature 60 Relation of sugar content to cold resistance 61 Methods of analysis — carbohydrates 64 Nature of water-retaining power in plants 66 Relation of pentosan content to cellular water-retaining power 68 Pentosan content in the hardening process in vegetable plants 70 3 4 Missouri Agr. Exp. Sta. Research Bulletin 48 Method of pentosan analysis 70 Pentosan content in garden plants 72 Pentosan content in plants watered with salt solutions 73 Rate of increase in pentosan content 73 Relation of hot water soluble pentosans to the hardening process 75 Factors influencing the imbibitional capacity of plant colloids 77 Acidity 77 Salts and Sugars 80 Summary 81 Conclusions S3 Applications 81 Acknowledgments 85 Bibliography 85 Plates 91 INVESTIGATIONS ON THE HARDENING PROCESS IN VEGETABLE PLANTS J. T. Rosa, Jr. This study was undertaken as one phase of a project on the transplanting of vegetable plants. The hardening process, whereby vegetable plants are made more resistant to cold and better able to withstand the hardships of transplanting from greenhouse or hotbed to the open field, is of great importance in the practice of growing certain vegetables which are customarily transplanted. In the pro- duction of early crops, hardiness also is especially important be- cause of the low temperatures to which transplanted plants are ex- posed upon their removal to the field in early spring. Furthermore, since the hardening process in vegetable plants results in a condition of acquired hardiness, developed rather quick- ly by subjecting plants to certain treatments, experiments with such material throw considerable light on the general problem of cold re- sistance in plants. This question, in connection with that of the nature of the process of killing of plants by low temperature, has received the attention of numerous investigators during the past one hundred years. Though much information has been accumu- lated, the whole problem is in a somewhat undefined state. It is the purpose of this paper to propose a theory comprehensive enough to explain satisfactorily the known facts as to the cold-resistance of living plants and to present data on the nature of the response of plant-tissues to treatments which result in increased hardiness. The injurious effects of temperature slightly above the freezing point on the growth of plants are not dealt with in this paper. REVIEW OF LITERATURE. The Physical Process of Freezing in Plants. — An early theory as to killing of plants by cold, advanced by Duhamel and Buffon 27 * in 1737, held that death was due to the rupture of the tissues, bursting of the plant cells, by the expansion of ice crystals form- ing within the cells upon freezing. •This and subsequent superscript numerals refer to literature cited in the Bibliography. NOTE. — Also submitted to the Faculty of the Graduate School of the University of Mis- souri as a thesis in partial fulfilment of the requirements for the degree of Doctor of Philos- ophy. 5 6 Missouri Agr. Exp. Sta. Research Bulletin 48 Geoppert 32 in 1829, found that ice formation upon freezing of plant tissue was not confined to the interior of the cells and con- cluded that the killing of plants by cold was not due to cell rupture. A few years later, Morren 73 substantiated Geoppert ’s conclusion in that he found no organ of the plant torn by freezing. He consid- ered that injury from freezing was due mostly, to the separation of air from the plant sap. In 1860, Sachs 105 using improved technique, observed that in the process of freezing, water was withdrawn from the cell and ice-crystals formed for the most part in intercellular spaces. In 1860, Nageli 89 showed by calculation, that the expansion caused by freezing all the water in the cell, would not be sufficient to cause a rupture of the cell-wall. Prillieux" in 1869, found that water was extruded from the cells upon freezing. Muller-Thurgau 74 found that ice formed within the cell to some extent, when the lowering of the temperature was very rapid, but in case of gradual cooling to the point of ice formation as in nature, the crystals were found exclusively in the intercellular spaces. Wiegand 28 noted similar results upon freezing Spirogyra and Nitella. Thus the find- ing of ice crystals within the cells by earlier investigators, who froze the plant tissue very quickly, is explained. Cavallero 19 confirmed the work of the German writers, as he found that cell rupture in winter was very rare, the cells themselves never freezing, though ice formation occurred in the intercellular spaces of both hardy and tender plants. Geoppert 32 noted that plants which were frozen to death lost water rapidly upon thawing. Sachs 106 observed that upon thawing, water remained in the intercellular spaces until reabsorbed by the cells or lost by evaporation. Under certain conditions considerable time elapsed before the water was reabsorbed and the protoplast re- gained its turgid condition. Prillieux 100 describes experiments on freezing pieces of potato and beet, showing that water was lost from these tissues upon thawing. He recognized also that water was lost from the tissues while still frozen, by evaporation from the sur- face of the ice crystals. Prunet 101 found that moisture is lost by evaporation from the surface of the leaf on thawing, rather than by normal transpiration through the stomata. Abbe 1 stated that as plant tissues were cooled, water exuded from the cells into the intercellular spaces, and after sufficient under- The Hardening Process in Vegetable Plants. 7 cooling, this water froze. The concentrated sap left within the cell did not freeze until cooled still lower. If the water is withdrawn from the cell before freezing in the intercellular spaces, it is important to find how this withdrawal takes place. Wiegand 131 offered two theories to account for cellular water loss upon freezing, ‘ ‘ extrusion ’ ’ and ‘ ‘ attraction. ’ ’ Extrusion . — This hypothesis is that the cell actively gives up water at low temperature by contraction and squeezing. Greeley 34 showed that cooling to near 0°C. caused Stent or to contract and become cyst-like. Under the same conditions Spirogyra became much plasmolyzed. Livingston showed that when mounted in oil, this plasmolysis was accompanied by extrusion of droplets of water. Wiegand thought that the most probable explanation of this method of water loss from the cell was by change in permeability of the protoplast to the sap solute. A recent report by Pantanelli 96 sup- ports this idea. In experiments with the pericarp of the mandarin cooled almost to the freezing point of this material (-6°C.) he observed a progressive increase in cellular permeability, as shown by rapid loss of water and exomosis of substances from the tissue. Osterhout 93 has shown that freezing as well as treatment by various anesthetics, greatly increases cellular permeability. Attraction. — Wiegand 139 considered his so-called attraction the- ory as the more probable explanation of water withdrawal from the cell. Thus in ordinary plant tissue Wiegand pictured the fol- lowing arrangement: (1) A film of pure, or nearly pure, water adhering to the outer surface of the cell wall, bordering on the intercellular spaces. (2) The inert cell-wall cellulose material filled with water of imbibition, which is continuous with that of the protoplast. (3) A more or less narrow strip of protoplasm adhering close- ly to the inner surface of the cell wall and containing water of im- bibition, continuous with that of the vacuole. (4) The vacuole, containing an aqueous solution of salts, sug- ars and other substances. Normally this system is in equilibrium. According to Wiegand, upon lowering the temperature below the freezing point, the film of pure water on the outer surface of the cell walls freezes first. The tendency will then be to restore equilibrium by drawing water from the interior of the cell to replace the surface film. This water will be drawn first from the cell wall, which in turn will draw on the 8 Missouri Agr. Exp. Sta. Research Bulletin 48 protoplast, which in turn will draw on the sap in the vacuole. The water of the vacuole is held by the force of solution alone, whereas the cell wall and protoplasm hold water by the stronger force of imbibition. If the temperature remains constant, this readjustment will continue until the force of crystallization is equalled by the in- creased force with which the remaining water is held within the cell. After equilibrium is established between the forces of crystallization and the water-retaining power of the cell, at any given temperature, no more water freezes unless the temperature is lowered further, thereby increasing the force of crystallization. However, since the force with which the remaining water is held increases rapidly with the progressive loss of water, Wiegand predicted that the amount of water frozen at each successive degree for which the temperature is lowered would be smaller and smaller. This was shown to be ap- proximately true by the experiments of Muller-Thurgau 74 with ap- ples, and the work of McCool and Millar 80 with green plants sug- gests the same conclusion. Bouyoucos 11 working with soils, found that little more water was frozen at -78 °C. than at -6°C. The foregoing hypothesis as to the conditions under which ice is formed in living plant tissue has been substantiated by work of Effect of Glucose Solutions on Cold Resistance in Sections of Red Cab- bage Leaves. Temperature Concentration of Solution. 2M M M/2 M/4 M/8 M/16 Water - 5.2°C. all living Vz cells alive - 7.8°C. all living 14 cells alive single cells alive all dead •ii.rc. all living y 2 cells alive single cells alive all dead -17. 3°C. all living y 2 cells alive single cells alive all dead -22.0°C. all living single cells alive all dead -32°C. Vo cells | alive 1 single cells alive all dead The Hardening Process in Vegetable Plants. 9 Maximow. 66 In extensive experiments with red cabbage and Trades- cantia discolor he found a marked ‘ ‘ protective ” action when sec- tions were frozen in solutions of salts, sugars, and other organic ma- terials, provided the substance used was not toxic and its eutectic point did not lie too near the freezing point. Although the con- ditions of Maximov/ ’s experiments cannot be duplicated in nature, his results are of interest. The following table, taken from Maxi- mow’s work, is typical of the results he secured. Evidently red cabbage cells, which ordinarily are killed at a little below -5°C., survive a temperature as low as -32°C. in 2-mol. glucose solution. Maximow concluded that this apparent protective action of the solution could not be explained by the depression of the freez- ing point, since the resistance to cold always increased with the strength of the solution much more rapidly than this depression. The degree of protection was found however, to be closely related to the eutectic point of the solution, substances having a high eutectic point showing no protective effect. Isotonic solutions of different substances with low eutectic points possessed nearly the same degree of protective action. Maximow found no relation between the rate of penetration of the protective substance and the degree of protec- tion afforded, and that just as much protective action was exerted by the various solutions when sections were immersed in them and frozen immediately, as when the tissue had been soaked several hours in the solution before freezing. (Hence there could have been no effect on cell sap concentration or in preventing precipitation of the cell proteins.) If we consider Maximow’s work in connection with Wiegand’s hypothesis of freezing, we have a condition differing from the usual, in that the film of pure water on the outer surface of the cell wall is replaced by a more or less concentrated solution. In the first place, this would lower the initial freezing point somewhat. More im- portant still, the fact that the cell is surrounded by a more or less concentrated solution should mean that in the process of water with- drawal and ice formation at any given temperature, a state of equi- librium between the ice-ciystal and the cell system would be reached sooner than in the case of cells not surrounded by such solutions, if the ‘‘attraction” theory of water loss as advanced by Wiegand be ac- cepted. Somewhat less water would be frozen at a given temperature in the cells of tissue immersed in salt or sugar solution. If the amount of water frozen per degree of temperature lowering becomes smaller and smaller, it would be necessary for a “protective” solution to 10 Missouri Agr. Exp. Sta. Research Bulletin 43 effect a very small reduction in the amount of water freezing at the lower temperatures to enable the cell to stand cooling several degrees below the usual death-point. Recent work by Yass (122) on bacteria leads to the same conclusion. He found a distinct protective action exerted by glycerine and glucose solutions on freezing bacteria, as shown in the following table. Vass’ Results on Freezing of Bacteria at -5°C. Strength of solution Percent of bacteria killed In glycerine In glucose 0.00 (water) 96 — 0.01% 92 98 0.05% 87 95 0.1 41 89 0.5 45 74 1.0 0 58 5.0 0 35 10.0 0 4 Vass concluded, in agreement with Maximow, that the protective action of these solutions was due to their power to keep a film of un- frozen water in contact with the outer layer of the protoplast, the plasma membrane. Nature of the killing of plant-tissue by cold. — From the fore- going review, the evidence appears conclusive that cell rupture cannot be the cause of killing of plants by cold, but that water-loss from the cells by ice formation in the intercellular spaces is an invariable accompaniment in such killing. According to Muller-Thurgau, 76 Mo- lisch 71 and others, death cannot be due directly to absolute cold, and there is little if any evidence of death due to shock or other re- action attributable to ‘ ‘ cold-rigor. ” Thus, both Muller-Thurgau 76 and Voightlander 123 showed that plant tissues could be undercooled several degrees below the freezing point without injury as long as ice formation did not take place. Wright and Taylor 122 have recent- ly shown that potatoes can be cooled several degrees below their freez- ing point and warmed up again without injury, provided no ice formation took place. However, jarring undercooled potatoes caused ice-formation to take place and resulted in typical frost injury. Chandler 20 found evidence that tender plants exposed to tem- perature slightly below freezing when the surface of the leaves was wet, killed to a greater extent than if the leaves were dry. This result is explained by Harvey's “injection" theory, according to which undercooled tissues are caused to freeze in spots where droplets of free water on the surface crystallize and inoculate the tissue just beneath with the growing crystals. The Hardening Process in Vegetable Plants. 11 These facts strengthen the view that killing by cold depends on ice formation, rather than on the effect of low temperature in itself. Just how death is caused by the freezing process is a question of interest. Four distinct theories have been advanced. ( a ) Direct result of water loss — “ desiccation — Miiller-Thur- gau 74 believed that death was the direct result of the water loss, that is, death ensues when so many molecules of water are withdrawn from the protoplast that its living structure is permanently destroyed. Wiegand 131 concurred in this hypothesis, with the additional sugges- tion that “probably every cell has its critical point, beyond which water withdrawal causes death.” Cavallero 19 attributed killing to the wilting upon thawing, due to rapid evaporation of melting ice in the intercellular spaces. He mentioned an opinion generally held by practical gardeners., that under conditions favoring slow thawing or slow evaporation, such as shade or moisture, severe injury to the plant might be prevented by the re-entry of water into the cells. However, Miiller-Thurgau 74 and later Molisch 71 found no difference in extent of killing, between rapid and slow thawing. Chandler 20 also concluded from a considerable number of experiments that the rate of thawing generally had no influence on death from freezing. We should distinguish here between the loss of water from the cell and its loss from the plant as a whole. If the cells are killed directly by loss of water on freezing, or if they are killed by changes taking place as a result of this water loss, then the rate of thawing would have no effect on the killing. However plants capable of standing some ice formation within their tissues, would take back more of this water if thawed slowly, whereas they might lose the most of it if thawed rapidly. This explains two things, the wilted condition often observed in frozen plants upon thawing, and the cumulative effect of successive freezing and thawing, whereby a fraction of the plant’s water content is permanently lost by the plant on each thawing. Nelson 90 thought that rapid loss of moisture was the principal cause of winter-killing of shrubs in high, dry sections. Kylin 56 in a recent study of the cold resistance of marine algae, concluded that death from cold was conditional upon actual formation of ice and that such death was primarily due to withdrawal of water from the cell. Matruchot and Molliard 04 described the successive changes in arrangement of the chromatin strands of the nuclei in leaf cells of the snowdrop subjected to freezing temperatures. They stated that water was withdrawn from the protoplast and nuclear material of the cell, and that this continued if the temperature was sufficiently low, 12 Missouri Agr. Exp. Sta. Research Bulletin 48 until these portions of the cells contained less water than the mini- mum necessary to vitality. They 63 also subjected plant tissue to freezing, to drying and to the action of solutions of high osmotic concentrations. They observed a marked parallelism in the effects of these treatments, hence they concurred with Molisch and Miiller- Thurgau in that death of the cell was due to rapid loss of water. Adams 3 working with moist seeds, observed that in freezing, water was drawn from the cells and solidified in the intercellular spaces and if the freezing did not go too far, upon melting the water was reabsorbed slowly, without injury having been done to the cells. Wiegand 129 made extensive observations on the freezing of leaves and buds. In buds in winter: “Ice was always found in broad, prismatic crystals arranged perpendicularly to the excreting surface and usually formed a single continuous layer throughout the meso- phyl of the scale or leaf, to accommodate which the cells were often separated a considerable distance. The cells near the ice mass having lost their water, were in a state of collapse, but upon thawing they reabsorbed the water and resumed their normal condition.” This was also true of evergreen leaves, in which he observed that ice crystals first lined the spaces of the spongy parenchyma, later filling these spaces and, in leaves of high water-content, the crystals fused into a sheet of ice completely separating the upper and lower por- tions. In observations on thawing of frozen leaf sections, Wiegand no- ticed in hardy tissues, not killed by the freezing, that upon thawing the water was drawn back into the cells, but in tender tissues killed by the freezing process, water was not drawn back into the cells to any extent. Pantanelli 94 concluded that the suffrance of each cell is directly proportional to the outgo of water during cooling. He also attaches great importance to the condition of the roots with reference to ready water absorption in determining whether or not plant re- covers from freezing. (b) Injury to the plasma membrane by water withdrawal . — Maximow 66 concluded as a result of an extensive series of experiments, wherein sections of plants were frozen in solutions of various salts and inorganic materials, that killing by cold is not due to low tem- perature as such, but to physico-chemical changes set up in the col- loids of the plasma membrane during ice formation therein. This is really a modification of Miiller-Thurgau ’s theory, limiting the in- jurious effects of water-loss to the outer layers of the protoplast. Chandler 20 also concluded from his exhaustive researches that The Hardening Process in Vegetable Plants. 13 “killing from cold is more likely a mechanical injury due to with- drawal of water from the protoplasmic membrane than an injury resulting from a precipitation of proteins. ” (c) Protein precipitation through “ salting out.” — Gorke 36 con- cluded that killing was due to irreversible precipitation of the pro- teins of the cell. He accounts for this precipitation by the greater concentration of the salts in the sap as water is withdrawn from the cell by formation of ice, since certain proteins are precipitated in strong salt solutions. He found that approximately l/ 3 of the pro- teins were precipitated in frozen cereal plants. Gorke found also that hardiness of certain plants bears some relation to the ease with which their proteins were precipitated. In the tender begonia he obtained protein precipitation at -3°C., in winter rye at -15 °C. and in pine needles at -40 °C. Schaffnit 110 also concluded that protein precipitation was the cause of death. He found that the proteins of rye plants grown in the open at low temperature were not as easily precipitated upon freezing as those of tender greenhouse plants. The effect of low temperatures on the hardiness of plants grown in the open was ascribed to a transition from less stable to more stable forms of the proteins by splitting. He found that he could prevent the precipitation of proteins from the sap of tender greenhouse plants by addition of sugar, to which he ascribed a protective action against protein precipitation and consequently against injury of the plant from cold, although it was not proven that these two are always related. Chandler 20 was rather disinclined to accept the idea of killing by “salting out” of proteins. He found that the hardiness of plants was increased by growing them in salt solutions, such as zinc sulphate, which is an excellent protein-coagulating agent. However, Chand- ler’s work on this point cannot be held to disprove the protein-pre- cipitation idea, since he showed no evidence that the protein-precipi- tating salts were taken up by the plant, or if they were taken up, that they existed in the plant in a form which would precipitate proteins upon concentration. However, the fact that Chandler did not find appreciable protein precipitation on freezing the extracted sap of apple twigs indicates that killing may not always be accom- panied by protein precipitation, although his technique on this point may be open to question. (d) Protein precipitation by increase in acidity. — Changes in color of plant sap due to change in reaction upon freezing are well 14 Missouri Agr. Exp. Sta. Research Bulletin 48 known. Gorke 35 noted an increase of acidity in sap upon freezing. He believed this was a factor in the precipitation of the plant pro- teins, since the acidity of the medium is important in determining the state of such colloidal materials. Harvey, 42 in a recent paper dealing with cold injury to cabbage plants, extended this theory. He found definite evidence of increased acidity as a result of freezing cabbage plant juice, by measuring the hydrogen-ion concentration before, during and after freezing to definite temperatures. He noted protein precipitation when the actual acidity was increased from Ph 5.65 to Ph 5.26. It is especially interesting to note that Harvey found a similar increase in the acidity of juice expressed from leaves exposed to wilting, though he does not state if the leaves were wilted beyond recovery. Harvey demonstrated that if phosphoric acid was added to the ex- pressed sap until the Hydrogen-ion concentration was increased as much as it would have been by freezing, a precipitation of the protein occurred, thus implying that the parallel effect of water loss by wilt- ing or by freezing and addition of acid, was protein precipitation and death. Harvey repeated Gorke ’s experiment on the precipitation of pro- tein from expressed sap by freezing. Samples of juice were taken from hardened and not hardened cabbage plants and frozen to -4°C., a temperature which would kill the non-hardened, but not the har- dened plants. It was found that 9.4 percent of the protein in the juice of the hardened plants was precipitated and 31.2 percent in the tender plants. Repeating the experiment and adding sufficient acid to change the reaction of the juice the same amount as it would be changed by freezing to -3°C. he found that 11 percent of the protein was precipitated in the juice of hardened, and 44 percent in tender plants. He also made complete analyses of hardened and tender cabbage plants, finding that of the water-soluble fraction of nitrogen about 35 percent was amino-nitrogen in hardened plants, and only 17 percent in tender plants, having about the same amount of water-soluble nitrogen. Harvey thought this increase in amino- nitrogen to be a very significant result of the hardening process, though he said it was not necessary that complete cheavage of the proteins to the amino acids should occur, to prevent their precipita- tion on freezing. Relation of water- withdrawal from the cells to killing by cold. — No matter which agency is chiefly operative in the actual freezing and killing process, they all depend on the withdrawal of The Hardening Process in Vegetable Plants. 15 water from the cell. Irreversible coagulation of colloids, such as protoplasm, is itself essentially a dehydration process. It is, then, by means of factors affecting water-withdrawal from the cell by ice- formation that the differential killing of plant tissues by low tem- peratures may be explained. Schaffnit 110 classified plants in three groups, according to their cold-resistance and ability to withstand desiccation. 1. Plants for which water is absolutely essential. This we take to include such plants as tomatoes, which are killed once extensive ice formation actually takes place. 2. Plants which withstand a certain degree of desiccation. These would be such plants as the cabbage which can survive a cer- tain amount of ice-formation in the tissues without injury. It is this group with which we are mostly concerned in discussions of harden- ing or cold-resistance. 3. Those which withstand complete drying — seeds, spores, etc. This classification can be taken to include all plants, except those which are killed by cold above the freezing point. Such killing is probably due to inability to carry on their normal metabolic func- tions at low temperatures, as suggested by Molisch, rather than to direct effect of cold. Relationship to cold resistance of factors influencing the water- retaining power of cells. — If the killing of plant tissue by cold is primarily due to water-withdrawal from the cells beyond a certain minimum point, then the difference between hardy and tender tissues may be ascribed largely to the relative water-retaining power of the cells in the two types of tissue. There are two main forces concerned in the water- retaining power of plant cells. (1) Osmotic concentration, due to sap solutes in the vacuole, and (2) Imbibition, a force exerted by some con- stituents of the cell wall, nucleus, plastids, and especially by the colloidal cytoplasm. The importance of either of these forces in the water-retaining power of cells may be influenced by various factors. Osmotic concentration and water -retaining power. — Since the freezing point of a solution is lowered in proportion to its molecular concentration, several workers have sought a correlation between coM resistance and the molecular concentration of the sap as measured by the depression of the freezing point. Lindley 01 in reviewing the work of Morron and others in 1852, 16 Missouri Agr. Exp. Sta. Research Bulletin 48 was probably the first writer to connect the depression of the freez- ing point of the sap with cold resistance. Chandler 20 directed much attention to the relation of osmotic concentration to hardiness, although he admitted that the force of imbibition may be the more important factor in the water-retaining power of plant tissue. He found in most cases that the hardier plants had the more concentrated sap. To explain the relation of a slight difference in freezing point depression to a considerable difference in hardiness, Chandler reasoned that, since in a solution containing one gram molecule the freezing point is -1.86° C., and in a M/2 solution, -0.93°C., in the latter solution at a temperature of -0.93°C. all the water would be unfrozen, at -1.86 °C. one half would be un- frozen, and at -3.72 °C. one-fourth would be unfrozen, and so on. If this held true for the water contained in a plant, the sap of which is equivalent to about one-half gram molecular concentration, we would then expect 75 percent of the water to be frozen at -3.72 °C. However, Chandler’s conjecture on this point does not apply in all cases since McCool and Millar found in their dilatometer experiments that nearly as much water is frozen at -4°C. in wheat plants having a freezing point depression of 1.107 °C. as in corn plants having a depression of only 0.578 °C. Ohlweiler 92 in studying the effect of a late spring frost on vege- tation at St. Louis, found that plants which showed the greater osmotic concentration of the sap were generally injured the least, although there were some exceptions. He found, for example, that in twelve species of Magnolia, the order of hardiness paralled the order of sap concentration fairly well. Harris and Popenoe 40 found that on the average, the hardier species of avocado had slightly the greater sap concentration. Lewis and Tuttle 59 working on evergreen leaves in Canada, found that in Picea Canadensis , the freezing point low- ering varied only slightly from October to April, the maximum lowering being in March. In the bark of Populns and the leaves of Linnaea and Pyrola , the maximum depression of the freezing point was also found to be in March, after the coldest weather was over. The freezing point depression was found to parallel the accumulation of sugars during the winter months, the maximum sugar content being found April 2nd., just before spring growth started. They found little correlation between cold resistance and sap concentra- tion, as measured by the depression of the freezing point. Pantanel- li 95 likewise, was unable to establish a relation between osmotic con- centration of the cell sap and resistance to cold. The Hardening Process in Vegetable Plants. 17 Salmon and Fleming 109 found no relationship between sap concentration and winter hardiness in several common cereal crops in Kansas. Thus on November 27th., hardy Kharkov wheat gave a freezing point depression of 1.230° C. and tender Culberson oats 1.199°C. On December 17th., the freezing point depression of the wheat was 0.935°C. and of the oats 1.260°C. They explain these results by the supposition that oats are less able to secure sufficient water from the soil to supply that lost by transpiration, the ground being frozen at the time of the second determination. This resulted in water-depletion in the oat plants, giving a higher cryoscopic value to their sap. Wiegand 131 thought osmotic concentration of plant sap to be of importance in relation to ice-formation at the inception of freezing only. Imbibition and icater-retaining power . — The term “imbibition” will be used in this paper in the general sense, as applying to the absorption of water by colloidal materials and the holding of water by finely divided solids by means of surface phenomena, such as adsorption, adhesion or molecular capillarity. De Candolle 75 (quoted by Lindley in 1855) formulated the fol- lowing laws of temperature in relation to plants : “1. The power of the plant to resist low temperature is in inverse ratio of the water content. “2. Hardiness is in direct proportion to the viscidity of the plant’s fluids. “3. Hardiness is in inverse ratio to the rapidity with which the fluids circulate. “4. Tenderness is greater in proportion to the size of the cells.” Considering that De Candolle had few or no experimental data from which to draw conclusions, and that he wrote many years be- fore the classical researches of Miiller-Thurgau, his views on the resistance of plants to low temperature are remarkably near present conceptions. Wiegand 131 considered that the force of imbibition was to a large extent the cause of the water-retaining power of plant cells. According to Pfeffer 140 this force increases with decreasing moisture content. Although Wiegand made no quantitative measurements, his theories were the result of keen observation and sound reason- ing and are of very great importance to an understanding of the differential killing of plants by cold. He pointed out that the water of crystallization in frozen plant tissue was practically pure, sepa- 18 Missouri Agr. Exp. Sta. Research Bulletin 48 rating from the other cell constituents upon freezing. The progres- sive dehydration of the cell by the withdrawal of water to form ice crystals, was thought by Wiegand to increase the combined forces of osmosis and imbibition holding the remaining molecules of water. He advanced the hypothesis that the degree of cold necessary to form ice was proportional to the force which held the w^ter in the tissues, which force (osmosis plus imbibition) was thought to depend largely on the water content. Wiegand believed that in succulent tissues of high water-content, most of the water would be frozen out near the initial freezing point and a smaller portion would be frozen in less succulent tissues. Wiegand 129 observed that no apparent ice formation took place in the buds of Quercus, Castanea, Hicorea, Juglans , and Fraximus, at -18 °C. The buds of these species were observed to differ from many others in which ice formation took place at a higher tempera- ture by: (1) lower water content, (2) smaller cells, (3) thicker cell walls. He considered that these factors favored the retention of cell moisture by a relatively greater force of imbibition than in buds lack- ing such characteristics and in which ice forms at a higher tempera- ture. Wiegand also observed that the ice crystals in frozen beets and potatoes were smaller near the periphery than in the center of these organs. The cells of the peripheral regions in these roots being smaller and poorer in water, were thought to have a greater capacity for retaining water against the formation of ice crystals. Recent work by Parker 97 strengthens Wiegand ’s hypothesis that decreasing water content increases the force of imbibition. He found that finely divided materials in suspension held a considerable amount of water as capillary surface films, and the force with which this capillary water was held increased rapidly with decreasing moisture content. That moisture content has a marked influence on the force of imbibition is indicated also by the work of Reinke, 139 who found that a pressure of sixteen atmospheres would squeeze water from a frond of Laminaria when the moisture content was 73 percent, but when the moisture content was reduced to 48 percent, it required a pres- sure of 200 atmospheres to extract water. If decreasing moisture content increases the force with which water is retained by plant cells, a direct connection is indicated be- tween such water-retaining power and cold resistance, for several investigators working with a wide variety of plants have shown that hardiness is usually associated with low moisture content. Thus, Lindley 61 recognized the fact that decreasing the moisture content The Hardening Process in Vegetable Plants. 19 tended to increase cold resistance and that the removal of some water in the “ripening process’’ made the plant’s tissues better able to withstand cold. Detmer 26 stated that such parts of plants as are poor in water withstand low temperature best. He found that air- dry seeds of Triticum and Pisurn germinated normally after exposure to temperature of -5° to -10° C., while turgid seeds were killed under the same conditions. Gorke 35 noted that the more hardy plants had the greater per- centage of dry matter and slightly lower sap freezing point. Schaff- nit 110 found a gradation in the amount of dry matter in different varieties of wheat in direct proportion to their resistance to low temperature. He concluded that high dry-matter content was cor- related with high frost resistance. Rivera 103 found that all cultural conditions which tended to increase the percentage of dry matter in wheat decreased the tendency to lodging and increased hardiness. Hedlund 44 found that under like cultural conditions, those varieties of winter wheat having a higher percentage of dry matter in autumn are generally more winter-hardy than those having a low percentage. He found also that cultural conditions that make for high percentage of dry matter favor winter hardiness. Hedlund attributed the high dry-matter content of hardy plants to their large carbohydrate con- tent. Shutt 114 found that a correlation existed between percentage of dry matter and hardiness in apple twigs. A set of samples gathered on the Canadian Experiment Farm in midwinter had moisture con- tents ranging from 45.1 percent in terminal parts of twigs of Yellow Tiansparent (hardy) to 51.59 percent in the same portion of the Blenheim Pippin (tender). He recommended the use of cultural practices to regulate the moisture content, as indicated by the degree of maturity in the fall. It is now a pretty well recognized fact that the ability of a variety of the apple to survive in Northern sections depends on its maturing thoroughly before winter — in other words, developing a condition of low moisture content and maximum water retaining power. Webber 127 and his co-workers observed after a very severe freeze in the citrus regions of California that trees and por- tions of trees which were dormant or inactive were much less injured than those actively growing and functioning. Trees which had been rather dry for some time also were more hardy than those recently irrigated while trees suffering badly from drought were injured worst. Batchelor and Reed 5 found that winter-injury of the distal end of the branches of the Persian walnut in California could be pre- 20 Missouri Agr. Exp. Sta. Research Bulletin 48 vented by bringing the trees to early maturity by with-holding water, followed with heavy irrigation during the winter. Johnson 51 found a marked seasonal increase in water content of peach buds in Maryland, correlated with the increased tenderness of buds in spring. The variety Greensboro had a lower water content than the Elberta, which is a tenderer variety. West and Edlefsen 128 also working on peach buds, pointed out that buds might escape injury from cold by under-cooling below the freezing point without ice formation, when the amount of moisture in the buds was small. Chandler 20 and more recently Carrick 18 found that apple roots which had been allowed to absorb moisture for several hours were injured by cold a little more than normal roots, whereas partial drying increased their cold resistance. Beach and Allen 6 found that drying apple twigs before freezing lessened the injury by cold. They also found that the hardier va- rieties of apples have the lower moisture content during the growing season but after prolonged freezes in winter, these hardy sorts may contain more moisture than tender varieties. In other words, the hardy twigs undergo a smaller water loss during freezing. Salmon and Fleming 109 performed an interesting experiment with greenhouse-grown cereal plants, which demonstrated that cold resistance may be increased by decreasing the amount of water in the tissues by slight wilting. Wheat plants were dug up, wilted for two or three hours, and exposed to freezing temperatures. Turgid plants killed much worse than slightly wilted plants at a temperature of -2 to -3°C. for 20 to 30 minutes. Chandler 20 compared the relative extent of killing by cold in turgid and wilted plants. He included in his experiments a large number of tender plants which are incapable of withstanding ice formation and which cannot be expected to show much response in the way of hardiness to any treatment. His experiments were made in summer, hence the killing at temperatures only slightly below freezing. Though Chandler concluded that on the whole, wilting does not increase cold resistance, yet the following table, taken from his data, indicates that under certain conditions, wilting may do so. In the case of lettuce, it seems that the wilted plants were killed the worst by slight freezing, -2°C. At the lower temperatures, how- ever, the percentage killed increases very rapidly in the turgid plants, and slowly in the wilted plants, so that the killing of turgid leaves considerably exceeds that of the wilted when the temperature of The Hardening Process in Vegetable Plants. 21 Effect of Wilting on Killing by Cold, Compiled from Chandler, p. 196. Plant Condition Temperature -2°C. -3°C. -* & C. -4.5°C. Lettuce turgid wilted 12y 2 % killed 47% killed 66.6% 55.5% 83% 62% Red Clover turgid wilted 17% killed 34% killed 100% 66.6% Rose Geranium turgid wilted 97% killed 60% 100% 100% Red Cabbage turgid wilted 65% 44% -4.5 °C. is reached. The same thing is indicated in the case of red clover. Chandler remarks that brief wilting does not increase the total amount of material in the cell sap which might function in hold- ing water in solution, yet it seems that the hardiness of the plants may be materially affected. Wiegand 131 states that the greater the water content, the thicker the film of water on the surface of an imbibing substance, such as the plant cell, and the weaker the force by which the outer layers of this film are held, hence more easily withdrawn to form ice. Parker 07 has furnished some experimental data, which substantiates Wiegand ’s suggestion. Kiesselbach and Ratcliff 52 in experiments with seed corn, found that death from freezing was directly proportional to the moisture content of the kernel and to the duration of exposure to cold. Seed corn maturing in the natural way was found to become cold-resistant progressively as the moisture content decreased. The following table taken from their data, illustrates the relation between moisture con- tent and killing by cold as measured by the germination of the seed. Kiesselbach and Ratcliff found that the temperature as which ice formation commences in the corn kernel depends very largely on the moisture content. Immature seed containing 60 to 80 percent mois- ture, froze just below 32 °F., whereas in air-dry seed, containing 18 percent moisture, no ice formation could be detected at -10°F. Usual- ly where ice formation took place in the seeds and they remained in the frozen condition 24 hours, the vitality was weakened or destroyed, but in some cases ice formation within the seed was not followed by 22 Missouri Agr. Exp. Sta. Research Bulletin 48 Relative Germination of Seed Corn of Varying Moisture Content After Exposure to Low Temperatures. (After Kiesselbach & Ratcliff) Temperature Percent moisture content of grain to which exposed Degrees F. 10 15 20 25 30 35 40 45 50 60 to to to to to to to to to to 15 20 25 30 35 40 45 50 55 65 32—28 100 85 75 71 69 — 33 0 24—20 100 96 77 67 13 12 12 6 0 16—12 100 88 34 12 0 0 0 0 0 8—4 100 98 47 7 0 0 0 0 0 0 0— -5 97 63 0 0 0 0 0 0 0 0 death. They show that air-dry seed are uninjured by low tempera- ture, and that ice-formation does not take place therein. The observations of G-orke, Schaffnit, Rivera, Hedlund, Shutt, Webber, Wiegand, Beach and Allen, West and Edlefsen, Batchelor and Reed and Johnson, indicate that individual plants, species or varieties having a low moisture content are usually hardier to cold than those having a high moisture content. The work of Chandler, Carrick, Beach and Allen, Salmon and Fleming, and Kiesselbach and Ratcliff indicates that reducing the moisture content of a given plant or part of a plant increases its cold resistance. This, it seems, may be partly accounted for by Wiegand ’s hypothesis and Parker’s recent work, in that the force with which water is held by plant cells increases with decreasing water content. Removal of some water by drying before freezing should increase the force with which the remaining moisture is held. In other words, if plant tissues become more cold resistant upon slight drying out, such increase in hardiness may be ascribed to the increased power of imbibition on the part of the plant’s cells. Relation of factors influencing water loss by the plant as a whole, to hardiness. — The foregoing discussion has shown the re- lation of some factors to the water-retaining power of plant tissue, as measured by the effects of low temperature. It is indicated that increasing the water-retaining power of the cell, either by increasing the concentration of its sap, or by increasing its power of imbibiton, or both, results in greater resistance to low temperature because of the increased force of crystallization necessary to withdraw the re- quired amount of water to cause death or bring about the changes which cause death. If the ability of the individual cell to retain some moisture when exposed to freezing is the significant point of differ- The Hardening Process in Vegetable Plants. 23 ence between tender and hardy tissues, then the plant as a whole may show the same difference in water-retaining power and resistance to water loss, but this does not imply necessarily that hardiness and drought resistance go together. Salmon 108 remarks that some hardy grasses thrive best in damp localities. In drought-resistant species, the plant as a whole may be protected against water-loss by morpho- logical differences in structure, such as special water storage tracts, few or small stomata, thick integument, bark, scales, xerophytic characters in general; yet the individual cell may possess little water-retaining power which would prevent the excessive withdrawal of water upon freezing. While a low transpiration rate due to morphological modifica- tions would undoubtedly be of great assistance to plants in withstand- ing injury from physiological drought, a low transpiration rate also may be associated with high water retaining power of the cells. Beach and Allen 6 observed a loss of four to nine percent in weight of apple twigs during a single week in January with a mini- mum temperature of -15°F. They found that in general the hardiest varieties are most resistant to the loss of water. Strausbaugh 117 found that coincident with the breaking of the rest period in semi-hardy varieties of the plum in midwinter, the moisture-retaining power of twigs and buds decreased rapidty, while in the hardy variety Assiniboine, which remained dormant until early spring, the water-retaining power remained constant. This is sig- nificant, since increased tenderness to cold, especially of the flower buds, follows the break of the winter rest. Sinz 115 concluded as a result of experiments at the University of Goettingen that those varieties of winter wheat which seemed able to prevent rapid transpiration, were among those most highly re- sistant to cold. Weaver and Morgensen 126 in Nebraska found that in winter the water losses of coniferous trees with their needles intact, are relative- ly no greater than are the losses from deciduous trees after leaf-fall. This indicates great water-retaining capacity in the foliage of coni- fers, most of which are very hardy. Some writers have likened hardy to desert plants because of their xerophytic characters, by which water loss is reduced to a minimum. Thus Schimper 112 states that desert plants frequently have a strong resemblance in their structure and habit of growth to those of polar regions, as would be expected if resistance to cold depended on the reduction of water loss to a minimum. What Schimper probably 24 Missouri Agr. Exp. Sta. Research Bulletin 48 had in mind was the form of injury due to physiological drought, where above-ground plant tissues are killed by desiccation resulting from their inability to obtain water from a frozen soil or through a frozen stem. Storber 118 states that “ winter leaves’ ’ of herbs are quite xero- phytic in structure, enabling them to survive the severe conditions to which they are exposed. He points out a fact that seems to have been hitherto overlooked — that the low water content and high os- motic concentration in hardy plants may insure to them more ready absorption of soil water. This would certainly be of great impor- tance to plants in winter, in overcoming physiological drought, as well as increasing the resistance to the direct effects of freezing. Dachnowski 22 observed xerophytic developments in plants exposed to physiological drought conditions in bogs. Modifications were found enabling certain plants to survive in bogs in spite of slow water ab- sorption due to toxicity of bog waters. The following are the chief modifications to which Dachnowski ascribes resistance to rapid w T ater loss in leaves of bog plants. 1. Reduction in size of leaves. 2. Thick- walled epidermis. 3. Cuticle, wax, and hairs. 4. Mucilaginous and resinous bodies in leaves and roots. Groom 36 stated that the function of mucilages and tannin in buds is to help hold the water in the young shoots. Chandler 20 found that the bud scales of the peach had no influence on the resistance of the embryonic tissue to low temperature, but that they served as protection against drying out by repeated freezing and thawing. Wiegand 130 recognized that loss of water from the plant might take place by evaporation from the ice masses in frozen tissues, and sug- gested that bark and bud scales serve as protection against such loss. As pointed out earlier in connection with the rate of thawing, pro- tection against such loss of water would be most important in tis- sues exposed to repeated freezing and thawing, as buds undoubtedly are in winter. In a number of recent experiments on the raspberry in Ne- braska, Emerson* found that by coating the canes with paraffin, winter-injury could be prevented. He observed that untreated canes killed only to the snow-line. Emerson’s results indicate that me- chanical protection against loss of water by the plant as a whole, * Emerson, R. A. Cornell University, Ithaca, New York, Personal correspondence with F. C. Bradford. The Hardening Process in Vegetable Plants. 25 may prevent the form of winter injury due to local physiological drought, wherein parts of plants exposed to repeated freezing and thawing and consequently to loss of water which cannot Le replaced because of frozen stem or frozen or dry soil, are eventually kb led by the progressive desiccation of the tops. This type of cold injury is distinct from the direct effects of low temperature, }et some of the factors which increase the water-retaining power of the tissues in the latter case may also be of importance in enabling the plant to withstand the former. Irmscher 49 attempted to correlate the cold resistance of certain peat mosses with their ability to withstand long drying out. He found that most species could stand a temperature as low as -20° C., but they were all killed at -30°C. He states that “no thoroughgoing parallel was found between cold resistance and ability to survive long slow drying.’ ’ However, he found that any particular species could be made more resistant to frost by previous drying out. Mosses growing in a dry location were found more hardy than the same species in moister places. Irmscher attributed to a 1 1 regenerative cell-complex” the means by which these mosses were enabled to survive both extreme cold and extreme drying. A higher osmotic concentration and greater cold resistance was observed in species of moss growing at low temperature. STATEMENT OF PROBLEM. The work of the earlier investigators shows that freezing to death of plant tissue is associated with water-withdrawal from the cells — the actual death process being due to (a) the direct effect of water subtraction on the protoplast, or (b) precipitation of proteins because of the increased acidity, or (c) precipitation of proteins due to increased salt concentration, or perhaps to other processes which have not as yet received attention. Regardless of the particular theory which may account for the ultimate killing of plant tissue by cold, the consideration that the primary factor is water-withdrawal logically suggests the following questions. In general, would not cold resistance be proportional to the water-retaining capacity of the plant cells? Since the force of imbibition increases with decreasing moisture content and since also cold resistance in plants increases with decreasing moisture content, does not cold resistance depend largely on the imbibitional fo' ce with which the cells retain moisture? Do hardy plant cells actually re- tain more moisture when exposed to freezing than cells of tender 26 Missouri Agr. Exp. Sta. Research Bulletin 48 plants? Do tender plants exposed to hardening treatments acquire an increased cell-water-retaining power, and if so, is this the main factor concerned in their increased cold resistance? Also, how is this increased water-retaining power acquired and what changes in the living plant are concerned therein? In order to answer these questions, the following experimental work has been undertaken. EXPERIMENTAL WORK. Materials used. — Most of the experiments were performed with the cabbage, as a representative of a type of plant which is capable of being hardened so as to withstand considerable ice formation within the leaves. Leaf lettuce, head lettuce, kale, cauliflower and celery were used to some extent. These also are plants capable of being hardened so that they can be frozen stiff without injury. The tomato was used as the principal representative of a type of plant which cannot be hardened so as to withstand ice-formation, but which is capable of hardening to the extent that the freezing point is lowered slightly. Other plants used of this type were pep- pers, eggplant and sweet potatoes. In each series of experiments plants of the same variety and age were used. Methods of hardening. — Series E . — The plants were kept in a warm greenhouse until nearly large enough for transplanting to the garden. The plants to be hardened were then removed to an open coldframe where they were exposed to temperatures near freezing during the night and to full sunlight during the day. This method of hardening was followed both in early spring and in late fall. Samples were gathered for analysis usually at intervals of 5, 10 and 20 days after the beginning of the hardening treatment, as well as from some of the original lot of plants which had been kept in the greenhouse under favorable growing conditions. The soil moisture supply was kept as nearly as possible the same for the plants in the greenhouse and those being hardened in the frames, so that temperature would be the principal limiting factor in their development. Series A . — The soil moisture for plants grown in a warm green- house was varied. As soon as the seedlings were well established after transplanting from the seed flat, a number of potted plants of uniform size were selected and divided into lots which were given The Hardening Process in Vegetable Plants. 27 different treatment only in so far as water supply was concerned. One lot, Al, was given liberal moisture — these plants were kept in rapidly growing condition and were always the tenderest plants in the experiments. Another lot, A2, was given moderate moisture, so that the plants grew at a moderate rate. Another lot, A3, was given just enough water to keep the plants growing slowly. They frequent- ly wilted somewhat in the middle of warm, bright days. This lot usually showed nearly the same degree of hardiness as those plants that had received the maximum degree of hardening in the cold- frame. A fourth lot, A4, was included in some of the experiments, these plants being watered liberally at first, then water was partially withheld for a week or ten days before samples were taken. Series B and C . — Plants were grown under uniform conditions in the greenhouse, in soils of different composition made up by mix- ing different proportions of sand and compost. Few data are re- ported on this series because it was found difficult to maintain uni- form moisture conditions in soils of such diverse texture. Also other factors, such as degree of root binding, were likely to be- come limiting before excess or deficiency of nutrients could exert much effect. However, it was definitely shown that growing plants in poor soils would increase their cold resistance, other conditions being the same. Such plants were smaller and grew more slowly than the more tender plants in the better soils. This series of ex- periments might have been more successful if the plants had been grown in a uniform soil-medium to which varying quantities of nutrient solution were added. Series H . — The treatment consisted of severely pruning the roots by running a knife close to the stem on one or both sides of the plant. This treatment checked the growth of the plants quite materially for a short time and increased the cold-resistance some- what. Series F . — A quite effective method of hardening was watering with M/10 solutions of various salts. The plants were grown under uniform conditions in a warm greenhouse and the test lots were watered with the various salt solutions whenever the soil became rather dry or whenever the plants wilted badly. In some cases, as under high transpiration conditions, the wilting point was reached while the moisture content of the soil was high. It is not altogether clear whether the hardiness resulting from these salt applications was due to their specific action, to a condition of mild physiological 28 Missouri Agr. Exp. Sta. Research Bulletin 48 drought, or to the toxicity of such concentrated solutions to the roots. This will be discussed in more detail later. EFFECT OF HARDENING TREATMENTS ON PLANTS. External appearance.— Cab bage . — Tender (wet-grown) green- house plants were usually about twice as large as those hardened by withholding of water, as shown by the relative green weights of A1 and A3 in Table 2. Plants hardened by withholding moisture were usually darker green, covered with heavy waxy bloom, with slight pink tints in the stem and petioles, but not as heavily pigmented as the coldframe hardened plants. The leaves were tough and leathery, in contrast to the brittle, crisp texture of tender plants. Cabbage plants hardened by exposure in coldframe were smaller and stockier than unhardened greenhouse plants and nearly always showed more or less pink pigment (probably anthocyanin) in the stems, petioles and leaf veins. Coldframe hardened plants were tough and leathery in texture. In most of the experiments the maximum degree of hardening by this method enabled cabbage to withstand a temperature of -5°C. to -6°C. for at least one hour, whereas non- hardened plants would be killed between -3°C. and -4°C. In a few experiments hardened cabbage withstood temperatures as low as -8°C. to -10°C. over night. The development of pink color, especially in the stems and peti- oles, was conspicuous in all hardened plants. According to Knud- son 55 the “work of Ewart, Overton, Wheldale and others indicates a close relationship between the sugar content of the plant and pig- ment production. ” Throughout Knudson’s experiments on the ef- fect of carbohydrates on green plants, a tendency to anthocyanin pro- duction was observed, plants fed on glucose and maltose (M/20 solutions) showing heavy coloration, which disappeared within a week when they were placed in diffuse light. These results are of special interest in connection with the large sugar content found in hardened plants, discussed later. Nicholas 91 was of the opinion that “the production (in leaves) of anthocyanin is correlated with the formation of organic acids. The connection known to exist between oxidation and pigmentation in- heres in the production of these acids, accompanied by the formation of the red pigment/’ The conspicuous development of the waxy bloom on cabbage plants has been considered by Harvey 42 of some importance in rela- tion to cold resistance in that it may permit the undercooling of the The Hardening Process in Vegetable Plants. 29 leaf several degrees below the freezing point. He suggests that it prevents the ‘ ‘ inoculation ’ ’ of the moisture in the leaf by droplets of water freezing on the surface. Cabbage plants hardened by other methods showed much the same changes as did those in the series mentioned above. In all cases hardiness was in direct proportion to the external changes noted. Wherever the growth of the plant was materially checked, even for a few days, hardiness was increased in proportion to the checking. Cauliflower and kale showed about the same changes on harden- ing as cabbage. Leaf lettuce. Both small potted plants and large plants ap- proaching maturity in the greenhouse and coldframe were used. The leaves become tougher, thicker and of more leathery texture upon hardening. Pigmentation was not conspicuous. When hard- ened by drying, the crinkling of the leaves was more pronounced and the color deeper green. Tomato. Leaves of hardened plants became very dark green with much pigmentation on the under side, were much smaller, tended to curl on the midrib; the stems and petioles became very heavily pigmented, tough and woody in texture. Hardening tomato plants in the greenhouse by any of the methods of checking growth had about the same effect on external appearance. The same was true of the coldframe-hardened plants, except that when hardening was long-continued at low temperature, the lower leaves turned yellow and fell, until the plant was nearly defoliated. This is prob- ably similar to the form of killing by temperatures above the freez- ing point noticed by Molisch and attributed by him to the inhibition of metabolism by the low temperatures. Morphological difference in hardened plants. — Schaffnit 110 was unable to find any structural differences in varieties of wheat vary- ing in degree of cold resistance. Salmon and Fleming 109 could find no difference in cell structure in hardy and tender varieties of cereals. On the other hand, Briggs 10 found that the cells were somewhat smaller in the pistils of hardy varieties of peaches. Walster 124 ob- served that in barley grown at 15°C., there was greater lignification of the xylem bundles than in plants grown at 20° C. This would make the plants grown at the lower temperature stiffer and stronger. To determine whether the hardening process affects the size of the cells in vegetable plants, sections were made of hardened and 30 Missouri Agr. Exp. Sta. Research Bulletin 48 not hardened cabbage and tomato leaves and the palisade cells meas- ured. Portions of young leaves which had made most of their growth during the process of hardening were used in each case. Hence the differences in the cells here reported do not represent an “ acquired ” condition, but differences in development between plants under favor- able growing conditions and those subjected to hardening. Portions were taken from corresponding locations on leaves of about the same size, killed and fixed in the usual way. Transverse sections were made with the rotary microtone, mounted, stained and measured. In the tender tomato leaf numerous large air spaces were ob- served, while in hardened leaves the cells were more compactly ar- ranged and filled with starch grains. Starch grains were less plen- tiful in hardened cabbage leaves, but the compactness of the cellu- lar arrangement was marked. Table 1 gives the measurements in two dimensions of the palisade cells and the cross-section area com- puted therefrom. These data are the averages obtained from meas- urements of several different sections. Table 1. — Measurements of Leaf Palisade Cells in Plants Hardened and not Hardened. Thickness Thickness Thickness Width Length Area. of whole of pali- of paren- of of sq. m- leaf sade chyma cells cells Cabbage Not hardened . Hardened by- ....291m 134.5m 106. 3m 19.1m 36.3m 694.0 drying in g. h. ... .269 127.9 106.3 19.4 27.8 538.8 Not hardened . . 274 136.2 102.1 20.9 36.9 772.0 Hardened in coldframe ....312 JL68.6 118.0 19.9 31.1 619.5 Tomato Not hardened . 196.6 ! 76.2 76.2 21.0 57.5 1201.5 Hardened . . . . ....133.7 i 55.6 68.0 1 14.1 44.8 630.1 Judging from the data presented in Table 1, hardened plants are characterized by somewhat smaller and more compact palisade cells than are non-hardened leaves of the same sort. In tomato, leaves from plants given hardening treatments are considerably thinner than tender leaves, however, cabbage leaves hardened in coldframe gained in thickness. Effect of hardening treatments on rate of growth. — The growth of plants subjected to any of the hardening treatments was checked in proportion to the intensity of the treatment. Data are presented in Table 2, on samples gathered from lots of the same age, grown The Hardening Process in Vegetable Plants. 31 under otherwise uniform conditions, except for the various harden- ing treatments. With the average green weight of the plants as the criterion, it is evident that hardened plants are much smaller, indi- cating the extent to which their growth had been checked. Thus, in the series gathered March 12, 1920, wet-grown greenhouse cabbage plants (tender) averaged 23.1 grams, plants hardened by partial withholding of moisture averaged 6.8 grams and plants hardened in coldframes three weeks averaged 7.67 grams. The differences in dry weight are not so great, since the smaller and hardier plants pos- sessed a larger percentage of dry matter. In the tomato, the rate of growth could be roughly measured by the increase in height from week to week. Accordingly a number in each of the lots subjected to the various treatments were measured each week. The retardation of growth, when any of the hardening treatments became operative, is shown in figure 1. Effect of hardening treatments on percentage of dry matter. — The data given under percent of dry matter in Table 2 indicate con- siderable increase of dry matter in all of the experimental lots of plants exposed to hardening treatments. Conversely, the water con- tent decreased in hardened plants, roughly in proportion to the ex- tent to which their growth was checked by the hardening treatment. 32 Missouri Agr. Exp. Sta. Research Bulletin 48 The possible significance of decreased water content in relation to the water-retaining power of the cell when exposed to water-loss by freezing has already been indicated by Wiegand and more recently by the work of Chandler, Salmon and Fleming, Carrick and indi- rectly by Parker. It may be repeated here that decreased water con- tent would be associated with increased force of imbibition, and with increased concentration of the cell sap, which forces tend to retain water in the cell during freezing. It is realized that the total loss in weight upon drying of leafy tissue does not truly represent the actual water content of the plant, but the difference is probably so small that this loss is taken as the moisture content throughout these experiments. Effect of hardening treatment on depression of freezing point. — In several experiments, the freezing point depression of the expressed sap of leaves of hardened and non-hardened plants was determined with the usual Beckman’s apparatus. Potted plants from each ex- perimental lot were brought into the laboratory to insure having fresh tissue for each determination. All of the leaves were taken from two or three plants and ground. The sap was then squeezed from the macerated pulp and duplicate or triplicate freezing point determinations were made at once. The data given in the column “Depression of the Freezing Point” in Table 2 show that the de- pression was somewhat greater in the hardened plants, indicating greater osmotic concentration of their sap. Similar data have been obtained by Chandler and by Harvey, working on the same sort of material, hence it was not deemed worth while to make a larger number of these determinations. The differences found here in freezing point depression are somewhat greater than those obtained by Chandler 20 and much greater than those reported by Harvey 42 for hardened and not hardened cabbage. This is due perhaps to the extremes in the treatments used in these experiments. Heavy water- ing made Series A1 somewhat more tender than ordinary non-hard- ened plants and Series A3 attained maximum hardiness through the application of the minimum amount of water to keep the plants from wilting. It may be pointed out that the increased sap concentration in hardened plants is due probably to the combined effect of the follow- ing factors: (1) Decreased total moisture content. (See Table 2). (2) Increase in the amount of sap solutes. Numerous investigators have found an increase of soluble sugars in plants exposed to low The Hardening Process in Vegetable Plants. 33 72 72* £ 2 72 72 tH © tH < CO w 5 £ J ” 3 m d 3 tH ^ 0 O T^ d 0 JS d 0 A H tH d 8 fu T3 o O 72 aS TO +J -d 0 cq 63 tH © i +J 4J "tn d o -^1 1 H ^ (!) © © w fn Q ctf a U CO ° i •2, 03 c3 c3 ® to 'd V os c3 p tn T3 d -2a tH d tH d •i—j £.S c3 'd § 0) > i +j 02 © © 1 © "“ 1 d d d © H ^ d d — © >» tH fH ^ d S3^.^ © tn H-> d >2 >> 2 ►» d s tH d < S c5 TO © © b s.2, •d C .2 .2 -3 d S’ -d © ■s§i d3 2 ,rH d3 hhi be d --5 d be-- 5 ~ be _bfi be ^ OQ +j Ch « ^ •(— > 6 o o S3 rd P cc ^ 0 CO ZZZ W rn rn o T UJ 17 7 S3 bfl O g dd K 76 N 72 CD 5 o o lO LO co 00 LO o co oo co co co cq 02 t- 0 © 02 S CO OO lo H< t-H OOOH< Tfl CO CO CO O 10 O 02 D- t> O t-H cqo CO cq 02 cq cq LO co ■< 63 ■j y 2h z | oo oo od O T-i t-H rH d d cq’ rH rH © rH d 02 ’ rH rH T-i Cq’ rH r- 1 rH i-i cq’ rH rH cd cd rH 1 — 1 cq’ rH rH t-H 72 © oo b0 d © oo o c-* cq c- o o rH CO 00 cq 0 Tt< O CO CO IO CO £ cS >> d ^ <70 LO CO CO rH fc- LO t> 02 cq 0 00 02 LO CO rH LO LO a es u t- t; OO CO 00 00 rH lo in oo CO 02 CO CO 00 cq 02 O rH C— 63 O > z fc « © 'd 0 > ooh © T-i odd tH t-H rH d d ri d cq' d T-i d d ® H CO 63 S5 N £ © ti bO S3 d o o o o oo c— 0 LO 0 0 0 iO t- oj © c3 oO -r o T LO cq Tt< oo o 00 0 00 t— CO cq co 0 t— 0 00 K £ I> I> rH cq o cq o ^ cq CO 02 t— cq co C2 TJT CO CO rH Th © t-i ** > be . « ? 02 t> CO oo d ©in© cq’ cd d c- d cd t-i t- t-i t-i rH cd S ON EZIN( cq rH rH cq rH 1— 1 pH M GO 02 05 O 02 O 02 02 O O 0 0 02 02 O 02 O 02 O 02 02 Z PS 0) t-H t-H cq t— i cq rH rH Cq cq cq cq rH H Cq t— 1 cq rH Cq rH T— ( 63 fc m */■? 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Date I Average I Average j % of I Depression 36 Missouri Agr. Exp. Sta. Research Bulletin 48 The Hardening Process in Vegetable Plants. 37 temperature and, as shown later, there is an increased sugar con- tent in hardened vegetable plants. (3) Increased amount of water held in the absorbed state by the cell colloids. Since this absorbed water is probably nearly pure, the sap solutes of hardened plants are held in solution by a smaller volume of water — hence the greater concentration. Chandler 21 found that a large part of the depression of the freezing point in plant sap was due neither to sugars nor to electro- lytes. Recent work by Parker 97 showed that finely divided material in suspension exerted considerable influence on the freezing point and that this depression increases rapidly with decreasing moisture content. Though Parker ’s work was done with soils and dried, finely ground inorganic colloids, it may be supposed that the organic col- loidal particles of plant protoplasm have the same property of de- pressing the freezing point. Parker attributes the lowering of the freezing point by finely divided material to the force of “adhesion” by which films of the liquid are held on the surface of the solid ma- terial. The freezing point depression caused by the finely divided material decreases almost to zero in presence of high moisture con- tent. He explains this by the suggestion that as the amount of liquid increases some of it becomes so far distant from the solid par- ticles and is so weakly held that no depression of the freezing point occurs. This is very nearly the same as Wiegand’s theory as to the holding of water in plant cells by “molecular capillarity.” It may be that the increase in colloidally -held water alone would account for much of the depression of the freezing point in hardened plants. Therefore it may be said that the apparent increase in osmotic con- centration of the sap in hardened plants is merely coincident with the state of being hardy, rather than a cause of it. EFFECT OF HARDENING ON ICE FORMATION IN PLANTS. Previous investigations on freezing of plant tissue have shown that water is drawn from the cells to form ice in the intercellular spaces. Some plants are killed once this occurs. Others are capable of withstanding some ice formation, but are killed at lower tempera- tures. Unhardened cabbage plants are killed by freezing at -3°C. to -4°C. The same plants after being subjected to a hardening treatment, may withstand a temperature of -5°C. to -6°C. or even a few degrees lower. Miiller-Thurgau 74 has shown that by no means all of the water freezes in plant tissue when exposed to temperatures well below 38 Missouri Agr. Exp. Sta. Research Bulletin 48 the freezing point. His method of measuring the amount of ice in frozen plant tissue was based on observing the cooling of water in which tissue frozen to a definite temperature was placed, calculating that 80 calories are required to melt one gram of ice. In this way he was able to show that the amount of ice in an apple increased as the degree of freezing increased. The following table is taken from his data on frozen apples : at -4.5°C. percent total water frozen = 63.8 at -7.3 percent total water frozen = 68.2 at -8.0 percent total water frozen = 72.4 at -13.0 percent total water frozen = 74.4 at -14.8 percent total water frozen = 77.4 at -15.2 percent total water frozen = 79.3 It appears from these data that the amount of water frozen at each successive fall in temperature decreases fairly regularly until -13 °C. is reached, but below that temperature the rate of ice forma- tion increases somewhat. The latter temperatures are far below the killing point for apples, which may affect the results, and Miiller- Thurgau’s technique may be open to some experimental error. Miiller-Thurgau also made some interesting determinations of the time-rate of ice formation. Kohl-rabi leaves exposed to a tem- perature gradually declining from -5°C. to -8.25 °C. froze at the Fig. 2. — Relation of time to rate of ice formation in 100 grams kohl-rabi leaves (ar- ranged from Mttller-Thurgau’s data). The Hardening Process in Vegetable Plants. 39 end of 13 minutes, when the temperature of the leaves was only -4.3 °C. and that of the surrounding air was -7.3 °C. In the first half minute of freezing 0.69 grams of ice formed in 100 grams of leaves. In the next minute, 2.0 grams, the following two minutes, 1.5 grams per minute, and the next minute, 0.8 grams froze. There- after the amount of ice formed per minute gradually decreased un- til, at the end of one hour of freezing, 41.32 grams of ice had formed in 100 grams of leaves, probably a little over 50 per cent of the total water content. This experiment is illuminating as to the re- lation of the time factor to freezing of plants. It has been observed that in plants, such as kohl-rabi, which can withstand some ice for- mation, injury at a temperature near the death point is proportional to the duration of exposure. This fact has been observed frequently in experiments here and Harvey 42 presents in his paper an excellent series of photographs illustrating the same thing. Figure 2 plotted from Miiller-Thurgau ’s data, illustrates graphically the rate of ice formation. We may regard the progressive increase in total amount of ice and the decrease in amount frozen per minute as being due to the balanced action of the force of crystallization and the water-re- taining power of the cells, the rate of ice-formation approaching zero as a limit. Foote and Saxton 28 used the dilatometer in experiments on the freezing of inorganic colloids and were able to show that in such materials water existed in three forms, viz., free water, capillary absorbed water and water of chemical combination. Recently, Bouyoucos 10 and McCool and Millar have made use of the dilatometer to measure the amount of water freezing in soils, in plants and in seeds. McCool and Millar in their latest report, state that the amount of water freezing in plants at -1.5 °C. decreased as the con- centration of the sap increased (as measured by the freezing point method). At -4°C. the amount of water freezing was considerably more than at -1.5°C. and its correlation with the freezing point of the sap almost disappeared. The following figures rearranged from their report illustrate this : Crop-plant Date Freezing point depression Amount of water freezin; in 5 grams of leaves. At -1.5°C. At -4°C. Wheat Nov. 14 1.107°C. 0.40cc. 2.65cc. Rye May 17 1.030 .86 2.40 Rye Nov. 24 .928 .90 2.50 Sweet clover Nov. 24 .906 1.22 2.82 Red clover May 15 .780 1.70 2.70 Corn June 10 .578 2.10 2.90 40 Missouri Agr. Exp. Sta. Research Bulletin 48 Unfortunately they did not express their results as percentages of the total moisture content in each of the different plant tissues, hence it is impossible to draw very definite conclusions. Also noth- ing is stated as to the source of the material, whether greenhouse or field grown. One interesting point to be noted here is that wheat and rye, which one would expect to be more cold-resistant than corn or clover, show a smaller amount of water frozen at -1.5 °C. and somewhat less at -4°C. It is rather surprising that such decided variation in sap density made so little difference in the amount of water freezing at -4°C. In another series of experiments with corn and barley McCool and Millar 80 showed that varying the soil moisture content affected the water relations in the plants. Freezing point depression and the percentage of moisture in the tops decreased slightly in plants grown with 15.53 percent soil moisture, as compared to those grown with 23.29 percent soil moisture. The amount of water freezing at -2.5°C. was decreased somewhat, and the amount freezing at -4.5°C. was decreased considerably in plants grown on the soils of lower moisture content. Hibbard and Harrington 45 found that the freezing point of the sap of roots and tops of corn plants fell regularly as the moisture content of the soil in which the plants were grown was decreased. The following data from their work illustrate the relation of soil moisture to freezing point depression of sap. Percent moisture in soil Freezing point depression of tops of roots 31 1.835°C. .492°C. 23 1.920 .600 16 2.027 .647 13 2.120 .942 11 2.204 .995 McCool and Millar 80 studied the effects of varying the concen- tration of the soil solution, when the moisture content was kept con- stant. They found a progressive increase in the freezing point de- pression of the tops and roots of plants grown in the greater con- centrations, but the amounts of freezable water showed little varia- tion. Earlier experiments by the same authors 78 showed that the freezing point depression of both tops and roots varied in the same direction as the concentration of the soil solution in which the plants were grown but not in proportion to it. They also studied the effect of varying the soil moisture content, keeping the concentration of the soil solution constant. Unfortunately, in their experiment The Hardening Process in Vegetable Plants. 41 the plants on the high moisture soils took up a larger quantity of the nutrient salts so that the concentration of the soil solution soon became less than in the low moisture soils. Therefore, it remains undetermined whether the effects of wet and of dry soils on freezing point depression and amount of freezable water in plants grown thereon are due to the variation in the water supply, or to variation in concentration of the soil which is involved, or to both. Method of measuring amount of water freezing in plant tissues. Though Miiller-Thurgau was able to obtain considerable data on this subject by measuring the latent heat of ice in frozen tissues, his method is laborious and perhaps open to some experimental error. The dilatometer method, as described by Bouyoucos 11 , presents great advantages in its directness, simplicity, and accuracy for measure- ments at different degrees of freezing. The use of the dilatometer is based on the fact that one gram of water increases approximately one tenth of its volume upon freezing. It has been used in this work essentially as described by Bouyoucos, and by McCool and Millar. A definite weight (4-6 grams) of fresh leaves, is placed in the bowl of the dilatometer, which is then filled with petroleum ether (boiling point 63°C.). The dilatometer is then stoppered with a rubber cork through which a thermometer is placed, so that the bulb is in contact with the leaf tissue and the scale convenient for reading. It was found at the beginning of this work that quicker and more accurate results can be obtained with plant tissues, by placing the loaded dilatometer in crushed ice for 15 to 20 minutes, to lower the temperature of the whole mass slowly and evenly to the neighborhood of 0°C. After this preliminary cooling, the dilatometer is plunged into an ice and salt bath, mixed in such proportions that the tem- perature is slightly below that which is desired in the dilatometer. Usually the temperature of the plant tissue in the dilatometer lags slightly above that of the freezing mixture. The dilatometer must be kept perfectly still after it is placed in the bath in order to secure uniform under-cooling of the plant tissue, but the freezing mixture can be gently stirred so as to keep all parts of the bath at uniform temperature. At first the column of petroleum ether in the graduated side-arm of the dilatometer falls rapidly due to the contraction of the contents on cooling. It is usually necessary to add a little more ether to bring the column up to the point on the scale where it can 42 Missouri Agr. Exp. Sta. Research Bulletin 48 be read easily. When the thermometer indicates that the contents of the dilatometer have been desired temperature for several minutes, the position of the column in the side-arm is read, then solidification of the under cooled water in the plant tissue is caused by tapping the dilatometer against the sides of the bath. As solidification of the water takes place, the column rises in the side arm, slowly at first, then more rapidly and then quite slowly for several minutes. It usually takes 5 to 10 minutes to es- tablish equilibrium, indicating that all the water is frozen which will freeze at that temperature. When the column of ether in the side arm becomes stationary, the reading is taken. The amount of expansion as a result of the freezing is the difference between the readings before and after freezing. In this work the side arm tube was graduated to 0.01 cc. and readings could be made to 0.005 cc. The expansion on freezing multiplied by 10 gives the number of cc. of ice formed. In some of the earlier experiments separate samples were used for moisture and dry matter determinations. Later it was found that these determinations could be made on the dilatometer sample after the freezing experiments had been performed. The water content of the tissues being known, the percentage of the total water frozen can be calculated by dividing the number of cc. of ice formed by the total water content of the sample. Effect of temperature on amount of water freezing in hardened and non-hardened cabbage leaves. — Cabbage leaves were used in most of the experiments, since these were available in varying de- grees of hardiness. It was found very difficult to secure rapid crystallization in hardened cabbage leaves at a temperature higher than -3°C., so that point was taken for the minimum reading. On the other hand, leaves could seldom be under cooled below -6°C. without ice formation, so that point was taken as the maximum limit of freezing in most of the experiments. Dilatometer determinations were made a number of times with each class of material under ex- periment. These determinations were distributed over a period of sev- eral months. The samples were taken at different times of day, but in each series samples were taken at the same time of all the different types of material which were being compared. Table 3 gives the results secured with leaves of non-hardened greenhouse cabbage plants thoroughly hardened in coldframe (9-12 days) and of plants har- dened in the greenhouse by partial withholding of water for two weeks or more. Each figure represents an average of several deter- minations, the individual determinations sometimes varying as much The Hardening Process in Vegetable Plants. 43 as 10 percent due to slight differences in the material and to the hour at which the samples were taken. Table 3. — Percentage of Total Moisture in Cabbage Leaves Frozen at Different Temperatures. Previous treatment of plants Percent of dry matter Percent of total moisture freezing at -3°C. -4°C. -5°C. -6°C. Wet-grown greenhouse plants, (tender) . . . 10.29 49.9 75.2 82.1 84.2 Dry-grown greenhouse plants, (hardy) .... 12.7 27.2 48.9 62.6 71.0 Hardened in coldframe 9-14 days, (hardy) . 14.67 29.8 49.6 : 58.7 64.°. Considerably more water froze in the tender plants than in the hardened at each temperature. The material used for the dry-grown hardy plant determinations perhaps was not quite so uniformly hardy in its nature as that of the two other types. The outstanding feature is the progressive in- crease in percentage of total moisture frozen as the tem- perature is lowered. This is brought out clearly in Figure 3 plotted from the data in Table 3. The increase becomes less and less for each degree of tempera- ture lowering. Thus in the case of the tender cabbage, 26.3 percent more water freezes at -4°C. than Fig. 3. — Relation of temperature to percent of total moisture freezing in tender and hardened cabbage leaves. at -3°C., 6.9 percent more freezes at — 5°C. than 44 Missouri Agr. Exp. Sta. Researcpi Bulletin 48 at -4°C. and 2.1 percent more freezes at -6°C. than at -5°C. In the coldframe hardened plants, 19.8 percent more water is frozen at -4°C. than at -3°C., 9.1 percent more at -5°C. than -4°C., and 5.6 percent more at -6°C. than at -5°C. Table 3 shows that the percentage of total moisture frozen at a given temperature is less in the hardened plants. Table 3A, constructed from the same data, on the basis of 100 grams fresh tissue, shows that the actual amount of water remaining unfrozen is greater also in the hardened leaves although there is a smaller total moisture content in such tissues than in tender leaves. Table 3a. — Amount of Water in 100 Grams of Cabbage Leaves Remaining Unfrozen at Different Temperatures. Treatment | 1 Percent dry matter I Percent moisture Grams water remaining unfrozen at -3°C. -4°C. -5°C. -6°C. Wet-grown greenhouse plants, tender ; 10.29 89.71 34.9 22.3 16.1 14.3 Dry-grown greenhouse plants, hardy i 12.70 87.30 63.5 44.6 32.6 25.3 Coldframe, hardened for 9-14 days ; 14.67 85.33 59.9 ! 42.9 35.2 30.4 Since the percentage of the total moisture which freezes at each temperature is materially less in hardy than in tender plants, and since the actual amount of water remaining unfrozen is greater in the hardy than in the tender plants, we may safely assume that the cells of the hardened plants possess a greater power to retain water when exposed to freezing. Although the amount of water frozen increases with the lowering of the temperature, we may as- sume that whatever the nature of the water-retaining force, it is overcome in successively smaller increments by the force of crystalli- zation as the temperature is lowered. The percentage of water re- maining unfrozen in the hardened leaves is approximately a logarith- mic function of the temperature. The hardiest plants used in this experiment probably could have been killed by long exposure to -6°C. to -8°C. However, it may be predicted from the rate of increase in the amount of water frozen at the lower temperatures, that if in some way the water-retaining power of the cells in these plants was increased slightly, a much lower temperature could have been sustained. Maximow has shown that sections of cabbage leaves which were injured at -5.2°C. when The Hardening Process in Vegetable Plants. 45 frozen in water, successfully withstood a temperature of -32° C. in 2-mol. sugar solution. Changes in amount of freezable water during the hardening process. — Harvey 42 stated that cabbage plants kept at 3°C. for 24 hours showed slightly increased hardiness and at the end of five days a considerable degree of hardiness was developed. In the present experiments, absolutely controlled conditions were not available ; however, it is generally considered that about two weeks’ exposure of greenhouse-grown cabbage plants in the open coldframe during March will bring about maximum hardening. To study the relation of the amount of freezable water to the hardening process, lots of tender cabbage plants were removed from the greenhouse to the coldframe at intervals and dilatometer determinations made on the leaves of these plants which represented progressive degrees of hardening. The results are presented in Table 4. Table 4. — Amount of Water, Freezing at -5°C. in Cabbage Leaves Hardened in Coldframe for Different Lengths of Time. Treatment Percent dry matter Percent water Percent total water frozen In 100 gran grams water frozen as of tissue grams water unfrozen Not hardened 9.91 90.09 82.1 73.96 16.13 Hardened 2 days .... 13.20 86.80 75.3 65.32 21.48 Hardened 4 days .... 13.90 86.10 62.8 54.47 31.63 Hardened 9 days .... 14.00 86.00 58.7 50.48 35.62 Hardened 14 days . . . 14.79 85.21 54.6 46.52 38.69 Hardened 16 days ... 18.7 81.3 51.0 41.46 39.84 Hardened 20 days . . . 19.35 80.65 47.9 38.63 42.02 It appears that the percentage of the total water frozen at -5°C. decreases as the plant tissue increases in hardiness. At the same time the amount of total moisture in the plants decreases, accom- panied by an increasing percentage of dry matter. The relation of degree of hardening to the percentage of freezable water and to dry matter content is shown graphically in Figure 4. The dates of de- crease in percentage of water frozen and of increase in percentage of dry matter proceed quite rapidly the first four or five days the plants are exposed to hardening in tli e coldframe. After this, these changes proceed slowly for some days longer. On the whole, it seems that there is a close correlation between the degree of hardiness and the percentage of total water retained in the unfrozen condition. The actual amount of ice per gram of fresh leaf tissue also decreases 46 Missouri Agr. Exp. Sta. Research Bulletin 48 with the degree of hardening, while the actual amount of water re- maining unfrozen increases as shown in Figure 5. The Hardening Process in Vegetable Plants. 47 Influence of time of day on percentage of water frozen. — McCool and Millar 80 found that the time of day influenced both the depres- sion of the freezing point and the amount of water frozen at a given temperature. Their experiments with various cereal plants showed that the freezing point depression of the leaves increases during the forenoon, declines slightly in the afternoon and almost reaches tho early morning value by midnight. Over the same period the per- centage of total moisture varied inversely with the depression of the freezing point, but to a much less degree. Shaded oat plants de- creased steadily in sap density during the day, while exposed plants showed the usual increase at mid-day. The slight difference in water content of plants is held by these writers to be insufficient to explain fully the increased sap concentration at mid-day, hence it seems that the products of photosynthesis must play a part. Bar- ley plants kept under bell jars in a saturated atmosphere, under con- ditions retarding transpiration but permitting photosynthesis, had 55> percent of the water in the tops and 62 percent of that in the roots frozen at -3°C. to -4°C. in the morning. At noon 43 percent of the water was frozen in the tops and 59 percent in the roots, at the same temperature. It has been shown by Dixon 133 that illumination increased the osmotic concentration in leaves and this concentration gradually fell when light was cut off. Chandler 20 also found that plants shaded 24 hours had decreased concentration. According to Drabble and Drabble, 134 a greater concentration of cell sap occurs in plants sub- jected to factors favoring rapid loss of water by transpiration. Under these conditions the increased concentration of cell sap is probably very largely the result of, as well as the means of protection against, rapid loss of water from the leaves. In the course of the experiments on the amount of water freezing in cabbage leaves of different degrees of hardiness, some data were obtained relative to the effect of the time of day on the amount of water freezing in leaves of the same hardiness. No attempt was made to provide specially controlled conditions ; they were the same as those previously described for the various hardening treatments, and were identical with those referred to in Table 2. 48 Missouri Agr. Exp. Sta. Research Bulletin 48 Table 5. — Effect of Time of Day on Amount of Water Frozen in Cabbage Leaves at -5°C. Material Time Percent moisture in plants Percent water frozen at — 5' C. Wet-grown greenhouse 9 A. M. 90.43 82.4 plants (tender) 2 P. M. 90.22 78.2 6 P. M. 85.9 Dry-grown greenhouse 9 A. M. 86.60 55.8 plants (hardy) 2 P. M. 86.39 47.1 Coldframe 9 A. M. 87.87 61.9 hardened plants 2 P. M. 84.12 55.5 It is seen that the amount of freezable water is somewhat greater in the morning than in early afternoon, but the differences are not as great as those found by McCool and Millar. The moisture con- tent is also somewhat less in the afternoon indicating the possibility of a greater power of imbibition at that time. Probably the larger factor in causing the slight difference in amount of frozen water is the increased concentration of sugars formed by the photosynthetic activities of the leaf. Both the moisture content and the concentra- tion of cell sap evidently have some influence on the amount of freezable water. Effect of watering plants with salt solutions on amount of easily frozen water in the leaves. — A method used to harden vege- table plants was watering with salt solutions. Only one of these ex- periments will be discussed here. On February 15, seedling cabbage plants were potted in 3-inch clay pots, which were plunged in soil on a raised bench in the greenhouse. One, series was potted in river sand, one in greenhouse compost soil and a third in compost plus rotten stable manure. Each series was divided into four plots and after the plants were well established one of each series was watered with: (1) tap water, (2) M/10 NaN0 3 , (3) M/10 KC1, (4) M/10 NaCl. These applications were repeated every few days, when water appeared to be required. After the second application, the rate of growth in the different plots was evidently being affected. All of the salt solutions depressed growth but particularly in the series grown in compost and in the compost and manure mixture. Plants growing in the sand showed some of this stunting effect, but much later than in compost soils. A test made March 30 showed that the plants grown in the compost soils and stunted by the salt applica- tions were much hardier than those receiving tap water and making normal growth. Little effect of the salts upon either the size or The Hardening Process in Vegetable Plants. 49 hardiness of the plants grown in sand could be observed. Plants grown in the compost soils and watered with NaCl were exposed to -6°C. for 45 minutes without injury. Plants in compost soils watered with KC1 and NaN0 3 were injured somewhat under the same con- ditions, and those receiving tap water were killed. Plants from all of the lots grown in sand were killed at -6°C., but when exposed to -3°C. to -4°C. for one hour, only those receiving water were much injured. The day following the freezing tests, dilatometer determina- tions were made on leaves of plants from some of the lots given different treatments, the results being shown in Table 6. Most of these figures represent only one determination. The samples were gathered about 1 :30 P. M. on a bright sunny day, which may explain why the percentage of water frozen in some of these plants is a little less than that shown in Table 3 for tissues of approximately the same degree of hardiness. Table 6. — Amount of Water Freezing at -5°C. in Cabbage Leaves From Plants Watered with Various Salt Solutions. iPercent dry matter Treatment of plants Percent moisture Percent total water frozen at -5°C. Grams water frozen in 100 grams of leaves In compost soil watered with tap water (medium tender) . . .10.86 89.14 61.2 54.55 In compost soil watered with *1/10 NaN0 3 (hardy) . .11.83 88.13 37.3 32.87 In compost soil watered with M/10 NaCl (hardy) .12.02 87.98 39.5 34.76 In compost and manure-watered with M/10 NaCl (very hardy) 14.03 85.97 27.2 23.29 In sand watered with tap water (very tender) . 8.24 91.76 79.8 73.22 In sand watered with M/10 NaCl (medium tender) 11.01 88.99 59.4 52.86 The percentage of water frozen is much less in the stunted plants — those found most hardy to cold. The amount of water frozen is correlated with the observed degree of cold-resistance and the extent to which growth was checked. Here again the percentage of water frozen varies inversely with the percentage of dry matter. Unfortunately the freezing point depressions of the plants used in this experiment were not taken. However, we know from Chandler’s work that the sap of plants watered with salt solutions has an in- creased osmotic concentration. Bartetzko 4 found that Aspergillus , Penicillium and other fungi grown in nutrient media of varying 50 Missouri Agr. Exp. Sta. Research Bulletin 48 concentrations increased their resistance to freezing in proportion to the increase in the osmotic strength of the medium. The question arises as to how the application of salt solutions to soils in which plants are growing checks growth, increases cold resistance and reduces the amount of freezable water. The stunt- ing might be due to: (a) The toxicity of the salt solution to the roots of the plants at the concentration used. However, since the salt solutions were not appreciably toxic to the plants grown in sand, it seems doubtful if the stunting and hardening of the cabbage plants in the compost soils can be attributed to this factor, (b) Absorption of the salts by the plants, causing a greater concentration of the sap, yet why should nutrient salts such as NaN0 3 cause a stunting of healthy plants? (c) Condition of physiological drought within the plant, at least at such times as the moisture content of the soil was low or the rate -of transpiration very rapid. Such a condition might easily arise, in treating a succulent plant such as cabbage, with rather strong salt solutions. If the tops of the plants suffered from phy- siological drought a considerable part of the time because the roots were unable to absorb water rapidly from the more concentrated soil solution, then a condition would exist more or less similar to that in ordinary soils wherein plants have been hardened by partially withholding moisture. The fact that the lots grown in sand did not show nearly so much of the checking or stunting effect as those grown in the finer soils containing more organic matter lends strength to this idea. To see whether or not the observed results might be due to variations in the concentration of the soil solution this was determined in each lot at the end of the experiment. The method of Bouyoucos 13 w r as employed, taking 15 grams of air-dry soil and 10 cc. of distilled water, determining the freezing point depression with the Beckman thermometer, and calculating that the soil solution contained 100 parts of solute per million for each 0.004° C. of freezing point lowering. The results are presented in Table 7. The Hardening Process in Vegetable Plants. 51 Table 7. — Concentration of Soil Solutions After M/10 Salt Solutions Were Applied Five Times. Treatment Sand Compost Compost and manure Freezing 1 point depression p.p.m. in soil solution Freezing point depression p.p.m. in soil solution Freezing point depression p.p.m. in soil solution Tap water . . .005°C. 125 .045°C. 1125 .159°C. 3975 M/10 NaNO s . . .020°C. 400 .277 6925 M/10 KC1 .. .020°C. 400 .210 5200 M/10 NaCl . . 1 .020°C. 400 .263 6576 : .367 9175 The results set forth in Table 7 show that at the conclusion of the experiment, just after samples for the dilatometer determinations had been taken, the concentration of the soil solutions had increased very markedly in the compost soils to which the salts were applied, as compared to the check of the same sort of soil, but receiving tap water. However, in sand the soil solution was much less concentrated and there was no great increase in the concentration of the soil solutions where the salts were applied. Bouyoucos 13 has shown that 52 Missouri Agr. Exp. Sta. Research Bulletin 48 soils containing much organic matter cause a large amount of prac- tically pure water to become “unfree” by means of capillary ad- sorption. At a given moisture content therefore, the free solution in such soils would be more concentrated than in a sandy soil having little organic matter and large soil particles. Comparing the data in Tables 6 and 7 it is observed that the amount of water frozen in plants at -5°C. varies inversely to the concentration of the soil solu- tion, but probably not in proportion to it. This point is illustrated by Figure 6. To show to what extent the growth of plants in this experiment was affected by the salt applications in the three soil media used, the average green weights, average dry weights, and percentages of dry matter are given in Table 8, for the plants in each lot at the end of the experiment (one day after the freezing determinations were made). Table 8. — Average Growth Made by Cabbage Plants in Soils Receiving Five Applications of M/10 Salt Solutions. Treatment Sand Sandy compost J Sandy compost and horse manure Green Dry % dry Green Dry % dry Green dry 1 % dry wt. wt. matter wt. wt. matter wt. wt. ! matter Tap water . . . 13.55 .967 6.78 8.12 .581 7.16 M/10 NaN0 3 . 8.58 .560 6.52 4.63 .431 9.31 M/10 KC1 . . 8.32 .523 6.29 5.38 .505 9.40 6.18 .631 ; 10.22 M/10 NaCl . . 8.56 .546 | 6.37 4.76 .455 9.57 5.53 1 .624 j 9.55 It is seen from Table 8 that the plants in the sand made nearly twice as much growth as plants receiving corresponding treatment in the compost soils, using the average green weights as the indicator. It should be remembered that in this experiment a rather high soil moisture was maintained in order to prevent the lack of water from influencing the results ; furthermore this experiment was ended before the lack of nutrient material in the sandy soils could become the main factor limiting their growth. The indications point to the conclusion that applications of rather strong salt solutions raised the concentration of the soil solu- tion to a point at which roots could take up water only slowly, and probably not at all when the total soil moisture content fell below a certain point. This developed a state of physiological drought in the tops due to the restricted water intake. Under these conditions, the leaves developed xerophytic characteristics to some extent, as indi- The Hardening Process in Vegetable Plants. 53 cated by the greatly increased water-retaining power on the part of the cells. This is shown by the smaller amounts of water frozen in the leaves of such plants, and as is shown later, by the lower trans- piration rate, and slower rate of drying in an oven. Another ex- ample of increased cold resistance apparently resulting from phy- siological drought was observed in the field in the spring of 1921. On March 31, the temperature fell to -8°C., and the following night to -6°C. Hardened cabbage plants set in the field 10 days previous, were very severely injured, but here and there thru the field small plants were observed after the freeze, the leaves of which were ap- parently uninjured. On examination, the stems of all such plants were found to be nearly severed by a “ damping off” fungus. Evi- dently the stem injury by the fungus had caused physiological drought in the top of the plant, resulting in considerable increase in hardiness. Relation of amount of freezable water to percentage of dry matter and freezing point depression in garden plants. — Three spec- ies of plants were used in these experiments, cauliflower representing a group- possessing potential hardiness, and tomatoes and sweet po- tatoes representing plants lacking potential hardiness. Leaves were gathered during June from plants growing under ordinary condi- tions in the garden. The soil was fairly moist at this time and the plants were making good growth. A portion of each lot of leaves was used for the dilatometer determination, and another portion for determination of the freezing point depression. This latter was made, not on the expressed sap, as were the previous determinations herein reported, but directly on the triturated leaf tissue, according to the method of Bouyoucos and McCool. 14 The results are given for two sets of determinations in Table 9. In the last two columns of this table are given the relative amounts of frozen and unfrozen water, calculated on the basis of 100 grams of fresh leaf tissue. The plants used in these experiments would probably have been killed by a brief exposure to -3°C., except the cauli- flower, which might have withstood a somewhat lower temperature. It may be seen from Table 9 that the percentage of total water freezing in cauliflower at -5°C. is somewhat less than in tomato and sweet potato, while at -3°C. this difference is much greater in favor of the hardier cauliflower. The amount of water remaining unfrozen is correspondingly greater in cauliflower. It appears that allowing cauliflower leaves to stand in 8 percent sucrose over night has in- creased the percentage of dry matter and the freezing point depres- sion and has decreased the amount of water freezing at -5°C. The 54 Missouri Agr. Exp. Sta. Research Bulletin 48 Table 9. — Amount of Water Frozen in Leaves of Garden Plants. In 100 grams leaf tissue frozen at - -5°C. % Freezing grams grams dry point % water water water Date Plant matter depression frozen frozen unfrozen June 8 Cauliflower (in 8 % sucrose 15.86 .780°C. 57.0 48.0 36.14 over night) Cauliflower 13.26 .413 77.4 67.1 19.64 (in water over night) Tomato 11.73 .650 79.3 70.0 18.27 99 99 Sweet Potato 17.5 83.0 69.4 13.1 Frozen at - -3°C. June 21 Cauliflower 17.7 1.300 28.3 23.3 59.0 " " Tomato 13.5 .915 43.7 | 37.9 48.6 " " Sweet Potato 14.84 .750 48.1 41.0 44.16 amount of water remaining unfrozen in the cauliflower leaves which had been in the sugar solution is nearly twice as much as in the check leaves kept in water. Since sucrose penetrates plant tissue quite slowly, the changes noted are probably not due to increased sugar content. However, the dry matter is 2.60 percent or nearly J greater in the leaves placed in sugar solution. The 8 percent sucrose solution is approximately equivalent to 0.25 molecular con- centration. Under these conditions, water may be withdrawn from the leaf, thereby decreasing the moisture content, increasing the per- centage of dry matter and presumably increasing the power of im- bibition with which the remaining moisture is held by the leaf cells. Rather tender cabbage plants, the roots of which were placed in 8 per- cent sucrose solution, wilted quickly, indicating withdrawal of water from the upper portion of the plant, or at least stoppage of intake to make up losses by transpiration. Resume.— It does not necessarily follow from the water-loss theory of killing by cold that there is a definite minimum moisture content below which the protoplasm of all plants dies. In view of experiments such as those of Adams 3 and of Kiesselbach and Rat- cliff 52 it seems quite likely that the minimum amount of water re- quired by plant cells to retain life varies with the state of physiologi- cal activity, the stage of development, perhaps with changes in either internal or external conditions, and probably differs in various spe- cies at the same stage of development and under the same conditions. Ewart 135 has shown that some seeds can he dried to a moisture con- The Hardening Process in Vegetable Plants. 55 tent of 1 or 2 percent without killing and there is reason to believe that if a tender cabbage leaf is killed by the loss of 50 percent of its water, a hardened leaf may be able to survive the loss of even a larger fraction at still lower temperatures. May not the hardening process in vegetable plants, the maturing process in woody stems and the ripening process in seeds involve changes which increase the stability of the protoplasmic structure as well as changes which make for increased water-retaining power? RATE OF WATER-LOSS BY TRANSPIRATION IN HARDENED AND TENDER CABBAGE. It is a commonly observed fact that non-hardened vegetable plants wiiLseverely. upon transplanting to the field and if conditions favor rapid transpiration or if the soil is dry they may die, due to excessive water loss. On the other hand, plants properly hardened by any of the methods mentioned in this paper withstand transplant- ing without serious wilting. To the practical grower the ability of hardened plants to survive transplanting without dangerous wilting is probably of greater importance than the increased cold-resistance developed by the hardening process. Plate 6, B and C, illustrates the marked difference in turgor of hardened and not hardened cabbage plants one day after transplanting to the field. These were potted plants, so the root systems were not disturbed much by transplanting. Of interest in this connection are the observations of Bergen 7 on the rate of transpiration of a number of evergreens, as Olea , Qiter- cus and Pistacia, compared to that of TJlmus and Pisum sativum. He found that the water loss in the former group was 25 percent less than in the latter. He concluded, however, that xerophytic leaf struc- ture (of the hardy evergreens) is not always incompatible with abundant transpiration, but sometimes exists only for use in emer- gencies, to protect the plant from injurious loss of water. Salmon 108 draws attention to the xerophytic structure of the hardiest types of winter cereals; winter rye, Turkey and Kharkoff wheats are characterized, for example, by a narrow leaf and pros- trate habit of growth. The same is true of Winter Turf, the hardiest variety of winter oats. Salmon found no differences in cell structure, epidermal covering, or mechanical ability to control transpiration, that could be correlated with the great difference in hardiness known to exist in cereals, except that Turkey wheat (hardy) had 25 percent greater root length than Fultz (less hardy) and 40 percent greater than oats and barley (least hardy). This character might enable the 56 Missouri Agr. Exp. Sta. Research Bulletin 48 plant to escape dangerous drying out when the ground is frozen to a certain depth. The relation existing between water-retaining power and re- sistance to cold is demonstrated by observations of workers 53 in the United States Forest Service, in a recent study of a chlorosis of coni- fer seedings. The chlorotic leaves were less turgid than normal leaves and wilted very quickly when the water supply was cut off; in fact, chlorotic leaves of the Douglass fir wilted so quickly that accurate leaf measurements could not be made. Plants having chlo- rotic leaves failed to harden properly in the fall, so that many were injured by early fall frosts and many more by winter cold. How- ever, in plots where the chlorosis was corrected in summer by spray- ing with ferrous sulfate, the plants became perfectly winter-hardy. Evidently chlorotic leaves are unable, because of absence of chlo- rophyl, to develop the usual water-retaining power and cold resistance of the species. It was considered desirable to determine the difference in rate of transpiration of non-hardened plants and plants hardened in various ways, because of the indications which might be obtained thereby as to the relative water-retaining power of plants of different degrees of hardiness. Four experiments were performed, using cab- bage plants in 4-inch clay pots. The pots were coated and sealed with a mixture of paraffin, vaseline and beeswax. Two to four plants were used from each experimental lot. Before sealing, the pots were brought to uniform moisture content. The experiments were con- ducted under different conditions, but in each experiment the plants were kept uniform with reference to external factors. Plants as nearly the same size as possible were used, but the hardened plants were usually smaller than the non-hardened. At the conclusion of each experiment the plants were weighed at once and the leaf area of each plant was measured with a planimeter. The results of the four experiments are presented in Table 10. The Hardening Process in Vegetable Plants. 57 Table 10. — Transpiration Experiments With Cabbage Plants. Treatment of plants No. plants j used | Av. leaf area per plant Av. Ajmit. transpired per plant , Transpiration in grains per hour j per sq. M. per plant ! leaf area Expt. 1 (outdoors) 3/15/21 partly cloudy, cool, moderate wind, 24 hours Dry-grown gh. plants 4 125.0 sq. cm. 13.15 g. 0.547 43.7 Wet-grown gh. plants 2 354.0 39.6 1.649 46.6 Expt. 2 (In cool greenhouse) 3/15/21 temp. 60-70 degrees F., 24 hours Dry-grown gh. plants 4 167.0 12.95 I 0.539 i 32.3 Wet-grown gh. plants 2 285.0 27.9 1.162 40.8 Expt. 3 (In warm greenhouse) 3/19/21 temp. 65-80 degrees F., 24 hours Coldframe hardened 1 for 5 days 2 347.0 34.60 1.441 41.5 Dry-grown gh. plants 4 ; 165.0 19.22 0.800 48.5 Wet-grown gh. plants 2 ! 1 315.0 I ! 41.15 1.714 54.4 Expt. 4 (Outdoors) 4/2/21, clear, warm, little wind, , 5 hours, 11:30 A. M. to 4:30 P. M. Coldframe hardened for 1 week 2 278.3 i 18.90 3.778 135.6 Med. dry-grown gh. plants 3 202.0 12.97 2.590 128.3 Med. wet-grown gh. plants 2 395.5 1 29.6 5.920 150.0 Greenhouse plants grown in compost soil and watered with M/10 NaCl (hardy) 2 153.6 j 10.5 2.100 136.6 Same, watered with tap water (tender) ...2 164.0 15.9 3.180 193.9 Greenhouse plants grown in sand, and watered with tap water (tender) . 2 | 252.8 1 23.9 4.780 189.0 The water loss per square centimeter of leaf area per hour is somewhat greater in tender plants than in those hardened by drying, by coldframe exposure, or by watering with salt solutions. The much greater total water loss of the non-hardened plants was due to a large extent to the fact that they were larger than the hardened plants, though of the same age. 58 Missouri Agr. Exp. Sta. Research Bulletin 48 The fact that the rate of transpiration per unit of leaf area was less in hardened plants is significant. If the rate of diffusion of water from the cells into the intercellular spaces determines the rate of transpiration, then a lower rate of transpiration would be as- sociated with a greater water-retaining power on the part of the plant cells. This water-retaining power would be exerted when the plant’s cells are exposed to water loss by freezing in the same way as when exposed to loss by transpiration or by drying. RATE OF DEHYDRATION IN HARDENED AND TENDER PLANTS. Since it was found that hardened plants exhibited a greater water-retaining power than non-hardened plants upon freezing, it was thought that the difference might be measured by the rate of water loss in similar tissues exposed to drying. Mr. V. R. Boswell 8 undertook a special investigation of the rate of dehydration of leaves from hardened and non-hardened plants during the winter and spring of 1921. The material used in his experiments was from the same lots upon which other results are reported in this paper. Leaves of uniform condition and from corresponding parts of plants were gathered from cabbage and tomato plants subjected to various hardening treatments. Lots directly comparable were gath- ered and dried at the same time. The samples were placed in stop- pered bottles, taken at once to the laboratory, weighed, and immediate- Table 11. — Rate of Water Loss by Drying at 60°C. in Hardened and Ten- der Leaveis. (In per cent of total moisture content) Tomato leaves Cabbage leaves Time Greenhouse Hardened Greenhouse Hardened Greenhouse plants in wet-grown in cold- wet-grown in cold- water NaNO 3 | NaCl minutes (tender) frame (tender) frame (ten- (medi- (hardy) der) um har- , dy) | 15 34.77 26.46 21.53 8.92 23.82 19.70 ’ 12.13 30 68.11 57.91 42.71 17.43 46.71 38.68 24.29 45 83.99 75.34 54.20 36.83 62.74 54.80 34.68 60 94.27 87.85 75.32 47.56 74.81 j 66.27 43.13 75 97.94 95.57 79.08 55.23 84.67 | 76.42 51.99 90 99.34 99.02 93.32 68.23 91.59 | 83.20 59.36 105 99.59 99.52 93.12 74.58 96.32 ! 89.27 66.79 120 99.62 99.61 99.66 86.01 98.73 ! 93.94 73.34 135 99.64 99.63 | 98.11 94.88 99.68 j 97.57 i 78.87 150 99.80 94.73 99.93 ! 99.37 84.48 165 1 98.88 99.95 1 99.87 88.91 The Hardening Process in Vegetable Plants. 59 ly placed in an electric oven at a constant temperature of 60°C. The leaves were spread out on wire gauze placed on the shelves of the oven until they were beginning to become brittle, after which they were transferred to the weighing bottles in which the dehydration was completed. Each lot of leaves was removed from the oven at intervals of 15 minutes, cooled and weighed. From the loss in weight for each period of drying, was calculated the percent of total mois- ture removed per period. Table 11 compiled from some of Boswell’s data, gives the percent of total moisture lost at the end of each 15- minutes interval in samples of hardened and tender plants. The data presented in Table 11 bring out a striking difference in the rate of water-loss by drying in hardened and non-hardened leaves of cabbage, especially at the beginning of the period of drying. This difference as not very great in the two lots of leaves of tomato. In cabbage, leaves from plants hardened by exposure in the cold- frame, by watering with salt solutions and by partially withholding water, show a much smaller loss of water for each period than do leaves of tender, well-watered plants grown in the greenhouse. This difference in rate of drying indicates a relatively much greater water-retaining power in hardened plants. Whatever the differences in the two types of plant tissue are, the greater water-retaining power of the hardy tissue evidently does not depend entirely on the or- ganization of living matter, but on the chemical and physical prop- erties of the substances of which the tissues are composed. Another point which may be seen from Boswell’s dehydration experiments is that tomato leaves dry out much more rapidly than cabbage leaves. Even the hardened tomato leaves give up water faster than the leaves of non-hardened cabbage. In view of the fact that the tomato is not susceptible of hardening to the extent of sur- viving ice formation, it seems that we have here an indication of the fundamental difference between the two types of plants. The tomato lacks the potential ability to acquire or develop increased water-retaining power to any great degree, while the cabbage and similar plants have this potentiality to a considerable degree. 60 Missouri Agr. Exp. Sta. Research Bulletin 48 Figure 7 shows graphically the relative rate of water-loss by diying in the different types of tissue. d >-50 O EL >.'! El e* H 7 70 o u i? U H \~ z iii — - — ^>1 """ I 1 / ' / / s' ./ X 1 / / / / ' 1 i / f / / / / / / X / / K X X // / X* X* / / / / , X X ' Te n t- Torr-.c-to Na.rie.-nea Ibrrtxto - - - — Ttidst- Cs-bVe-a®. . i / ~/~7 — X ' / + + + 4 Hat- Ae AC&-bbag« \ \ t_> <<• es fc- i f A /// X X X' t/y A," 1 O IS 3o 15 bo ns 90 /os 120 rss !Si /(•? DURATION OF DRYING IN THE OVEN AT GO 6 C. Fig. 7. — Rate of water loss from leaves of varying degrees of hardiness. CHANGES IN CARBOHYDRATES ON HARDENING OF PLANTS. Formation of sugar by low temperature.— Numerous investi- gators have noted increased amounts of sugars in plants exposed to low temperature. Mer 67 was probably the first to note the disap- pearance of starch and the accumulation of sugar in evergreen leaves in winter. Lidforss 60 noted in a number of evergreen plants in Sweden that starch was converted to sugar in the fall and reconverted into starch in spring. He found that tender seedlings placed in a sugar solu- tion for a short time w^ere able to withstand several degrees of lower temperature without injury. Lidforss thought the hardiness of the evergreen leaves and the sugar-treated seedlings w T as due to increased concentration of the cell sap resulting from the accumulation of sugar, to which he attributed reduced transpiration and low r er freez- ing point depression, as well as a protective effect of sugar on the precipitation of proteins of the cell. Gorke 35 found that he could prevent the precipitation of protein from expressed plant sap by adding sugar. The Hardening Process in Vegetable Plants. 61 Miyake 70 examined the leaves of evergreen plants in various parts of Japan in winter finding those of many plants to be starch- free during the coldest part of winter. Another group had very little starch in the mesophyll during cold weather (when the mean temperature was near or below the freezing point.) Plants in Northern Japan were markedly lower in starch than in the warmer sections. Schulz 111 examined one hundred species of plants in Ger- many, finding most of them starch-free in winter, while a few con- tained a little starch mostly in the fibrovaseular bundles and the surrounding cells. Recently, Swedish investigators 2 have shown that the hardier varieties of wheat have a larger sugar content in fall and winter. They found that the percentage of dry matter and the amount of sugar in winter wheat varies considerably during the winter, fluc- tuating with the temperature, but during the period from November 12 to February 15, no starch could be found in the leaves. Gasner and Grimme 31 upon analyzing the first leaves of wheat, found that seedlings germinated at 5-6 °C. had a greater sugar content than those germinated at 28 °C. They also found a higher sugar content in leaves of hardy winter wheats than in spring wheats germinated at the same temperature. Micheal-Durand 69 in extensive studies on the changes of carbo- hydrates in plants, found an enormous accumulation of sugars in leaves of certain evergreens in winter, while starch completely dis- appeared during the coldest weather. ITe explains this condition as follows : (1) In winter assimilation is low, but respiration is depressed still more by the low temperature. (2) Conditions in winter are unfavorable for translocation. (3) Low temperature prevents the condensation of the simple sugars into higher carbohydrates. (4) Breaking up (splitting) of polysaccharides. Miiller-Thurgau 75 and others, have measured the accumulation of simple sugars in potatoes at the expense of starch upon exposure to low temperature and the reconversion of the sugars into starch when higher temperatures are provided. This reversible chemical change seems to be generally associated with changing temperatures near the freezing point, probably due to shifting of chemical equi- librium by enzymatic activity. Relation of sugar content to cold resistance. — Since the ex- periments of Lidforss 00 and Gorke 35 the extensive formation of 62 Missouri Agr. Exp. Sta. Research Bulletin 48 >> s-4 00 rH O O O LO CO oq oo 00 tH rH rH rH oq LA LA t— CO oq © oq Q X to Oq H t- CO OOOH< 00 t— LO co oq CO oq 05 p p o oq co p p cq 00 00 © r-i r-i O oq rH rH d d 05 rH rH Oq co oq’ oq’ oq rH cd oq LA CD LA rH rH rH rH rH rH rH rH tH H H H rH tH tH HtHHH X *x X a> © a3 tH 00 rH rH CO 00 05 rH 00 00 CO t- • A OO th rH tH tH oq P 5 >> LA OS oq co oq 05 rH c— CO 05 CD 05 oq t- t- LA rH o- Eh o O xi 05 CO rH CO 05 t— O 00 t- c~ 05 OO O oq © oq c-; p © tH £ p Eh ft O © d r-i d d d rH o d d d d r-i r-i ri © © © © © W £ Eh be O G 0) X If 43 00 Oq 05 Oq rH 05 05 05 ^ CO rH © © LA © © 2h P CO o rH rH OO CO oq co oq 05 ©LA © TH L~- CO H X a> 00 oo oo co oq oq CO oq rH O Oq rH © tH TH d d d d d o d d d d © © © © © d o £ CQ fl _ X oq la rH Tf* 00 CO o oo 00 -eJL © LA rH © t- oq 00 l-H CO LO 05 00 CO c- LO co rH 00 00 © rH © t- rH M oq g 4-> oq t— i tH O c- CO CO oq tn; Lfl coo p p TH rH cq p cq < G a Eh S3 d d d d d d d th* d rH rH rH © © rH rH © © © X sh ojO o a> 1, 33 » oo t~ co oq o CD O tc- th © rH tH © oo oq oo oq 05 L£5 H- 00 00 Tt< CD rH rH LA © © CO t- P 33 be O r-4 CO CO oq rH 05 O © © th p p © rH © g © 33 d d d o d d O r-i r-i © © © © © © © a> x Si P cc X ^ *3 in m rH a TO <4 — J o Q- Ph bo 43 > Ch 73 rt< 00 oq co oq* oq" CO oq co d rH ^3 od od -a G "3 X 00 CO o t- oq oq 00 o t> o LA 05 LA © © -^ © t- O -+-> G be 33 X p CO o © CO th LO p © 1> LA t>. © © LA © rH rH o o ■ij g Q) C4 O H oq r-i d CO oo d CD rH H O OO tH tH tH Tji i-i 05 rH LA oo" ©’ 5 b/3 G X S-c S-c O © lo LA © 2 o *o G O LO o o LO 00 t— CD tr rH LA ers © LA Ph bfl rH Oq LO LA oq t-; tr oo oq oq co © TH rH © eq o n © a> tf G X rH tH cc LO oq* rH oo oo oo tH CO LA* CD r-i Oq *3 © 0)00 05 O 05 O O O o O O 05 05 o 05 O O © © © © © © © O a> a) rH oq oq rH Oq t-h oq oq oq oq oq oq rH rH Oq rH oq oq rH rH Oq cq rH \W w \w \ \\\ w w \ \ \ £; G a n fq o oq oq oO oq lo o oq oq LO © oq oq © oq o oq oo oq la rH rH rH g Oq © T-l Oq rH \rH \ rH rH oq \ cq oq oq oq Oq rH oq \ oq \ CO CO CO Q 33 03 W oq \th \ \ \ W W \oq \th N \ cc h oo oq rH 00 rH CO CO CO CO rH CO co co oq rH rH CO © © © 5 s? X rH l-H rH rH 1 rH rH rH rH o o o> a> X U1 ri u Jr C4— j- Lettuce X HH> 33 G — X ^3 o3 G P X X +-> c -ri rH 2 2 E- a> 3 as a> Cabbage wn wet, in ouse (tendei wn medium i greenhous wn dry in ouse (hardy ter partially eld for 2 w in greenhou enhouse plai ot hardened dened in co week O o 33 T3 o> J3 o> 73 X 0) rown wet ( ft o> X G O P G 0) j- G O h lenhouse pla rown in riel very tender) Eh | 2^ c .p O r; tr o3 43 (D jr U * (r 03 oq o3 00 d> be A t- be 2 ^ w 1 o o C5 £ o X s X 6 CD O £ X 6 1 ,-H oq co CO oq rH rH co ZD 0) 0) 03 •'- £ <* << <5 a a a a < < r* .AUI.E ±z. — itonunuea) The Hardening Process in Vegetable Plants. 63 3r cent on fresh weight basis % Dry i matter 1 5.13 LO oo in h 02 06 rH co c-P to (MOOO o in co c- o’ co’ rH 11.13 9.97 11.9 13.4 12.41 8.29 17.78 11.00 14.1 14.6 13.85 Total poly- saccharides t- to CO © 0.264 j 0.605 ! 0.527 0.860 cm cm co o’ 1.240 t- rH ▼H 0.419 0.691 1.346 1.711 0.662 i o rH © 3.416 A C2 f-t d -1-2 w 02 02 cq © 0.280 OO Tt< O LO rH CO © o’ 0.226 i 1 0.488 1.56 | 0.502 , LO LO t>- rH Total sugars o cm o’ 0.121 0.746 0.485 0.499 2.100 0.874 2.487 1.594 0.982 H8 ’I 0.285 0.320 0.313 00 00 rf d 0.335 to © d PU ox; d ai rj to 00 rH CM MOO co cm CO rH LO LO TjH oo 02 cm cm o O rf rt< LO 02 02 00 CO CO cq 00 oo © OlflH CM 00 CO LO CO LO 02 o rH CO rH CM 02 M o’ o’ o’ o' o o o’ © © o' © o’ d d © ©’ PS r< r-i J2 02 io oo 02 d a <12 rf co cm O 02 02 cm 02 cm Tt< rH 02 LO OO 03 rH co t> LO rH LO CO co co LO O LO rf CM d O M A tP to fc*“ C"P 00 LO’ o’ 00 co oo CO Hjl CO d E-* a? *h O d rH rH CM CM -<-> A Cl be 02 > o c3 OO LO CM CO CO 00 rH LO >> u m Id CO cm co rH lO CO d | d d is bfl q m — H t- o d d Wl 'd d a) cc PS 02 02 d P. Q 6 5 <22 o be d O “ o O o " a . 00 © © CM © LO C2 CO©l> 02 co O O CO t- CO CO 02 d ^ d t- o . ~ ^ hfl 02--O o CO H 0 W Hob; tc- o' M cm rH o OO © LO 00 © CO cm’ oo d 02 © o i-H cm cm \\\ 02 O Oq^jH HCON CO “ M d i2 03 5 d ft® 5 d 5 d tP 2 d a p, q d ^ P •d ^ d ft M ^ ~ . o o 1 X 02 02 CO* 02 a d 02 £ §2 g§ ►2 e .. §2 *d ad d T3 d ■g oS rt >>o u % 02 Th ^ d 'd 0r d ° > M’S d d d O 02 S-. -M be d d Q rH ICO ca 02 02 >- 02 d d S 'd 02 ■go *2 d tH d ffi ffi 64 Missouri Agr. Exp. Sta. Research Bulletin 48 sugars in leaves of plants- exposed to cold has generally been con- sidered to be related to their cold-resistance. However, Harvey 42 concluded that carbohydrate changes were not important in the hardening process with cabbage plants, since he found that cabbage plants could be hardened to some extent at least, by keeping them several days in the dark in a low temperature chamber, during which time there was little change in the carbohydrate equilibrium. How- ever, it has been shown by several investigators and notably by Lewis and Tuttle 59 that simple sugars form a large part of the osmotieally active cell contents. From the beginning of these experiments, samples were col- lected for carbohydrate analyses from some of the series of plants in each of the hardening treatments. The results of some of these determinations are given in Table 12. Methods of analysis. — The sugar analyses were made according to the modified Munsen and Walker method, as described by Hook- er, 46 the results being expressed as dextrose. One gram of the air dry, ground plant material was weighed, transferred to filter paper and washed thoroughly five times with distilled water. The insoluble residue was used for the starch determination. The filtrate, amounting to about 150 cc., was taken for determination of soluble sugars. After clearing with basic lead acetate the extract was made up to 250 cc. and filtered. Two hundred cc. of the filtrate was pipetted into a volumetric flask, excess lead precipitated with solid sodium carbonate, made up to 250 cc. and filtered. An aliquot of the filtrate (Solution A) was used for the determination of reducing sugars, while another portion was used for determination of the total sugars. Five cc. of concentrated HC1 was added to 75 cc. of Solution A and hydrolized at 70°C. for exactly ten minutes (Solution B). After cooling, this solution was neutralized with sodium hydroxide made up to 100 cc. and used for the determination of total sugars as dextrose. The sugar-free residue of the original sample was used for the starch determination. It was washed into a beaker, boiled five minutes to convert the starch into a paste and after cooling 3 cc. of Taka-diastase solution were added. The beaker was then placed in the oven at 40°C. for 24 hours, the starch being broken down to maltose and dextrin. The liquid containing these sugars was then filtered off, adding the washings to the filtrate, which w T as hydrolized with acid for 2 y 2 hours under a reflux condenser to break down further the products of digestion to dextrose. After cooling, the solu- tion was neutralized with sodium hydroxide, cleared and prepared for analysis as previously described. A blank with the same amount of Taka- diastase solution was run with each series of starch determination. Total polysaccharides were determined on a sample of the dry plant material washed free of soluble sugars with cold water. The filter paper was punctured and the residue washed into a 700 cc. flask. Eight cc. concentrated HC1. and enough water was added to bring the total volume to 150 cc. After boiling two and one half hours under reflux condenser, the contents of flask were cooled, transferred to a beaker and made neutral to litmus with sodium hydroxide. The solution was then prepared for analysis as previously described for Solution A. The Hardening Process in Vegetable Plants. 65 Discussion. — Table 12 presents evidence that: (1) The content of both reducing and total sugars increases in hardened plants. This increase seems to be greater in plants hardened by exposure to low temperature in the coldframe than in plants hardened by other methods. The increase in sugar is greater in hardened cabbage and lettuce than in the tomato, though there is no direct evidence that the absolute quantity of sugars present in the plant is directly re- lated to its cold-resistance. Thus some of the tender lettuce samples have more sugar than certain samples of hardy cabbage. Young lettuce plants contained much more sugar than plants approaching maturity and this may have something to do with the greater cold- resistance Chandler 20 found in the younger leaves of lettuce, where- as in most plants the young leaves were somewhat more tender to cold. (2) In lettuce, cauliflower and cabbage the amount of total polysaccharides is usually somewhat less in hardened than in non- hardened plants, which decrease may be attributed to the reduction in the amount of starch. In the tomato, on the other hand, the total polysaccharides show a large increase, apparently due mostly to the deposition of starch in large quantities in both stems and leaves of plants exposed to any of the hardening treatments. Kraus and Kraybill 54 found a similar increase of starch in tomato plants in a stunted condition. Hartwell 43 found a large accumulation of starch in plants, especially the potato, when the growth was checked by any limiting factor. Here is an interesting distinction between the chemical changes in a group of plants susceptible of considerable hardening to cold and a plant not susceptible of much hardening. In the group of plants possessing potential hardiness, exemplified by the cabbage, any hardening treatment causes a considerable increase in sugars and a decrease in starch, while the total polysaccharide figure re- mains nearly constant (on the fresh weight basis) because, as will be shown later, of an increase in pentosans. On the other hand in the tomato, lacking potential hardiness, the hardening treatments caused only a slight increase in the sugars and an enormous increase in polysaccharides due mostly to an increased starch content. An increased sugar content in the hardened plants would in- crease the osmotic concentration of the cell sap, depress the freez- ing point and perhaps serve to hold a somewhat larger amount of water in the unfrozen state when the plant is exposed to low tem- peratures. However, the importance of the increased content of sim- 66 Missouri Agr. Exp. Sta. Research Bulletin 48 pie sugars in cold resistance remains undetermined, nor is it known to what extent sugars may be responsible for the greater water-retain- ing capacity of hardened tissues. It appears probable that an in- creased sugar content in hardened plants is more likely one of the manifestations of the condition of being hardy than a direct cause of cold resistance. NATURE OF WATER-RETAINING POWER IN PLANTS. It has been shown by experiments with the dilatometer that: (1) the amount of water frozen in hardened cabbage plants is con- siderably less than in tender plants, (2) the increase in the amount of water frozen as the temperature is lowered becomes less and less, probably approaching zero, (3) the amount of water freezing at a given temperature (-5°C.) decreases as the degree of hardening in- creases. It has also been shown that hardened plants have a lower transpiration rate and that hardened tissues dry out more slowly than tender tissues. The greater water-retaining power of the cells of hardened plants must therefore be accepted as a fact. What factors are responsible for the development of this increased water- retaining power? Several investigators have attached great significance to the osmotic concentration of the sap as determined by the depression of its freezing point. Some data are also presented in this paper (Table 2) showing that in hardened plants this depression is greater than in non-hardened plants. However, concentration of the sap, even if entirely due to substances having a low eutectic point, would not be sufficient to account for the amount of water found to remain unfrozen in hardened plants (Table 3). Moreover, some of the sap solutes have a high eutectic point, for Harvey 42 found numerous large crystals of calcium malophosphate in frozen spots on leaves exposed to temperatures not low enough to kill the whole plant. Some investigators have stated that the increased sugar content of plants in winter was at least to some extent responsible for their cold-re- sistance because of the increased concentration thereby imparted to the cell sap. The highest percentage of total sugar found in cab- bage in these experiments (Table 12, 1.461 percent in Series El, gathered March 22, 1920,) is equal to only 1.68 percent sugar solu- tion in the plant sap. Considering half of this sugar to be glucose and half sucrose, we have a sugar solution equivalent to less than 0.075 molecular. This would not be sufficient to affect materially the amount of water frozen in the plant tissue at a point several degrees The Hardening Process in Vegetable Plants. 67 below 0°C., one gram-molecular weight of a non-electrolyte lowering the freezing point 1.86° C. It has been further pointed out (p. 35) that the greater depres- sion of the freezing point in hardy tissues is associated with certain changes in the plant cell upon hardening, the apparent increase in sap concentration being simply an accompaniment, rather than a cause of increased hardiness. It therefore is necessary to introduce some other factor to explain the difference in amount of water freez- ing in hardy and in tender tissues. It has been indicated that the force of imbibition may be a powerful factor in withholding water from freezing. This force varies inversely to the water content, but probably increases more rapidly than the rate of decrease in water content, as indicated by the slow rate of drying leaves and of col- loidal materials after they have been dried out to a certain extent. It is a pretty well recognized fact that tissues with lower water con- tent are more resistant to killing by cold. Plant protoplasm is not a compound of definite chemical com- position or even constant physical condition, but a colloidal mixture of the emulsoid type varying in consistency from a hydrosol to that of a hydrogel and containing different substances which may be pres- ent in greater or less amounts at different times and in different organs. According to Seif riz, 113 the change from one state to an- other is dependent upon, or coincident with, changes in physiologi- cal activity. Thus, in the eggs of Fucus, he found a progressive in- crease in viscosity with decreasing physiological activity. Straus- baugh, 117 as a result of recent investigations on the plum in Minne- sota, suggests that the prolonged dormancy and water-retaining power which he found in hardy varieties is due to a change in colloidal prop- erties creating an increased power of imbibition. The work of these in- vestigators is significant, since hardy plants are usually at a low state of physiological activity at the time of their greatest cold re- sistance. Water of imbibition may be held by molecular capillarity or in the absorbed condition by the hydrophilous colloids of the plant cells. Such water is not readily available for freezing, in other words, the force with which it is held must be overcome by a considerable force of crystallization before it can be drawn from the cell and frozen. Me Cool and Millar 80 have suggested the classification of plant mois- ture as “free’ or easily freezable and “unfree” or not easily freez- able, somewhat as Bouyouces has classified soil water. Such a classi- fication necessitates setting an arbitrary temperature of freezing, the 68 Missouri Agr. Exp. Sta. Research Bulletin 48 relative amounts of free and unfree water varying with the tempera- ture at which freezing takes* place. Yet it is convenient for our pres- ent purpose to refer to free and unfree water in the sense that the latter, for one reason or another, remains unfrozen at a given tem- perature. It seems from the work of Bouyoucos 10 and of McCool and Mil- lar 80 that the unfree water is held to a very large extent in the ad- sorbed condition by protoplasmic colloids. The water-retaining power of colloids and the quantity of certain colloidal materials in the cell are thereby suggested as an explanation of increased water-retaining power and cold-resistance in plants. * Relation of pentosan content to cellular water-retaining power. — Spoehr ’s work 116 on cacti suggests that pentosans may be the spe- cific substances which increase the water-retaining power in hardened plants. He found that the pentosan content of Opuntia increased considerably under xerophytic conditions and suggested that the large water-retaining power of the pentosans is largely responsible for their well-known ability to survive under such circumstances. The work of Livingston 138 and others has shown that the osmotic pressure in cacti and other desert plants is no greater than in many mesophytes, hence this factor probably plays only a small part in the water-holding power of most xerophytic plants. Spoehr found by analysis of desert plants that in cells under- going water depletion, other polysaccharides were changed to pento- sans, of which the plant mucilages are largely composed. Thus, 4 ‘un- due loss of water caused a change in the cell whereby the amount of water it may hold is greatly increased. ” Mac Dougal 86 considers a change of this sort to be the basis of xerophytism. Water of imbibi- tion was found by Spoehr to be closely related to the presence of the pentose polysaccharides. Pentosan formation increased decided^ with low and decreased with high water content. From April till June, while the weather was very dry, pentosans made up from 9 to 12 percent of the dry weight. During the rainy weather of July, the pentosan content fell to 4.39 percent of the dry weight, increas- ing to 12.5 percent again in the dry cool weather of the fall and falling to 4.37 percent when the winter rains set in. Not all cacti possess a large pentosan content. Spoehr gives analysis of two species of Opuntia growing at Tucson, as follows: Fresh weight basis % water % total % total % total % sugars polysach. pentose pentosan 0. Versicolor 82.15 1.97 1.50 0.36 0.230 O. Phaeacantha 78.70 3.53 3.22 1.64 1.550 The Hardening Process in Vegetable Plants. 69 In view of this difference in composition, it may be significant that 0. Pha^acantlia is listed in Bailey’s Encj^clopedia of Horticul- ture as a hardy variety and is reported by Shreve to grow in the mountains about Tucson to an altitude of 7500 feet. A striking property of the pentosans is their power of swelling and taking up an enormous amount of water, which the hexose poly- saccharides do not do to nearly so marked a degree. The occurrence of pectins in the middle lamella of the cell walls is well known. Spoehr believes that they are also distributed through the protoplasm and are used for a variety of purposes. The plant nucleo-protein has been found by Levine and Jacobs 136 to contain the pentose group as part of the nucleic acid radical. Tollens 137 showed that pentosans were widely distributed in plants and were limited to no special tissue, but abundant in roots, stems, leaves and seeds. He found further that pentosans showed all possible variations as to solubility in water. Swartz 119 obtained a water-soluble pentosan from Dulce. In the crude form, this was very hygroscopic, but this property was lost after several purifications. She found that the hemi-celluloses of ten species of marine algae were chiefly pentosans and galactans and concluded that pentosans and hexosans very commonly occur to- gether, not only intimately associated, but chemically combined. Mac Dougal 81 goes so far as to state that the “ plant protoplasm con- sists of a comparatively inert base of pentosans — in colloidal com- bination with proteins, amino acids, lipins, and salts.” As to the origin of pentosans, Spoehr 116 shows that pentoses can be formed from the hexosans as the first product of oxidation. This view is corroborated by the work of Ravena and Cereser, 102 who found no marked variation in pentosan content during the period of photosynthetic activity, but when the carbohydrate food consisted entirely of dextrose, the amount of pentosans increased greatly, es- pecially in light. The probability of pentosan formation from the hexosans is indicated also by the increased pentosan content in the presence of high total sugar and diminishing starch, as shown later in this paper. Davis, Daish and Sawyer 24 found no diurnal variation in the pentosan content of plants. However, they found that the amount of pentosan in the leaf of the Mangold ( Beta vulgaris) increased from August to October. Hornby 48 found that the pectin content varied in different parts of the same plant. More pectin was found in the epidermal tissue than in the cortex. Exposure to light, and mechanical injury to 70 Missouri Agr. Exp. Sta. Research Bulletin 48 tissues, were. found to result in increased pectin content in the ex- posed or injured part. Hornby suggested that pectin might have a protective effect on plants, especially against insect attacks. Hooker 47 has shown that the hardier parts of apple shoots, the bases, have a greater water-retaining power than the tips, which are less cold-resistant. He placed portions of the air-dried ground ma- terial in desiccators containing sulfuric acid, the concentration of which ranged from 100 to 36.69 percent. The air-dry material lost moisture in the desiccators containing the higher concentration of acid and this loss was greater in the tender material. But over the lowest concentration of acid used, water was taken up, the gain in weight being greater in the hardy material. This experiment indi- cates that hardy apple twigs contain a larger amount of some hygro- scopic material. Hooker attributed the greater water-retaining power of the hardy tissue to the larger percentage of total pentosan found therein. Pentosan content in the hardening process in vegetable plants. — In this work, a study was made of the pentosan content in an effort to throw light on the nature of the increased water-retaining power of hardened plants. For the pentosan determinations, sam- ples were taken from plants grown under the various hardening treatments previously described. Also a series of analysis were made on plant material gathered from the field at intervals during the fall of 1920. Method of Pentosan Analysis . — The method of analysis was that employed by Spoehr. 116 A two gram sample of the oven-drv material was liydrolized by boiling for three hours with eight cc. concentrated HOI in 150 cc. water. After cooling, the entire con- tents of the flask containing the products of hydrolysis were trans- ferred to a 400 cc. baker, neutralized with NaOH, a uniform amount of a suspension of yeast was added and the beakers placed in an oven at 35-40 °C. over night. The hexose sugars were fermented off, Reaving the non-fermentable pentose sugars in the solution. After fermentation the material was filtered and washed, the filtrate con- taining the pentose sugars was boiled ten minutes to drive off the alcohol, then prepared for analysis in the same way as described for sugar determinations. The result obtained was calculated from Munson and Walker’s tables, multiplying the glucose value by 0.85, since Spoehr found that the reducing value of the pentose sugars held that relation to glucose. The results on total pentosan content are given in Table 13. The Hardening Process in Vegetable Plants. 71 on dry wt. basis 2.12% rH co 6.489 3.29 1.61 2.19 5.10 ce) 4.20 4.20 on fresh i wt. basis 1 § o 0.402 0.564 0.131 0.126 0.230 0.568 head lettu 0.295 0.369 , , o O 05 O © © w 05 05 1 W 03 rH 03 03 03 rH rH © 2 \ x w w w +H C, lO lO LO 05 rH rH rH 05 rH © © \ \ \ 03 03 03 03 03 CO Q 9 o w w w © rH rH rH CO CO co rH O Vi rH rH tH >.•2 GO V o CO LO 05 tc- O rH CO OO t> oo H rH rH LO 03 rH 00 CO 05 rH HOC CO CO rj rH LOCOH ® H LO rH rH CO cq © 05 O rH t> co - 05 rH OO 05 CO oo LO OO t- 50 OO ZD SM .© S co co C~; rH rH f- t>inia to i> iq rH CO LO co iq cq io iq © © © © © © © © © © © © © © © © O cd © © © © o £ 0 rH (J5 O H 05 05 O H 05 C5 05 O 05 © rH © rH O rH rH mNh 03 03 rH rH 03 03 rH rH T- 03 rH 03 03 03 03 03 03 03 \\\ \w\ \\\W w \ . \ w w S »— i •V) <0 Cl CO O rH CO CO O 03 03 CO CO rH CO O co o co o o \ CO rH \COC3H \COHHH \ 03 CO \co \co CO • 9 rt o W lO \\\ WWW LO \ LO \ LO \ IO \ \ rH * 03 03 03 co rH IO 05 O O O rj rH o O rH t-I 03 03 03 03 03 03 03 03 © © \\\\\\ \\\ d ft 00 03 03 03 © 05 03 O 05 H H 03 03 r - ’ rH rl H rl Q c \\\\\\ \ w © Vi 03 CO CO CO CO CO tH CO 03 CO rH HMOioa^cqo^! ^ oo M r| rH 05 CO O ® rH id co id oo rH id id ^ H CO CO O CO CO H 13 Tj< oo IM ^ IM CO CO N H LOuOrHLOCDLOCOkiO 1 * 3 *. — ©5 LO 00 CO 05 © 03 cd id 03 t - rH 03 rH L© 05 o o O ® O O rH i H 03 03 03 03 03 03 03 03 ooooqoqoqifloco® Cq T HHOqo3\HHr l W\WrHW\ him cococo nco m I O |6 © t£.© © 03 05 i— I rH W co co I>- LO O O 05 05 03 tH 03 O o rH 03 03 03 © rtf M d « 72 Missouri Agr. Exp. Sta. Research Bulletin 48 Plants not hardened by any special treatment are low in total pentosans and hardened plants have a much larger amount, in some cases in cabbage an increase of about 200 percent. Plants given intermediate hardening treatments have a medium amount of pen- tosans. The increased pentosan content of the hardened plants is most striking if we consider the results on the fresh weight basis. This probably is the most suitable criterion to use in a study of the reactions which concern the living plant, especially since Parker has shown that the force with which water is held by finely divided materials depends largely on the moisture content. It may seem that the absolute amounts of pentosans, even in the hardened plants, are too small to influence very markedly the force with which the cells may retain water under conditions of stress. However, it should be borne in mind that in nature the pen- tose molecule probably exists in combination with four molecules of galactose or other hexose sugar. Hence the amount of pentosans in the plant is much greater than the analyses indicate. Pentosan content of garden plants. — Samples of leaves were gathered at intervals during the fall from cabbage, kale and celery plants growing in the open field. The seed had been sowed in July and the plants made considerable growth before the first light frost came on October 1. The month of October was mild, and the plants remained alive until heavy freezes the last of November. Exposed to steadily declining seasonal temperatures, these plants may be con- sidered to have undergone a kind of hardening treatment, for they were able to withstand light frost in October and heavy frost the early part of November. The results of the total pentosan deter- minations are given in Table 14. Table 14. — Total Pentosan Content of Garden Plants in Autumn. Kale Cabbage Celery Date sample collected % of fresh wt. | I % of i dry wt. I .% fresh wt. % dry wt. %of fresh wt. % of dry wt. Sept. 15 0.289 4.06 Oct. 7 0.511 3.93 ' 0.580 4.36 ! 0.567 4.42 Oct. 20 0.528 | 4.89 0.545 4.73 0.801 ' 4.26 Nov. 3 0.537 i 3.93 0.621 4.36 i 0.793 4.44 Nov. 10 Nov. 18 0.722 ! 1.064 j 4.95 6.48 0.782 ! 5.31 ! 1.029 | 5.58 Table 14 shows that the total pentosan content of these plants becomes high when thej^ are exposed to cool weather during the late 'The Hardening Process in Vegetable Plants. 73 fall. The pentosan content on the fresh weight basis increases fairly regularly up to date of last sampling. Pentosan content in plants watered with salt solutions. — An experiment wherein the hardiness of cabbage plants was consider- ably increased by watering them with M/10 salt solutions has been described. Plants hardened in this way were shown to have greater water-retaining power than unhardened plants. Samples from the salt treatment plots were analyzed for total pentosan content. The results are given in Table 15. Table 15. — Pentosan Content in Cabbage Plants Hardened by .Salt Solu- tions. Treatment of plants Percent total pentosans On fresh weight basis on dry wt. basis Compost soil, tap water 0.290 4.25 Compost soil, NaNO 0.471 5.05 Compost soil, KC1 0.451 4.24 Compost soil, NaCl 0.483 5.05 Sand, tap water 0.220 3.45 Sand, NaCl 0.288 4.25 The total pentosan content of the plants whose growth was checked by the application of the salt solutions and which were hardier to cold, show somewhat greater amounts of total pentosans on the dry weight basis and a considerable increase on the fresh weight basis, as compared to plants making a normal growth with tap water. It appears from Table 15, that the pentosan content of the plants grown in sand is considerably lower than for plants grown in com- post soil and receiving corresponding treatments. The plants grown in sand and receiving tap water were somewhat tenderer to cold than those grown in compost and likewise given tap water. Plants grown in sand and watered with M/10 NaCl show only a slight increase in pentosan content, as compared to plants grown in compost soil, like- wise watered with M/10 NaCl. Here again, pentosan content shows a close correlation with the hardiness of the plants, as determined by freezing experiments. Rate of increase in pentosan content. — The three groups of ex- periments just described having indicated a larger amount of pen- tosans in plants hardened in different ways, it was deemed desirable to determine their rate of development during the hardening process. Lots of potted cabbage plants were removed from the warm green- 74 Missouri Agr. Exp. Sta. Research Bulletin 48 house at intervals during March, and placed in an open coldframe. On March 19, samples were taken for analysis from all the lots which had been exposed to the hardening process for periods ranging from 3 to 20 days, as well as from some of the original lot which had been kept in the greenhouse under favorable growing conditions. The total pentosan content of the plants hardened for varying lengths of time is given in Table 16. Table 16. — Rate of Increase of the Total Pentosan Content in Cabbage Plants. Percent pentosan Treatment On fresh weight On dry weight basis basis Greenhouse plants, not hardened 1 0.260 2.97 Hardened in frame 3 days 0.374 3.56 Hardened in frame 5 days 0.442 3.86 Hardened in frame 10 days 0.750 5.00 Hardened in frame 20 days 0.776 5.84 Pays duration of Exposure in Coldfranc. Fig. 8. — Rate of increase in total pentosan content of cabbage leaves during the harden- ing process. The results of Table 16 are shown graphically in Figure 8. It appears that the increase in pentosan content proceeds quite rapidly and at a fairly uniform rate for ten days. After the first ten days of exposure in the coldframe the pentosan content increased only The Hardening Process in Vegetable Plants. 75 slightly in this experiment. Other experiments have shown that the cabbage plant acquires nearly its maximum degree of hardening in this time. The dry matter content likewise increases rapidly the first few days of the hardening process, and more slowly thereafter. In the dilatometer experiments, it was found that the amount of water frozen at -5°C. decreased with the duration of the hardening treatment in approximately the same order as the pentosan content is shown to have increased here. This seems to indicate a close relation- ship of pentosan content to water-retaining power and to cold re- sistance. The plants used in the dilatometer experiments were of the same lots as those from which the pentosan analyses were made. Table 17. — Relation of Hot-Water- Soluble Pectins to Total Pentosan Content in the Hardening Process. Percent pentosan on fresh weight basis Treatment Date sample taken Total Hot-water soluble Insoluble (by differ- ence) Cabbage Wet-grown green- house plants 3/12/20 0.215 0.075 0.140 Dry-grown green- house plants 3/12/20 0.423 0.292 0.131 Greenhouse plants not hardened 3/22/20 0.207 0.091 0.116 Hardened in cold- frame 2 weeks 3/22/20 0.530 0.408 0.124 Hardened in cold- frame 3 weeks 3/16/21 0.776 0.550 0.226 Tomato Wet-grown in greenhouse 5/3/20 0.693 0.070 0.623 Dry-grown in greenhouse 5/3/20 0.720 0.071 0.649 Greenhouse plants not hardened 5/3/20 0.384 0.051 0.333 Hardened in cold- frame 2 weeks 5/3/20 0.682 0.071 0.611 Sweet Potato Garden plant 10/7/20 0.477 0.127 0.350 Kale Garden plant 10/7/20 0.511 0.223 0.288 Garden plant 11/18/20 1.064 0.418 0.646 Celery Garden plant 10/7/20 0.567 0.236 0.331 Garden plant 1 11/10/20 0.793 0.423 0.370 76 Missouri Agr. Exp. Sta. Research Bulletin 48 Relation of hot-water-soluble pentosans to the hardening pro- cess. — In jelly-making a hot water extract of fruits is used. Ac- cording to Goldthwaite 33 a cold water extract of our common fruits contains little or no pectin. The total pentosan determinations given in the four preceding tables indicate the larger content of pentosans in hardened plants, but in the total pentosans is included probably a more or less considerable amount of the insoluble hemi-celluloses of the cell wall, which might not be expected to function to any great extent as water-retaining material, though undoubtedly a part of the power of imbibition of the plant cell is due to its walls. The experience of jelly makers indicates that the hot water extract of fruit contained the most of the jelly-forming pectins. It was thought, therefore, that a hot water extract of the plant material would yield approximately that fraction of the total pentosan which exists in the protoplasm and might function as the significant water-retain- ing material. Accordingly, analyses were made from some of the samples, varying the procedure from that described for the total pentosan determinations as follows: The weighed sample of dry material was transferred to a beaker with 150 cc. of distilled water. The slight acidity was neutralized by adding a bit of sodium carbonate, then the material was boiled for five minutes, and filtered hot through a Gooch crucible. This yielded a clear cherry-colored filtrate, con- taining all the hot-water-soluble pentosans, sugars, and other soluble carbohydrates. Hydrolysis, fermentation, clearing and analysis were carried out with this filtrate in the same way as previously described for the whole sample in the total pentosan determinations. The re- sults are given in Table 17. In cabbage plants exposed to hardening treatment, the water- soluble pentosans increase considerably while the insoluble (hemi- cellulose) fraction is nearly constant, regardless of the degree of hardiness. In hardened cabbage plants the amount of soluble pen- tosans is relatively large, in fact the increase in the total pentosan content is very largely due to the increase in the water-soluble frac- tion. In tomatoes, on the other hand, the hot water soluble fraction is very small and does not increase much in plants subjected to hardening treatments. The relatively large amount of total pen- tosans in the tomato, therefore, is largely insoluble, probably exist- ing mostly as hemi-cellulose or in the middle lamella. The sweet potato resembles the tomato, in that it has a relatively large total The Hardening Process in Vegetable Plants. 77 pentosan content, but only a small soluble fraction. Tlie sweet po- tato, like the tomato, is very tender to frost and is not susceptible of much increase in cold-resistance upon exposure to usual conditions of hardening'. The water-soluble fraction of the total pentosan content in gar- den plants of kale and celery is shown to increase considerably as they become hardier in the fall. These differences in the soluble pentosan content may give us an important clue to the reason for the previously shown difference in cold-resistance, susceptibility to hardening and water-retaining power in the two groups of plants represented respectively by the cabbage and the tomato. Factors influencing the imbibitional capacity of plant colloids. — In view of the increase in hardened plants of pentosans, especially in the hot-water-soluble fraction, and the possibility of these sub- stances being at least partly responsible for the increased water-re- taining capacity of such plants, factors which influence the water- retaining power of these and other hydrophilous colloids occurring in plants may be of great importance in relation to cold-resistance. Fig. 9. — Swelling of Agar as influenced by reaction of the solution. 78 Missouri Agr. Exp. Sta. Research Bulletin 48 Acidity. — Fischer 30 showed that the power of imbibition of col- loids was influenced very markedly by the reaction of the medium, as demonstrated by his experiments in which slight acidity increased the swelling of gelatin. He was able to alleviate oedema of the eye and other animal tissues by application of alkali and hypertonic sugar solutions. Fischer regards acidosis as one of the most im- portant causes of the presence of abnormal amount of water in cells. Dachnowski 23 found that seeds of beans and corn swelled more and retained more w 7 ater in N/800 acids than in water, but the amount of water absorbed and retained was not proportional to the concen- tration of acid, for a maximum was attained beyond which increased acidity decreases absorption. The addition of equi-molecular solu- tions of non-electrolytes, such as glucose and sucrose, did not increase the amount of water retained by seeds in Dachnowski ’s experiments. The amino-acid, glycocoll, was a striking exception in that greatly increased imbibition by seeds took place in the presence of this sub- stance. Upson and Calvin 141 have shown that the mixture of vegetable proteins which comprises the gluten of wheat, behave in the same way as Fischer’s animal proteins. They obtained maximum absorp- tion of water in 0.01 N hydrochloric acid and 0.04 N acetic acid, with marked depression of absorption by strong acids and by salts. Mac Dougal and Spoehr 87 found a greater swelling of agar in N/100 solu- tions of the amino-acids glycocoll, alanin, and phenylalanin, than in water. The same workers have shown that the imbibition of protei- naceous colloids, such as gelatin, could be increased considerably by dilute acids, whereas colloids such as agar, having a pentosan base, swelled less in N/100 HC1 than in distilled water. However, brief series of tests made by the writer on the swelling of agar as influenced by the reaction, indicate that the greatest swell- ing of this material occurs in about N/5000 HC1. Presumably^ it would require a much greater concentration of the plant acids to bring about the same degree of swelling as such a dilute HC1 solution Alkalinity, excess acidity, and the presence of salts depressed the imbibitional capacity of agar very markedly. The results of a du- plicate series of tests performed with shredded agar are presented graphically in figure 9. The results obtained by Mac Dougal 82 indicate that a mixture of agar and gelatin would exhibit maximum swelling in somewhat stronger acid than would agar alone. Since colloids of the pentosan type probably occur in plants in intimate association with proteina- ceous colloids, it is reasonable to suppose that the greatest power of The Hardening Process in Vegetable Plants. 79 imbibition would be exhibited by plant cells in the presence of slightly increased acidity. Mac Dougal and Spoehr 88 suggest that the in- creased acidity found in succulent plants may be a characteristic of a metabolic complex favorable to pentosan formation and to the development of succulence (a high degree of water-holding power). In connection with the increased water-holding power of some colloids, associated with slightly increased acidity and especially some of the amino acids, it is interesting to note that Harvey 42 found a marked increase in amino-nitrogen in hardened cabbage. May it not be possible that in developing hardiness, plants form some specific amino acid which would increase the water-retaining power of the cells ? Somewhat greater titratable acidity has been found in hardened cabbage, as shown in Table 18. Determinations of the hydrogen-ion Table 18. — Titratable Acidity in Hardened and Tender Plants. (cc. N/10 NaOH per one gram dry material in 100 cc. water) Treatment Cabbage Lettuce (a) Tomato (b) Greenhouse plants, tender . . . . 1.60 1.86 0.96 1.74 Coldframe plants, hardy . . . In coldframe, 2 weeks . . . 2.06 . . . 1.68 3.06 1.74 Grown dry in greenhouse (hardy) Grown wet in greenhouse . . . 1.30 — t — 0.96 1.00 (tender) . . . 0.82 — 0.66 1.44 concentration were also made on a few samples, but little variation could be detected by the Gillaspie method. It seems that there is a slight increase in acidity in plants as a result of the hardening process. This change may take place only in plants possessing potential hardiness such as cabbage and lettuce since the data in Table 18 indicate no correlation between acidity and hardening treatments in the tomato. Increased acidity might also influence the water-retaining power of plant cells to such an extent as to account at least partly for the cold-resistance of hardened plants, aside from any increase in the amount of hydrophilous cell colloids. However, too few data are available to draw a definite conclusion on this point. Salts and sugars . — It has been shown by Fischer, Dachnowski, Mac Dougal and his co-workers that the addition of salts to a solution greatly decreases the imbibitional capacity of gelatin, seeds and 80 Missouri Agr. Exp. Sta. Research Bulletin 48 agar. However, Free 29 found that gelatin swells a little more in 0.5 percent solutions of dextrose and glucose than in water, while a dis- tinct decrease of swelling occurred in solutions of 25 percent or over. Agar was found to swell a little more in two-percent sucrose than in distilled water, whereas dextrose had little effect, except that it depressed swelling in concentrated solutions. That dilute sugar so- lutions do not decrease the imbibitional capacity of such hydrophilous colloids as gelatin and agar is important, since a greater sugar content is found in hardened plants. According to Goldthwaite 33 pectin and acid are prerequisites for j edification of fruit-juice, while sugar is a necessary accessory. She was able to make an excellent artificial jelly with one percent pectin, 0.5 percent tartaric acid, and three- fourths volume of cane sugar. Furthermore, in her experiments, it was shown that increasing the proportion of sugar gave an increased volume of jelly. The work of these investigators suggests the pos- sibility of increased acidity and sugar content playing an important part in determining the state of the colloidal protoplasm. Perhaps sudden or extreme changes in some of these factors, which influence imbibitional capacity, might exert an important in- fluence on the water-retaining power and cold resistance of plant tissue. However, the capacity of plant organs to take up or imbibe large amounts of water must not necessarily be taken as an index of their power to retain water when exposed to conditions favoring un- due water loss, such as freezing or drying. SUMMARY. The work of previous investigators indicates that water-loss from the cells, by the formation of ice crystals in the intercellular spaces, is most generally the limiting factor in the killing of plant tissue by cold. Any treatment materially checking the growth of plants increases cold-resistance. In cabbage and related plants, hardiness increases in proportion as growth is checked. In tomato and other tender spe- cies, the checking treatments resulted in relatively slight increase to cold-resistance. The various means of hardening plants in these experiments have resulted in about the same type of changes within the plant. Cabbage plants hardened by various treatments contain a larger amount of “ unfree, ” or not easily frozen water, as measured by the dilatometer. The increment in unfree water corresponds to the The Hardening Process in Vegetable Plants. 81 extent to which growth is checked, both of these paralleling the de- gree of cold resistance. The amount of water frozen at different temperatures in leaves of varying hardiness was measured. The percentage of moisture frozen in hardened cabbage leaves at -3°C. and at -4°C. is about two-thirds of that frozen in tender cabbage leaves at the same tem- perature. The actual amount of water remaining unfrozen at a given temperature is greater in hardened than in tender leaves, al- though their total moisture content is less. The percentage of total moisture frozen in leaves increases for each successive degree of temperature lowering, but the increase be- comes rapidly smaller and smaller. The amount of water remaining unfrozen in hardened cabbage leaves is approximately a logarithmic function of the temperature. Cabbage plants exposed to low temperatures in a coldframe for varying periods have a progressively smaller amount of w^ater freez- able at -5°C., the longer they are exposed to hardening. The per- centage of freezable water decreased quite rapidly in the first four days after removal from the greenhouse, more slowly from four to fourteen days and very slowly thereafter* The rate of decrease in percentage of freezable water coincides with the observed rate of hardening. In other words, the hardening process in cabbage plants was accompanied by a proportional increase in the amount of water unfrozen at -5°C. The amount of water frozen at -5°C. is somewhat less in plants exposed to slight wilting at midday. The effects of watering plants with M/10 salt solutions are as- sociated with a condition of mild physiological drought. The degree of such drought is proportional to the concentration of the soil solu- tion, which in turn is influenced by: (a) the amount of water-soluble material present and (b) the power of the soil to hold a large part of the soil moisture unfree in the pure or nearly pure state. Hardened cabbage plants lose less moisture by transpiration per unit of leaf area than tender plants, under the same conditions. The amount of water lost by transpiration per plant for a given period is much less in hardened cabbage plants than in non-hardened plants of the same age because of: (a) the lower rate of transpiration and (b) the smaller size of hardened plants. This accounts for the fact that hardened plants can be transplanted to the field with less wilting. The rate of water loss from hardened cabbage leaves dried in an oven at 60°C. is much less than that from leaves of tender plants. In tomato, the rate of drying is only slightly less in hardened than 82 Missouri Agr. Exp. Sta. Research Bulletin 48 in non-kardened plants. Comparing the rate of water-loss from to- mato and cabbage leaves, it is found that hardened tomatoes lose water somewhat faster than tender cabbage leaves. The lesser amount of water lost by ice formation, the lower rate of transpiration and the slower rate of water loss upon drying in hardened cabbage plants, may be explained by the hypothesis that hardening develops an increased water-retaining capacity. The water-retaining power of plant cells is due to: (a) Osmotic concen- tration (b) Imbibition, and may be increased by means of either or both of these factors. Osmotic concentration of plant cells may be increased by: (1) Decreasing the total water content. (2) Increasing the amount of osmotieally active sap solutes. (3) Decreasing the amount of free water or conversely, by in- creasing the amount of unfree water held by colloidal adsorption. Osmotic concentration as measured by the lowering of the freez- ing point has been found to increase on hardening plants, varying inversely with the water content. Both reducing and non-reducing sugars increase with hardening. Sugars are found to increase more in cabbage and lettuce than in tomato. The increased sugar is not sufficient to account for much difference in the freezing point depres- sion or in the amount of water remaining unfrozen several degrees below the freezing point. The chief factor in increasing osmotic concentration in plants is considered to be the decrease in amount of free water, hence the observed increase in osmotic concentration would be a secondary result of the hardening process. The power of imbibition possessed by plant cells may be increased by: (1) Decreasing the total water content (or increasing the per- cent of dry matter). (2) Increasing the amount of hydrophilous colloids in the pro- toplasm. (3) Increasing the water-retaining power of such colloids by slight increase in acidity, etc. Decreased water content accompanies a condition of greater cold resistance in plants. During the hardening process, the per- centage of dry matter increases rapidly for a few days, and more slowly thereafter. The total pentosan content is greater in hardened th&n in tender plants, regardless of the kind of hardening treatment. The pentosan content of cabbage plants exposed to low temperatures in an open coldframe during March increases rapidly the first five The Hardening Process in Vegetable Plants. 83 days and more slowly thereafter. The pentosan content of cabbage, kale and celery plants growing in the open garden increases as the weather becomes colder during the fall. In cabbage, kale and let- tuce plants possessing potential hardiness, the fraction of the pen- tosan content soluble in hot water is larger than in tomato, eggplant and sweet potato, which do not possess potential hardiness. The hot water-soluble pentosan content is thought to represent more nearly the amount of pentosans in the protoplasm and these might function more specifically as water-retaining material. In the group of plants susceptible of considerable hardening to cold the increase in total pentosan content upon hardening is largely an increase in the hot water-soluble fraction, while in the tomato the hot water- soluble fraction does not increase upon subjecting the plants to har- dening treatments. CONCLUSIONS. The experimental data show that the hardening process in plants is accompanied by a marked increase in water retaining power, and that this water retaining power is due chiefly to the imbibitional forces of the cell. The amount of water frozen in hardy plants is less than in tender plants and cells of hardy plants actually retain a larger amount of unfrozen water than those of tender plants. It is believed that cold resistance in plants is due to the increased water-retaining power of the cells, which enables them upon freezing to retain a larger proportion of their moisture content in the unfrozen condition. The increased water-retaining power of hardened plants is as- sociated with the following changes: (a) decreased moisture con- tent, (b) increased amount of hydrophilous colloids, such as pen- tosans, (c) increased water-retaining power of such cell colloids be- cause of a slight increase in acidity or other internal changes, (d) increased amount of osmotically active substances as soluble sugars. The last factor probably is important only in plants hardened by prolonged exposure to cold ; the first three factors mentioned may become operative in a very short time, when the activity of the plant is limited by any factor. Perhaps the same changes which increase the water-retaining power also favor greater stability of the proto- plasm. The marked parallelism between pentosan content and hardiness indicates a causal relationship. However, pentosan content alone is not to be taken as an absolute index of cold resistance, since several 84 Missouri Agr. Exp. Sta. Research Bulletin 48 factors may affect tlie functioning of pentosans as water-retaining substances. Salt content, acidity, hydrogen-ion concentration, sugar, moisture, protoplasmic colloids other than pentosans and perhaps, other factors constitute a varying complex which may influence water- retaining power and hardiness. The differential reactions, when subjected to hardening treat- ments, of plants possessing potential hardiness as the cabbage and of plants lacking it as the tomato, indicate that the fundamental difference between hardy and tender species lies in their ability to initiate changes whereby the stability and water-retaining power of the protoplasm and consequently hardiness are increased. Hardy species and varieties of plants possess the ability to initiate such changes to a greater or less great degree, while tender species pos- sess it to a very slight degree or not at all. APPLICATIONS. \ In view of the connection between cell water retaining power and hardiness which has been found and the correlation between soluble pentosan content and hardiness, it seems that problems deal- ing with cold resistance of vegetables, cereals, fruits and shrubs may be attacked from a new angle. Furthermore, the association of water-retaining power of cells with their content of a specific material or group of materials, such as i pijlfogu is, may be important in the study of moisture relations an(^^vfTter!4novement in plants. Moreover it may lead to a better under standing\of^the cause and prevention of a group of physiologi- cal plant diseases usually associated with excessive water loss, such as Tipburn of potato and lettuce, and Blossom End Rot of tomato. Selection of plants for high soluble pentosan content may be helpful to the breeder of cold-resistant, drought-resistant, or disease-resistant varieties of crop-plants. The changes of the food value of fruits and vegetables subjected to long storage may be significant, since it seems that in living plant tissues, exposed to water deficit or to cold the hexosan carbohydrates are converted into pentosans, which have a much lower coefficient of digestibility. However, the use of such vegetables as have a high water-retaining power may be important dietetically in the alleviation of certain digestive disorders. The Hardening Process in Vegetable Plants. 85 ACKNOWLEDGMENTS To Messrs. V. R. Gardner, H. D. Hooker, Jr., and F. C. Brad- ford of the Department of Horticulture and W. J. Robbins of the Department of Botany, I am indebted for suggestions, constructive criticisms and some of the references of the literature. The work on measurement of cells and on the swelling of agar was performed in the Botanical Laboratories under the direction of Dr. Robbins. The writer wishes to acknowledge gratefully the assistance, sug- gestions, criticisms, and kindly encouragement so generously extended by all of these gentlemen, without which this work could never have been performed. BIBLIOGRAPHY A more complete list of the older works on killing of plants by cold will be found in Chandler: Mo. Agr. Exp. Sta. Res. Bui. 8. 1. Abbe, C., Influence of Cold on Plants, a Resume. Exp. Sta. Record 6: p. 777, 1895. 2. Ackerman, A. Johnson, Hj. & Platon, B., Sveriges utsadesferenigs Tid- skrift, pp. 216-224, 1918. Rev. by Malte, M. O., Sugar Content and its Relation to Winter Hardiness; Agr. Gaz. of Canada, p. 329, April 1919. 3. Adams, J., The Effect of Very Low Temperature on Moist Seeds, Sci. Proc. Roy. Dublin Society, N. S. Vol. 11; p. 1, 1905. 4. Bartetzko, H., Untersuchungen uber das Erfrieren von Schimelpilzen. Jahrb. Wiss. Bot., Bd. 47, Heft 1, pp. 57-90, 1909. 5. Batchelor, L. D. & Reed, H. S., Winter Injury or Die-back of the Walnut. Calif. Agr. Exp. Sta., Circ. 216, 1919. 6. Beach, S. A. & Allen, F. W. Jr., Hardiness of Apple as Correlated with Structure and Composition, Iowa Agr. Exp. Sta., Research Bui. 21, 1915. 7. Bergen, J. Y., Transpiration of Sun-leaves and Shade Leaves of Olea europaea and other Broad Leaved Evergreens. Bot. Gaz., 38: pp. 285-296, 1904. 8. Boswell, V. R., Unpublished data, Univ. of Mo., Dept. Horticulture, 1921, 9. Bouyoucos, G. J., An Investigation of Soil Temperature and Some of the Most Important Factors Affecting It. Mich. Agr. Exp. Sta. Tech. Bui. 17, 1913. 10. — , Measurement of the Inactive or Unfree Moisture in the Soil by Means of the Dilatometer Method. Jour. Agr. Research, 8: No. 6, pp. 195-217, 1917. 11. , Classification and Measurement of the Different Forms of Water in the Soil by Means of the Dilatometer Method. Mich. Agr. Exp. Sta., Tech. Bui. 36, 1917. 12. , Degree of Temperature to Which Soils Can be Cooled With- out Freezing, Jour. Agr. Research 20: pp. 267-269, 1920. 13* , Concentration of Soil Solution Around the Soil Particles, Soil Sci., 11: pp. 131-138, 1921. 14 , & McCool, M. M., Determination of Cell Sap Concentration by the Freezing Point Method. Amer. Jour. Agron. 8: p. 50, 1916. 15. , Measurement of the Amount of Water that Seed Cause to Become Unfree, and Their Water-soluble Material. Jour. Agr. Research, 20: No. 7, pp. 587-593, 1921. 16. Briggs, R. G., Relation of Physical Structure of Fruit Buds of the Peach to Hardiness. Master’s Thesis, University of Mo. 1913. 18. Carrick, D. B., Resistance of the Roots of Some Fruit Species to Low Temperature, Cornell Univ. Agr. Exp. Sta., Memoir Bui. 36, 1920. 86 Missouri Agr. Exp. Sta. Research Bulletin 48 19. Cavallero, Sebastin: Gior. Agr., March 1888, also Gaz. Montava, Jan. 1891. Abs. in Exp. Sta. Rec., 6: p. 777, 1895. 20. Chandler, W. H., The Killing of Plants Tissue by Low Temperature, Mo. Agr. Exp. Sta., Res. Bui. 8, 1913. 21. r — , Sap Studies with Horticultural Plants, Mo. Agr. Exp. Sta., Research, Bui. 14, 1914. 22. Dachnowski, A., Physiologically Arid Habitats and Drouth Resistance in Plants, Bot. Gaz. 49: pp. 325-339, 1910. 23. , Effect of Acid and Alkaline Solutions upon the Water- Relation and the Metabolism of Plants. Amer. Jour. Bot. 1: pp. 412-435, 1 1914. 24. Davis, W. A. & Daish, A. J., The Estimation of Carbohydrates. Jour. Agr. Sci. 5: p. 437, 1913, also 6: 152, 1914. 25. De Candolle, quoted in Lindley, J.; “Theory of Horticulture.” Book, 2nd. American edition, by A. J. Downing., 1855. 26. Detmer, W., Influence of Moisture, Temperature and Light Conditions on the Process of Germination. In Rept. of International Meteorological Congress, Chicago, 1893. 27. Duhamel du Monceau, H. L. & Buffon, G. L. L., Observation des differ- ents effects que produsuent sur les vegetaux les grandes gelees d’hiver et les petetes gelees du printemps. Mem. Math, et Phys. Acad. Roy. Soc. (Paris) pp. 233-298, 1737. 28. Foote, H. W. & Saxton, B., The Effect of Freezing on Certain Inorganic Hydrogels, I. Jour. Amer. Chem. Soc. 38: pp. 588-609, 1916. II. Same, 39: pp. 1103-1125, 1917. 29. Free, E. E., Swelling of Agar and Gelatin Gels in Solutions of Sucrose and Dextrose. Science, N. S. 46: p. 142, 1917. 30. Fischer, M. H., Oedema, Book, Cincinnati, Ohio, 1910. 30A. r , & Sykes, A., Non-electrolytes and the Colloid-Chemical Theory of Water Absorbtion. Science, N. S. 38: pp. 486-487, 1913. 31. Gasner, G. & Grimme, C., Biettage zur Frage der frostharte der Getriedepflanzen, Ber. d. Deut. Bot. Gesell., 31: 507-516, 1913. 32. Geoppert, H. R., Uber der warme entwickelung in dem pflanzen; deren gefrieren und die schutzmittel gegen dasselbe. Book, 274. p. Breslau, 1839. Also see translation in Edinburgh, Jour. Nat. & Geol. Sci. 1831, p. 780. 33. Goldthwaite, N. E., Contribution on the Chemistry and Physics of Jelly Making, Jour. Indus. & Eng. Chem. 1: pp. 333-349, 1909, also 2: pp. 457-462, 1910. 34. Greeley, A. W., On the Analogy Between the Loss of Y/ater and Lower- ing the Temperature. Amer. Jour. Physiol., 6: pp. 122-128, 1901. 35. Gorke, H., Uber Chemische Vorgange beim erfrieren der Pflanzen. Landw. Vers. Stat., Bd. 65, Heft, i/ 2 , p. 149-160., 1906. 36. Groom, P., Bud Protection in Dictoyledens, Trans. Linn. Soc. II, 3: p. 255, 1893. 38. Haas, A. R. C., The Reaction of Plant Protoplasm. Bot. Gaz. 63: pp. 232-235, 1917. 39. Harris, J. A., On the Osmotic Concentrations of the Tissue Fluids of Phanerogamic Epiphytes. Amer. Jour. Bot. 5: pp. 490-506. 1918. 40. — , & Popenoe, W., Freezing Point Lowerings of the Leaf Sap of the Horticultural Types of Persea Americana. Jour. Agr. Res.. 7: pp. 261-268, 1916. 41. • , & Gortner, R. A., Calculation of Osmotic Pressure of Ex- pressed Vegetable Saps from the Depression of the Freezing Point. Amer. Jour. Bot. 1: p. 75, 1914. 42. Harvey, R. B., Hardening Process in Plants and Developments from Frost Injury. Jour. Agr. Res., 15: pp. 83-112, 1918. 43. Hartwell, B. L., Starch Congestion in Plants, R. I. Agr. Exp. Sta. Bui. 165, 1916. 44. Hedlund, T., Ueber die Moglichkeit, von der anusbildeng des weizens in Herbst, auf die winterfestigheit der verschiedenen sorten zu schlies- sen. Revieiced in Bot. Centralbl. 135: 222-224, 1917. The Hardening Process in Vegetable Plants. 87 45. Hibbard, R. P. & Harrington, O. E., Depression of the Freezing Point in Triturated Plant Tissues and the Magnitude of this Depression as Related to Soil Moisture. Phyiol. Researches, 1: pp. 441-454, 1916. 46. Hooker, H. D. Jr., Seasonal Changes in the Composition of Apple Spurs. Mo. Agr. Exp. Sta. Res. Bui. 40, 1920. 47. , Pentosan Content in Relation to Winter Hardiness. Proc. Amer. Soc. Hort. Sci. 1920, pp. 204-207. 48. Hornby, A. J., Pectins in Various Plants, Jour. Soc. Chem. Indus. 39: p. 246, 1920. 49. Irmscher, Edgar, Uber die Resistanz der Laubmoose gegen austrocknung und Kalte. Jahrb. F. Wiss. Botanik, 50: pp. 387-449, 1910. 50. Jones, L. R., Miller M., and Bailey, E., Frost Necrosis of Potato Tubers, Wis. Agr. Exp. Sta. Res. Bui. 46. 51. Johnson, E. S., An Index of Hardiness in Peach Buds. Amer. Jour. Bot. 6: pp. 373-379, 1919. 52. Kiesselbach, T. A., and Ratcliff, J. A., Freezing Injury of Seed Corn. Neb. Agr. Exp. Sta. Res. Bui. 16, 1920. 53. Koestian, C. F., Hartley, C., Watts, F., and Holm, G. G., A Chlorosis of Conifers Corrected by Spraying with Ferrous Sulfate, Jour. Agr. Res., 21: pp. 153-171, 1921. 54. Kraus, E. J. and Kraybill, H. R., Vegetation and Reproduction in the Tomato. Ore. Agr. Exp. Sta. Bui. 149, 1919. 55. Knudson, L., Influence of Certain Carbohydrates on Green Plants, Cornell Agr. Exp. Sta. Memoir Bui. 9: 1916. 56. Kylin, H., Cold Resistance in Marine Algae. Ber. Deut. Bot. Gesell, 35: pp. 370-384, 1917. 57. Leclerc du Sablon. Researches Physioiogiques sur les Matieres de Reserves des Arbres. Rev. Gen. Bot., 16: p. 41, 1904. 59. Lewis, F. J. and Tuttle, G. M., Osmotic Properties of Some Plant Cells at Low Temperature. Ann. Bot. 34: pp. 405-416, 1920. 60. Lidforss, B., Die Wintergrune Flora. Eine Biologische Untersuchung. Lunds Univ. Orsskrift, N. F. Bd. 2, Afd. 2, No. 13, 1907, Abs. in Bot. Centbl. Bd. 110, pp. 291-293, 1910. 61. Lindley, J., Philosophy of the Destruction of Plants by lrost. Trans. London Hort. Soc. Ser. 2, Vol. 2, Part IV. Reprinted in The Horticul- turist, 7: pp. 405-411, 1852. 62. Matruchot, L. and Molliard, M., Sur Certain Phenomena presentes par les Noyaux sous laction der froid. Comp. Rend. Acad. Sci. (Paris) 130: pp. 788-791, 1900. 63. Matruchot, L. and Molliard, M., Sur l’identite des Modifications de structure produites dans les cellules vegetales par le gel, a Plasmolyse, et la fanaison. Compt. Acad. Sci. (Paris) 132: pp. 495-498, 1901. 64. Matruchot, L. and Molliard, M., Modifications produites par le gel dans le structure des cellules vegetales. Rev. Gen. Bot. 14, pp. 463-482. 1902. 65. Maximow, N. A., Chemische Schutzmittel der pflanzen gegen erfrieren, Abstract in Ber. Deut. Bot. Gesell, Bd. 30: (1) pp. 52-65. (2) pp. 293-305. (3) pp. 504-516, 1912. 66. • , Experimental^ und Kritische untersuchengen uber das gefrieren und erfrieren der planzen, Jahrb. Wiss. Bot. Bd. 53: Heft. 3, pp. 327-420. 67. Mer, E., De la Constitution et des functions des feules Hibernalis. Bill. Soc. Bot. France, 23: p. 231, 1876. 68. Mez, C., Einige Pflanzengeographische folgerungen aus einer neuen theorie uber das erfrieren eis-bestandiger pflanzen. Bot. Jahrb. (Engler) Bd. 34: pp. 40-42, 1905. 69. Michel-Durand, E„ Variation des substances hydrocarbonees dans les feuilles. Rev. Gen. Botanique, 31: pp. 53-60; pp. 143-156; pp. 250-268; pp. 286-317; 1919. 70. Miyake, K., On the Starch of Evergreen Trees and its Relation to Photo- synthesis During the Winter. Got. Gaz. 33: pp. 321-340, 1902. 71. Molisch, Hans, Untersuchung uber das erfrieren der pflanzen, Book. 1897. 88 Missouri Agr. Exp. Sta. Research Bulletin 48 72. ? , Erfrieren der pflanzen. Vertrage des Verins zur Verbrei- tung Naturwissenschaftlechen Kenntnisse in Wien, 51 Jahrgang, Heft, 6, 1910. 73. Morren, Bulletin de l’academie Royal de Bruxelles-Vol. 5, Quoted by Bindley in the Horticulturist 7: p. 406, 1852. 74. Muller, H., (Thurgau)., Ueber das gefrieren und erfrieren der pflanzen, Landw. Jahrb. 9: pp. 133-189, 1880. 75. , Ueber Zuckeranhaufig in pflanzentheilen infolge niederer temperatur. Landw. Jabrb, 11: pp. 751-828, 1882. 76. , Ueber das gerfrieren und erfrieren der pflanzen (II Thiel), Landw. Jahrb, 15: pp. 453-610. 78. Me Cool, M. H. and Millar, C. E., Water Content of the Soil and the Composition and Concentration of the Soil Solution as Indicated by the Freezing Point Lowerings of the Roots and Tops of Plants. Soil Sci. 3: pp. 113-138, 1917. 78. — , Further Studies in the Freezing Point Lowerings of Soils and Plants, Soil Sci. 9: pp. 217-233, 1920. 80. , Use of the Dilatometer in Studying Soil and Plant Rela- tionships. Bot. Gaz. 70: pp. 317-319, 1920. 81. MacDougal, D. T., Year Book No. 17, p. 56-57, Carnegie Institute of Wash- ington, 1918. 82. , Imbibitional Swelling of Plants and Colloidal Mixtures, Science, N. S., 44: pp. 502-506, 1916. 83. < , Auxographic Measurements of Swelling of Biocolloids and of Plants, Bot. Gaz. 70: pp. 126-136, 1920. 84. , Colloidal Reactions Fundamental to Growth, Science, N. S. 51: pp. 68-70, 1920. 85. ■ — , Pub. 297, Carnegie Inst. Washington, 1920. 86. , and Richards, H. M., and Spoehr, H. A., Basis of Suc- culence in Plants, Bot. Gaz. 68: p. 405-416, 1919. 87. • , and Spoehr, H. A., Swelling of Agar in Solutions of Amino Acids and Related Compounds, Bot. Gaz. 70: pp. 268-276, 1920. 88. , Origin and Physical Basis of Succulence in Plants, Car- niegie Inst. Washington, Yearbook 17, pp. 85-86, 1918. 89. Nageli, Ueber der Wirkung des frostes auf die Pfianzenzellen, sitz der Konig Bayer, Akad, d. Wiss. Munchen, I. p. 264, 1861. 90. Nelson, A., The Winterkilling of Trees and Shrubs, Wyoming Agr. Exp. Sta. Bui. 15, 1893. 91. Nicholas, G. R., Relationship between Leaf Anthocyanin and Respira- tion, Rev. Gen. Bot. 31: pp. 161-178, 1919. ( abs . in Exp. Sta. Record, 42: p. 227, 1920). 92. Ohlweiler, W. W., Relation Between the Density of the Cell Sap and the Freezing Point of Leaves. Ann. Rept. Mo. Bot. Gard. 23: pp. 101- 131, 1912. 93. Osterhaut, W. J. V., Effect of Anesthetics on Permeability, Science, N. S. 37: pp. 111-112, 1913. 94. Pantanelli, E., Influence of Nutrition and the Root Activity on the Collapse and Desiccation Produced by Cold. Atti R. Acad. Lincei, 5 Ser. 29: pp. 5771, 1920. (Abs. in Chemical Abstracts, 14: pp. 26-53). 95. — , The Resistance of Plants to Cold. Atti. R. Acad. Lincei. 5 Ser. No. 27: pp. 148-153, 1918. 96. — , Alterations in Cellular Permeability and Exchange at Temperature Near Freezing. Atti. R. Acad. Lincei, 5 Ser. V. 28; pp. 205-209, 1919. 97. Parker, F. W., Effect of Finely Divided Material on the Freezing Point Depression of Water, Benzene, and Nitrobenzene. Jour. Amer. Chem. Soc. 43: pp. 1011-1018, 1921. 98. Potter, G. F., Freezing of Apple Roots. In Wis. Agr. Exp. Sta. Bui. 319, p. 29, 1920. 99. Prilleaux, E., Sur la formation de glacons a l’interiur des plantes. Ann. Sci. Nat. Ser. 5, 12: p. 125, 1869. (Quoted by Wieeand in Plant World, 9: p. 25). The Hardening Process in Vegetable Plants. 89 100. , De l’influence de la congelation sur le poids des tis- sues vegetaux. Compt. Rend. Acad. Sci. (Paris) 74: pp. 1344-1346, 1872. 101. Prunet, Quoted "by Abbe in Exp. Sta. Rec. 6: p. 777, 1894. 102. Ravenna and Cereser, Origin and Physiological Function of Pentosans in Plants, Atti. R. Acad. Lincei, Ser. 5, 18: p. 177, 1909. (Abs. in Jour. London Chem. Soc., 96: p. 1046, 1909). 103. Rivera, U., Uber dies Ursach des lagums beim Weizem, Internat. Agri. Tulin. Rundschau, 7: p. 524, 1916. 104. Rosa, J. T. Jr., Pentosan Content in Relation to Hardiness in Vegetable Plants. Proc. Amer. Soc. Hort. Sci., pp. 207-210, 1920. 105. Sachs, J., Krystallbildungen bei dem gefrieren und veranderung der zellhaute bei den aufthauen saftige pflanzenthiele. Landw. Versuch. 2: Heft. 5, pp. 157-201, 1860. 106. , Ueber die Ausseren Temperaturen der pflanzen Flora, 1864, p. 37, (Quoted by Chandler). 107. — , Textbook of Botany-English Edition, by S. H. Vines. 108. Salmon, S. C., Why Cereals Winterkill, Jour. Amer. Soc. Agron, 9: pp. 353-379, 1917. 109. and Fleming, F. L., Relation of the Density of Cell Sap to Winter-Hardiness in Small Grains. Jour. Agr. Res. 13: pp. 497-506, 1918. 110. Schaffnit, E., Studien Ueber den Einfluss nieder temperature auf die pflanzliche Zelle. Mitt. Kaiser Wilhelms Inst. Landw. Bromberg, 3: pp. 93-115, 1910. 111. Schulz, E., Uber Preserwestoffe in immergrunen Blattern, Flora, 71: p. 223, 1888. 112. Schimper, A. F. W., Plant Geography upon a Physiological Basis. Trans, by W. R. Fischer, p. 25-41, The Clarendon Press. Oxford, 1903. 113. Seifriz, William, Viscosity of Protoplasm as Determined by Microdis- section. Bot. Gaz. 70: pp. 360-378, 1920. 114. Schutt, F. T., On the Relation of Moisture Content to Hardiness in Apple Twigs, Proc. & Trans. Royal Soc. Canada 11, 9: Sec. IV, pp. 149- 153, 1903. 115. Sinz, E., Beziechungen zwischen Trocksubstanz und Winterfestighalt bei verschiedenen winter-weizen Varietatur. Jour. Landw. 62, pp. 301- 335, 1914. (Abs. in Exp. Sta. Record, 33: 235, 1915.) 116. Spoehr, H. A., Carbohydrate Economy of Cacti. Publication 287, Car- negie Inst. Washington, 1919. 117. Strassbaugh, P. D., Dormancy and Hardiness in the Plum. Bot. Gaz. 71: pp. 337-357, 1921. 118. Storber, J. P., Comparative Study of Winter and Summer Leaves of Various Herbs. Bot. Gaz. 63: pp. 89-111, 1917. 119. Swartz, Mary D., Nutrition Investigations on the Carbohydrates of Lichens, Algae, and Related Substances, Trans. Conn. Acad. Arts and Sci. 16: pp. 247-382, 1911. 120. Tuttle, G. M., Induced Changes in Reserve Materials in Evergreen Herbaceous Leaves, Ann. Bot. 33: pp. 201-210, 1919. 121. Uphof, J. C. Th., Cold Resistance in Spineless Cacti. Ariz. Agr. Exp. Sta. Bui. 70, 1916. 122. Vass, A. F.. Influence of Low Temperature on Soil Bacteria. Cornell Univ. Agr. Exp. Sta. Memoir Bui. 27, 1919. 123. Voightlander, H., Unterkuhlung und Kaltetod der pflanzen, Beitr, Biol. Pflanzen, Bd. 9, Heft, 31. 124. Walster, H. L., Formative Effect of High and Low Temperature upon Growth of Barley, Bot. Gaz. 69: pp. 97-126, 1920. 126. Weaver, J. E. and Morgensen, A., Relative Transpiration of Coniferous and Broad Leaved Trees in Autumn and Winter. Bot. Gaz. 68: pp. 393- 424, 1918. 127. Webber, H. J. et al. Effect of Freezes on Citrus in California. Calif. Agr. Exp. Sta. Bui. 304, 1919. 128. West, F. L., and Edlefsen, N. E., Freezing of Peach Buds. Utah Agr. Exp. Sta. Bui. 151. 90 Missour Agr. Exp. Sta. Research Bulletin 48. 129. Wiegand, K. M., Some Studies Regarding the Biology of Buds in Winter. Bot. Gaz., 41: pp. 373-424, 1906. 130. , Occurrence of Ice in Plant Tissue. Plant World 9: p. 25, 1906. 131. , The Passage of Water From the Plant Cell During Freez- ing. Plant World, 9: pp. 107-118, 1906. 132. Wright, R. C. and Taylor, G. F., Freezing Injury to Potatoes when Undercooled.. U. S. Dept. Agric., Dept. Bui. 916, 1921. 133. Dixon, H. H., Transpiration and the Ascent of Sap. In Prog. Rei. Bot 3: pp. 1-66, 1910. 134. Drabble, E. and Drabble, H., The Osmotic Strength of Cell Sap in Plants growing under Different Conditions. New Phytologist, 4: pp. 189-191, 1905. 135. Ewart, A. J., On the Power of Withstanding Desiccation in Plants. Proc. Liverpool Biol. Soc. 11: pp. 151-159, 1897. 136. Levene J., and Jacobs, W., Ueber die Pankreas-Pentose, Ber. d. deut. Chem. Gesell, 43: 3147-3150, 1910. 137. Tollens. Untersuchungen uber Kohlenhydrate. Landw. Versuchs-Sta- tionen, 39: p. 401, 1891. (Quoted by Swartz, see Bib. No. 119). 138. Livingston, E., Role of Diffusion and Osmotic Pressure in Plants. Pamphlet, 75p., Chicago, 1903. 139. Reinke, Quoted by Pfeffer-Physiology of Plants, Vol. 1, p. 73. 140. Pfeffer, W., Physiology of Plants, Second Edition. English Trans, by A. J. Ewart, Vol. 1, p. 73-75, Oxford, 1900. 141. Upson F. W. and Calvin, J. W., The Colloidal Swelling of Wheat Gluten in Relation to Milling and Baking. Nebraska Agr. Exp. Sta., Research Bui. 8. The Hardening Process in Vegetable Plants 91 Plate 1 . — Effect of Exposure in Open Frames on Coed Resistance. A. Coldframe hardened vs. greenhouse plants frozen at -4°C. for 2 '/, hours. Nov. 17, 1919. Vi. Cabbage plants after freezing at -8°C. for 2 y z hours, March 28, 1921. (1) Hardened in coldframe two weeks. Lower leaves broken off for samples. (2) Non-hardened greenhouse plant. 92 Missouri Agr. Exp. Sta. Research Bulletin 48 Plate 2. — Effect of Variation in Soil Moisture on Cold Resistance of Cabbage. A. Plants grown in greenhouse with varying supply of water; after freez- ing at -4°C. for 2y 2 hours. Nov. 17, 1919. (1) Dry grown (2) Medium dry (3) Wet grown. B. (1) Medium-dry-grown greenhouse cabbage plants. (2) Medium wet grown greenhouse cabbage plants after freezing at -4°C. for 30 minutes, March 28, 1921. C. After freezing at -4°C. for 30 minutes, March 28, 1921. (1) Watered heavily until one week before this test, thereafter wilted slightly for five days. (2) Plant from same batch as (1) but not subjected to preliminary wilting. The Hardening Process in Vegetable Plants. 93 Plate 3. — Effect of Varying Soil Moisture on Hardiness of Tomato. A. Greenhouse tomato plants after freezing at -2°C. for 2 hours. Sept. 29, 1919. II. (1) Dry-grown Greenhouse tomato plants after Sept. 22, 1921. (1) Dry-grown (2) Wet-grown, freezing at -2.25°C. for 2/4 hours, (2) Medium-drv-grown (3) Wet-grown. 94 Missouri Agr. Exp. Sta. Research Bulletin 48 Plate 4 — .Effect of Watering Plants Grown in Sand in Greenhouse With M/10 Salt Solutions. A. After freezing at -6°C. for 30 minutes. (1) NaCl (2) KC1 (3) NaNO, B. After freezing at -3°C. for 30 minutes (1) NaCl, (2) KC1. (3) NaNO, (4) Tap water (4) Tap water The Hardening Process in Vegetable Plants. 95 Pi.ate 5. — Effect of Watering Cabbage Plants Grown in Greenhouse With M/10 Salt Solutions, A. Grown in compost soil and watered with: (1) NaCl (2) KC1 (3) NaNO, (4) Tap water. After freezing at -6°C. for 30 minutes. II. Grown in Compost soil plus rotten manure, watered with: (1) NaCl (2) KC1 (3) NaNO :f (4) Tap water. After freezing at -6°C. for 30 minutes. 96 Missouri Agr. Exp. Sta. Research Bulletin 48 Plate 6. — Relative Wilting of Hardened and Tender Cabbagei Plants. A. Cabbage plants from transpiration experiment No. 4 after 5 hours of exposure to high transpiration conditions. (1) Greenhouse plant, watered with M/10 NaCl, hardy. (2) Greenhouse plant, watered sparingly with tap water, hardy. (3) Hardened in coldframe 5 days, hardy. (4) Greenhouse plant watered heavily, tender. B. Coldframe hardened cabbage plants C. Greenhouse non-hardened one day after transplanting to field, plants, handled other- March 27, 1918, weather fair, warm, wise the same as those dry. in B. The Hardening Process in Vegetable Plants. 97 Plate 7. — Tender Cabbage Plant From Greenhouse Frozen at -5°C. Fob 30 Minutes, March 31, 1921. Droplets of water exuding from stem and petioles upon thawing. The leaves were covered with a film of smaller droplets. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 49 EXPERIMENTS IN FIELD PLOT TECHNIC FOR THE PRELIMINARY DETERMINATION OF COMPARATIVE YIELDS IN THE SMALL GRAINS (Publication authorized December 2, 1921.) COLUMBIA, MISSOURI DECEMBER, 1921 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY, P. E. BURTON, H. J. BLANTON, St. Louis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION F. B. MUMFORD, M. S., DIRECTOR J. C. JONES, Ph. D., LL.D., PRESIDENT OF THE UNIVERSITY AGRICULTURAL STATION STAFF DECEMBER, 1921 CHEMISTRY RURAL LIFE O. R. Johnson, A. M. S. D. Gromer, A. M. E. L. Morgan, A. M. C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, A. M. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. Sieveking, B. S. in Agr. AGRICULTURAL ENGINEERING J. C. Wooley, B .S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. E. F. Hopkins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale. B. S. in i\gr. W. W. Swett, A. M. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. C. McBride, FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, A. M. O. W. Letson, B. S. in Agr. B. M. King, B. S. in Agr. A. C. Hill, B. S. in Agr. Miss Bertha C. Hite, A. B. 1 Miss Pearl Drummond, A. A. 1 Ben H. Frame, B. S. in Agr. horticulture V. R. Gardner, M. S. A. H. D. Hooker, Jr., Ph. D. J. T. Rosa, Jr., M. S. F. C. Bradford, M. S. H. G. Swartwout, B. S. in Agr. POULTRY HUSBANDRY H. L. Kempster, B. S. Earl W. Henderson SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M. W. A. Albrecht, Ph. D. F. L- Duley, A. M. 3 R. R. Hudelson, A. M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr. Richard Bradfield, A. B. O. B. Price, B. S. in Agr. veterinary science J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Sercretary S. B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsham, Librarian E. E. Brown, Business Manager *In service of U. S. Department of Agriculture, Seed Testing Laboratory. 2 On leave of absence. CONTENTS Page The Problem 6 Plan and Method of Investigation 9 Terminology 9 Procedure 11 Work of 1919 12 Work of 1920 16 Work of 1921 18 Competition as a Source of Error in Preliminary Tests 23 Previous Investigation 23 Experimental Results 25 Illustrations of Effects of Competition 26 Relation of Competition to Various Characteristics of the Com- peting Varieties 31 Discussion 40 Size and Replication of Plots 43 Previous Investigation 43 Experimental Results 44 Size of Plots 44 Replication of Plots 50 Adjustment of Yields by Means of Check Plots 54 Previous Investigation 54 Experimental Results 56 Method Used in Adjusting Yields 58 Relative Variability of Actual and Adjusted Yields 60 Difference in Results Obtained by Adjustment with Different Check Varieties 63 Value and Limitations of Adjusting Yields by Means of Check Plots 71 Concluding Remarks 73 Summary 75 Acknowledgment 77 References Cited 78 TABLES Table Number Table Page 1 Yields of Barley Varieties 1919 13 2 Yields of Oats Varieties 1919 14 3 Yields of Oats Strains 1919 15 4 Yields of Wheat Varieties 1920 16 5 Yields of Wheat Varieties 1921 17 6 Yields of Wheat Varieties and Mixtures 1921 18 7 Yields of Oats Varieties 1921 21 8 Yields of Oats Strains 1921 22 9 Relative Yields of Two Small Grain Varieties When Compared in Al- ternate Rows and in Blocks (Kiesselbach) 24 10 Correlation of Competition with Various Characteristics in Barley Va- riety Test 1919 35 11 Correlation of Competition with Various Characteristics in Oats Va- riety Test 1919 35 4 12 13 14. 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 Tables Correlation of Competition with Various Characteristics in Oats Strain Test 1919 36 Correlation of Competition with Various Characteristics in Wheat Va- riety Test 1920 . . 37 Correlation of Competition with Various Characteristics in Wheat Va- riety Test 1921 37 Correlation of Competition with Various Characteristics in Wheat Mix- ture Test 1921 38 Correlation of Competition with Various Characteristics in Oats Va- riety Test 1921 39 Summary of Effects of Competition in All Tests 41 Correlation of Yield with Dates of Heading and Maturity in Variety Tests of Barley, Oats, and Wheat 42 Yield and Variability of 1-row, 3-row, and 5-row Check Plots in Bar- ley Variety Test 1919 45 Yield and Variability of 1-row, 3-row, and 5-row Check Plots in Oats Variety Test 1919 46 Yield and Variability of 1-row, 3-row, and 5-row Check Plots in Oats Strain Test 1919 47 Yield and Variability of 1-row, 3-row, and 5-row Check Plots in Wheat Variety Test 1920 47 Yield and Variability of 3-row and 5-row Check Plots in Wheat and Oats Test 1921 48 Yield and Variability of 3-row and 5-row Test Plots in All Tests 50 Relation of Plot Variability to Size of Experiment Field in Wheat Va- riety Test 1920 51 Relation of Plot Variability to Size of Experiment Field in Wheat Va- riety Test 1921 52 Relation of Plot Variability to Size of Experiment Field in Oats Va- riety and Strain Tests 1921 52 Soil Heterogeneity of an Experiment Field as Determined from Yields of Two Check Varieties 53 Effect on Plot Variability of Adjusting Yields by Check Plots (Kies- selbach) 55 Reduction of Variability by the Use of Check Plots Equivalent to That Probably Attainable with the Same Number of Plots by Replication . 57 Relative Variability of Actual and Adjusted Yields in Barley Variety Test 1919 59 Relative Variability of Actual and Adjusted Yields in Oats Variety Test 1919 60 Relative Variability of Actual and Adjusted Yields in Oats Strain Test 1919 61 Relative Variability of Actual and Adjusted Yields in Wheat Variety Test 1920 62 Relative Variability of Actual and Adjusted Yields in Wheat Variety Test 1921 64 Relative Variability of Actual and Adjusted Yields in Wheat Mixture Test 1921 ....65 Relative Variability of Actual and Adjusted Yields in Oats Variety Test 1921 66 Relative Variability of Actual and Adjusted Yields in Oats Strain Test 1921 67 Relative Variability of Actual and Adjusted Yields of Kherson and Red Rustproof Oats Each in 120 Distributed Plots, in Oats Variety and Strain Tests 1921 68 Summary of Relative Variability of Actual and Adjusted Yields of Interior Rows in All Tests 1921 71 EXPERIMENTS IN FIELD PLOT TECHNIC FOR THE PRELIMINARY DETERMINA- TION OF COMPARATIVE YIELDS IN THE SMALL GRAINS* L. J. Stadler During recent years the investigation of the reliability of field experiments has become an important phase of agronomic research. Field experiments as ordinarily conducted have been shown to be affected by many gross errors. In the light of these investigations it has become apparent that the results of many of the older experiments are inconclusive or even misleading. Various expedients have been suggested for counteracting experimental error. Some of these have been quite successful, while others have probably done more harm than good. The pioneer investigations in this field have been of great value in directing attention to the important sources of error and in suggesting possible means for their control. Doubtless at the present time most of the major sources of error are recognized. But the true extent of the errors and the actual practical value of the methods of counteracting them can be determined only by numerous investiga- tions of experimental methods under different conditions. The present paper is concerned with experimental error and field plot technic in preliminary variety and strain tests with the small grains. The same type of test is extensively used in small grain im- provement, not only in the preliminary testing of varieties, but also in the comparison of strains and selections. Although the small plot test is particularly subject to errors of certain sorts, it has a decided advantage over tests in larger plots in the possibility of extensive replication, which is probably the greatest single factor in the reduc- tion of experimental error. It should be possible, consequently, to obtain extremely accurate results in small plot tests without the use of large experimental areas, when the errors peculiar to the small plot are understood and controlled. *Also submitted as a thesis in partial fulfilment of the requirements for the degree of Doctor of Philosophy. ( 5 ) 6 Missouri Agr. Exp. Sta. Research Bulletin 49 THE PROBLEM. At present the type of plot most commonly used for the pre- liminary testing of small grain varieties and strains is probably the “rod-row.” The methods of conducting rod-row tests described by Love and Craig 8 may be considered typical. The varieties or strains are sown by hand in rows one foot apart, usually opened and covered with a wheel hoe or similar implement. The seed for each row is weighed out in a quantity equivalent to ordinary rates of seeding in field practice. In harvesting, six inches or a foot at the end of the row is discarded, to prevent increase in yield by reason of the more favorable space conditions at the ends of the rows. The list of va- rieties is repeated in several series, and the results averaged to reduce the error from plot variability. A check variety is grown in every tenth row to indicate the variability of the field. The use of rod-row tests involves several errors, derived principally from the modified conditions under which the plants are grown. The object of the test is to discover the relative value of the strains under field conditions, and therefore any modification of field conditions which may favor some sorts more than others introduces error. The wide spacing between rows, with consequently heavier seeding in the row for any given rate of planting; the hand seeding and covering, re- sulting usually in slightly ridged rather than slightly furrowed rows; and the growing of different varieties in single rows, in competition with other varieties rather than with their own kind, are examples of typical conditions which may be expected to favor some varieties more than others. Consequently the best varieties in the rod-row test are not necessarily the best varieties under field culture, even when soil and seasonal variability are reduced to the minimum by replication of plots and repetition of the test through a series of seasons. Such sources of error as those mentioned do not necessarily affect the variability of the yields of replicate plots, as Kiesselbach 5 has pointed out, and are therefore more likely to escape notice. They are systematic errors affecting the yields of replicate plots similarly. Marked superiority of Turkey wheat over Fulcaster in a variety test in Kansas does not indicate the superiority of Turkey over Fulcaster in Illinois, no matter how low plot variability in the variety test may be, because the growing conditions in Illinois are different from the growing conditions in Kansas. Similarly the superiority of Turkey wheat over Fulcaster in a rod-row test may not mean its superiority under field conditions in the same locality, because here again growing Experiments in Field Plot Technic 7 conditions are different. The error in applying the results, though of course much less in degree, is similar in kind. And, since the rod-row test has no purpose but to indicate the relative value of the strains tested, for field conditions, any pronounced tendency to favor some varieties at the expense of others is fatal to its object. Ordinarily, however, the rod-row test is only the first stage in variety testing, and final recommendations are based upon results of tests under conditions which approach those of field culture more closely. When the elimination of varieties in the rod-row tests is not extremely strict a considerable latitude may be allowed, and under these conditions the rod-row test has served a valuable purpose. It is of course desirable nevertheless to reduce these errors to the greatest possible extent. Probably the most important of the errors mentioned is that arising from the competition between different varieties, in the single-row test. Obviously a variety grown in a single row between two different va- rieties may yield considerably more or less than the same variety grown between two rows of its own kind. Various expedients for re- ducing varietal competition have been suggested. Sometimes the order of varieties is changed in each series to bring together different va- rieties and thus tend to equalize the effects of competition ; sometimes an attempt is made to grow the varieties in such order as to bring together those of similar habit, and thus to reduce the effects of competition. Probably the most effective method is to grow border rows which may be discarded, and some investigators therefore use three-row or five-row blocks, in which the outer row on each side is discarded. The principal objection to the use of border rows in the increased area required to test the same number of strains, and the large pro- portion of the crop which is not harvested for yield. This is par- ticularly true when 3-row blocks are used, since in this case two- thirds of the field is used for border protection. The border rows may be used for seed, but two-thirds of the field is of course much more than is required ordinarily for this purpose. When 5-row blocks are used the proportion of the crop harvested for yield is increased from one-third to three-fifths, though it is an increase in size of plot, with some decrease in replication, so that there may be no gain in accuracy. There is a possibility that the effect of competition on the yield of 5-row blocks may be slight enough to permit the harvesting of all five rows for yield, particularly if the varieties may be effectively arranged for the reduction of competition. At any rate, in such plots the error from competition may be expected to be much less than that in single- 8 Missouri Agr. Exp. Sta. Research Bulletin 49 row plots, since only two of the five rows are subject to competition with a different variety, and each of these is subject to such compe- tition on only one instead of on both sides. Another phase of the question which should not be overlooked is the effect of adding border rows on the error from soil variability. If, for example, each rod-row is to be protected from competition by two border rows, the test will require three times as large a field as the same test without the border rows. This can hardly fail to in- crease materially the variability of the yields of replicate plots, to an extent which will vary with the uniformity of the field concerned. The use of border rows may thus necessitate the use of an even greater number of replications for the same degree of accuracy, as far as plot variability is concerned. It is possible that 3-row plots (whether or not provided with border rows) may require less replications for a given degree of accuracy than single-row plots, on account of their larger size. It is possible also that 5-row plots, because of their size, may have an advantage over 3-row plots in reducing va- riability, great enough to justify in practice harvesting all five rows for yield, rather than harvesting the interior three rows and discarding the border rows. The importance of any practice that will reduce the variability of the replicate plots is thus increased when border rows are introduced. A familiar method for this purpose is the adjustment of yields by means of distributed check plots. In following this method the yields of check plots are considered measures of the productivity of the soil, which is usually assumed to vary uniformly between them. The yields of the experimental plots are adjusted on the basis of uniform productivity of the field as a whole. Of late this method has rather lost favor among agronomists. In some cases the adjustment actually increases rather than decreases the variability of the replicate ex- perimental plots. Check plots have not been used extensively in ad- justing yields in rod-row tests, principally because of the great in- crease in computation necessary in adjusting the yields of such a large number of plots. Experiments in Field Plot Technic 9 PLAN AND METHOD OF INVESTIGATION The experiments here reported were designed to obtain informa- tion on several factors affecting the accuracy of preliminary variety and strain tests, with a view to devising, if possible, an improved technic for this important phase of crop improvement work. The data obtained bear directly on the following points : 1. The extent of error from varietal competition in bor- der rows, and the relation of such competition to the charac- teristics of the varieties, 2. The relative variability of plots of 1, 3, and 5 rows, and the number of replications necessary for a given degree of precision with plots of the three sizes, and 3. The effect on variability of adjusting yields by means of check plots. Terminology. — In this report the term plot will be used to des- ignate an area on which a single variety or strain is grown, in com- parison with other varieties or strains, in other plots. The plot may consist of one or more rows. A plot of more than one row may also be referred to as a block. The single outside rows of the block are the border rows. A single-row plot protected from competition by border rows, which are to be discarded, will be spoken of as a protected single-row plot. A protected single-row plot is therefore a 3-row plot with border rows discarded, and a protected 3-row plot is a 5-row plot with border rows discarded. The phrase “3-row plots replicated five times” will be used to refer to 3-row plots in five systematically dis- tributed locations, not in six. The area on which a complete variety or strain test is conducted is spoken of as an experiment field, or simply a field. A group of plots including one plot of each variety or strain tested is a series. When four replications are used there are four series of plots. The group of contiguous plots from one side of the field to the other constitutes a range. The ranges are separated by alleys. Thus the field shown in figure 1 consists of sixteen ranges, each range including twenty-nine 5-row (or protected 3-row) plots. Ninety- six varieties were tested on this field, each replicated four times. Ranges I to IV, inclusive, make up the first series, V to VIII the sec- ond, IX to XII the third, and XIII to XVI the fourth. Each of the four strips running lengthwise of the field and separated by the check plots may also be considered a series. All yields are expressed in bushels per acre by weight, computed on the basis of 60 pounds per bushel for wheat, 48 pounds for barley, 10 Missouri Agr. Exp. Sta. Research Bulletin 49 B CK I 17 33 43 65 81 CK 5 2/ 37 53 69 65 CK 9 25 41 57 73 89 CK ft 29 45 61 77 93 CK B CK 2 18 34 50 66 62 CK 6 22 38 54 70 86 CK 10 26 42 58 74 90 CK 14 30 46 62 7 8 94 CK 8 CK 3 IS 35 5/ 67 83 CK 7 23 3 9 55 7/ 87 CK II 27 43 59 75 91 CK 15 31 47 63 79 95 CK B CK 4 20 36 52 68 64 CK 8 24 40 56 72 63 CK 12 28 44 60 76 92 CK 16 32 48 64 80 96 CK B CK 5 2/ 37 53 69 85 CK 9 25 41 57 73 83 CK 13 29 45 61 77 93 CK 17 33 49 65 8/ CK B CK 6 22 33 54 70 86 CK 10 26 42 58 74 90 CK IK 30 46 62 78 94 CK 2 18 34 50 66 82 CK B CK 7 23 39 55 71 87 CK // 27 43 59 75 91 CK 15 31 47 63 79 95 CK 3 19 35 51 67 83 CK B CK <9 24 45 56 72 88 CK !2 28 44 60 76 92 CK J6 32 4 8 64 80 96 CK 4 20 36 52 68 84 CK 3 CK 9 25 4/ 57 73 89 CK 13 29 45 6/ 77 93 CK I 17 33 49 65 8! CK 5 21 37 53 69 85 CK B CK 10 26 42 58 74 SO CK /4 30 4 6 62 78 94 CK 2 18 34 50 66 82 CK 6 22 38 54 70 86 CK B CK // 21 43 59 75 SI CK 15 31 47 63 79 95 CK 3 13 35 67 83 CK 7 23 39 55 71 87 CK B CK !2 28 44 60 76 92 CK 16 32 46 64 80 96 CK 4 20 36 52 68 84 CK 6 24 40 56 72 86 CK B CK ft 29 45 61 77 93 CK / 17 33 45 65 81 CK 5 21 37 53 69 85 CK 9 25 41 57 73 89 CK B CK H 30 46 62 78 94 CK 2 18 34 50 66 82 CK 6 22 38 54 70 86 CK 10 26 42 58 74 90 CK B CK 15 31 47 63 73 95 CK 3 19 35 5/ 67 83 CK 7 23 39 55 7/ 87 CK II 27 43 59 75 9/ CK B CK 16 32 43 64 00 96 CK 4 20 36 52 66 m CK 8 24 40 56 72 88 CK 12 28 44 60 76 92 CK Figure 1.— Planting Plan of Wheat Variety Tests 1920 and 1921. Legend: B, border. CK, check. Numbers 1-96, planting numbers of varieties tested as given in Tables 4 and 5. Experiments in Field Plot Technic 11 and 32 pounds for oats. The measures of variability used are the average deviation, the standard deviation, and the probable error. These were computed according to the following formulae: n E=±.6745 cr . in which A.D. = average deviation, o-=standard deviation, E = prob- able error (of a single determination), d = the deviation of a single variate from the mean, and n = the number of variates. The correla- tion coefficient r was determined by the formula / S(d x d y ) and the probable error of the correlation coefficient E r by the formula E r .6745 (1 — r 2 ) V" The tests reported are of two kinds, variety tests and strain tests. The variety tests were comparisons of commercial varieties, most of which were taxonomically distinct. A number of pure line selec- tions were included in the wheat variety tests. The strain tests were comparisons of a considerable number of commercial lots of the same variety obtained from different sources. These strains, so-called for convenience, are not, except in a very few cases, pure lines. Some of them are possibly identical, and all the strains of any one variety are of course very similar, since they are taxonomically the same. Procedure. — In the seasons of 1919, 1920, 1921, tests of va- rieties and strains of oats, barley, and wheat were conducted in blocks consisting of five rows ten inches apart and usually 18 feet long. From 24 to 96 varieties were included in each test, and from three to six (usually four) replications were used. The planting order in each case was designed on a plan similar to that illustrated in figure 1. It will be noted that the check plots were in continuous strips, that each variety was represented in each quarter of the field, whether divided from east to west or from north to south, and that in all four series each variety occupied the same position with relation to the check plots, and had the same varieties adjoining it on either side. 12 Missouri Agr. Exp. Sta. Research Bulletin 49 The rows in some cases ran east and west, and in some cases north and south. All these plots were seeded with a 5-row nursery drill, built from plans furnished by Professor T. A. Kiesselbach of the Nebraska Sta- tion. This is a hoe drill designed for rapid and thorough cleaning between plots. Photographs of it have been published in reports of earlier work on field plot technic at the Nebraska Station (Mont- gomery 14 page 57, and Kiesselbach 5 page 16). Its use resulted in uni- form seeding and covering and accurate spacing between rows, with a close approach to ordinary field conditions in the state in which the field was left after seeding. Each field was seeded in a single day. All plots were harvested by hand with sickles, a foot at each end of each row discarded, and the remainder (usually 16 feet) tied in a bundle and hung in a ventilated shed to dry. In 1919 and 1920 each row was bundled and threshed separately; in 1921 the border rows of each 5-row block were bundled separately and the three in- terior rows bundled together. Yields were determined by weighing in grams at the time of threshing. All final yields were converted to bushels per acre and are so expressed. Work of 1919. — In 1919 tests were conducted with barley and oats. Thirty varieties of barley were grown, each in 3 replicate plots. The test comprised three ranges of 185 rows each, including 21 check plots, or one in every sixth plot. The barley was drilled at the rate of eight pecks per acre, on March 21, in rows running north and south. The rows were 14 feet long and 10 inches apart. They were cut to 12 feet in harvesting. The planting plan is shown in figure 2. Conditions CK / 2 3 9 5 cn 6 7 8 9 10 OH II 12 13 19 15 a 16 17 18 19 20 CK 21 22 23 29 25 CK 26 27 28 29 30 CK B CK 2/ 22 23 2i 25 cn 26 21 28 29 30 CH 1 2 3 9 5 cn 6 7 8 9 10 CH 11 12 13 19 15 CH 16 n !8 19 20 CK B CK // 12 /3 M 15 CK 16 17 18 !9 20 CK 2/ 22 23 29 25 CK 26 27 28 29 30 GK / 2 3 9 5 CH 6 7 8 9 to CH B Figure 2. — Planting Plan of Barley Variety Test 1919. Legend: B, border. CK, check. Numbers 1-30, planting numbers of varieties tested, as given in Table 1. were fairly favorable, and the yields of the adapted varieties were slightly higher than the average obtained under the conditions at Co- lumbia. Two varieties, Italian and Australian White, gave extremely low yields and were excluded. Another, Sandrel, was represented only in two series, and was also excluded. The yields of the remain- Experiments in Field Plot Technic 13 in g 27 varieties are shown in Table 1. The planting numbers given in this table correspond to those shown in the diagram of the field (figure 2 .) Table 1. — Yields oe Barley Varieties. In Bushels per Acre. 1919. Planting number Variety Average Yield 3 interior rows 5 rows 1 Hanna 906 12.55 12.57 2 Steigum 907 19.90 19.65 3 Luth 908 23.65 23.40 4 Eagle 913 20.40 20.13 5 Italian 914* 6.70 6.57 6 Servian 915 19.85 19.86 7 Odessa 916 13.75 13.41 8 Lion 923 21.75 22.14 9 Australian White 925* 1.45 1.74 10 Horn 926 21.25 21.54 11 Odessa 927 20.80 19.53 12 Summit 929 23.05 24.03 13 Mariout 932 18.75 18.15 14 Odessa 934 10.30 9.84 15 Peruvian 935 22.25 20.55 16 Trebi 936 30.90 30.96 17 Sandrel 937* 35.90 33.48 18 Oderbrucker 940 23.35 23.79 19 Frankish 953 22.50 22.05 20 Manchuria 956 30.80 30.03 21 Oderbrucker 957 29.45 29.52 22 Manchuria x Champion of Vermont 959 18.30 17.49 23 Luth 972 25.05 26.28 24 Red River 973 27.25 28.14 25 Featherston 1118 28.25 27.00 26 Featherston 1119 25.80 25.83 27 Featherston 1120 34.35 35.49 28 Hanna x Champion of Vermont 1121 13.75 13.92 29 Manchuria 1125 20.35 20.94 30 Malting 1129 17.25 16.44 Mean 22.06 21.95 Forty varieties of oats were compared in 1919, but only 24 of these could be replicated 4 times and the remaining 16 were duplicated. The planting plan was therefore arranged as for 32 varieties, and these 16 varieties grown in two plots each in place of eight varieties ? H i and . A 1 u . stralia ? White 925 were omitted from all computations because of their extremely low yields, and Sandrel 937 because omitted in the third series. 14 Missouri Agr. Exp. Sta. Research Bulletin 49 CK / 2 3 4 5 6 7 3 CK 9 10 n 12 13 14 15 16 CK 17 16 19 20 2/ 22 23 24 c/r 25 26 27 26 89 30 31 32 CH B CK 33 <34 33 36 37 38 39 40 CK / 2 3 4 5 6 7 8 CK 9 /O II 12 13 14- 15 16 CK n 16 19 20 21 22 23 24 CK B CK 17 18 13 20 2/ 22 23 24 CK 25 26 27 28 29 30 31 32 CK I 2 2> 4 5 6 7 8 CK 9 to II 12 13 14 15 16 CK B CK 9 10 II 12 13 /4 15 16 CK 17 18 19 20 2/ 22 23 24 CK 33 34 35 36 37 36 39 40 CK / 2 3 4 5 6 7 6 CK B CK 5 E E D n U L T / P L 1 C 8 T I 0 N CK B CK / 2 3 4 5 CK 6 7 8 9 10 CK II 12 13 /4 15 CK / 2 3 4 5 CK 6 7 8 9 70 CK II 18 13 14 15 CK B CK // 12 13 K 15 CK I 2 3 4 5 CK 6 7 6 9 10 CK II 12 13 /4 15 CK / Z 3 4 5 CK 6 X X 9 10 CK B CK 6 7 8 9 /O CK II 12 13 14 15 CK / Z J 4 5 CK 6 7 8 9 10 CK // 12 13 14 15 CK X Z X X X CK B Figure 3. — Planting Plan oe Oats Variety and Strain Tests 1919. Legend: B, border. CK, check. Numbers 1-40 in first four ranges, planting numbers of oats varieties, as given in Table 2. Numbers 1-15 in last three ranges, planting numbers of oats strains, as given in Table 3. X, test plots planted to check variety because of insufficient supply of seed. Table 2. — Yields of Oats Varieties. In Bushels per Acre. 1919. Planting number Variety Average yield Four series in interior rows Three series 1 A. Sterilis nigra 30.0 31.7 2 Black Mesdag 44.2 44.7 3 C. I. 602 35.4 38.1 4 C. I. 603 53.9 55.1 5 C. I. 620 13.1 14.1 6 Early Champion 55.5 53.9 7 Early Gothland 54.1 52.8 8 Garton 473 30.6 31.7 9 Garton 585 21.7 23.0 10 Golden Giant 42.0 44.9 11 Irish Victor 69.6 70.2 12 Japan Selection 47.9 50.9 13 June 43.1 44.5 14 Kherson Selection 67.2 63.1 15 Fulghum 042 60.9 57.1 16 Lincoln 51.5 50.3 17 Monarch 56.0 53.4 18 North Finnish 51.0 49.5 19 Scottish Chief 59.3 60.1 20 Sparrow bill (Missouri) 39.8 41.3 21 Sparrow bill (Cornell) 42.3 45.7 22 Tobolsk 1 52.6 57.3 23 Tobolsk 2 46.1 51.9 24 White Tartar 49.7 50.3 Mean 46.6 47.3 Experiments in Fieed Plot Technic 15 in four plots each, as shown in figure 3. The rows were 14 feet long and were cut to 12 feet in harvesting. This is a convenient size of plot for oats tests with 10 inches distance between rows, when the border rows are discarded, since the total yield of three rows in grams, divided by 10, gives the yield in bushels per acre. The oats were planted at the rate of 10 pecks per acre, on March 18, in rows running north and south. The season was favorable and a good yield of the better varieties was obtained. The yields of the 24 varieties replicated four times are shown in Table 2. The oats strain test was conducted on the same field, as shown in figure 3, directly south of the oats variety test. In planting, these two tests were handled as one ; and the rate, date, and method of planting were the same. The strains tested were 15 strains of oats obtained under the name Red Rustproof from various experiment stations and seedsmen. Three of these strains, 0121, 0124, and 0127, were not true to name, but the remainder were taxonomically Red Rustproof oats, as described by Etheridge 2 . The oats strains were tested in six series, with check plots in every sixth plot. The line of check plots on the west, however, gave abnormally low yields, probably because they were located partly on a dead furrow at the edge of the experiment field. On account of shortage of seed some of the varieties could not be planted in the last series. The first and last series were therefore dis- Table 3. — Yields of Oats Strains (Red Rustproof). In Bushels per Acre. 1919. Planting number Accession number Average yield 3 interior Rows 5 Rows 1 0119 49.58 49.41 2 0120 45.83 44.51 3 0121* 49.43 53.01 4 0122 47.85 49.59 5 0123 53.55 53.47 6 0125 50.18 49.19 7 0126 44.85 45.81 8 0127* 38.55 36.67 9 0124* 63.90 67.46 10 0133 48.00 46.49 11 0128 53.55- 53.15 12 0129 49.35 49.01 13 0130 52.73 51.89 14 0131 48.60 47.84 15 0132 55.13 55.44 Mean 50.07 50.20 # Not taxonomically Red Rustproof. 16 Missouri Agr. Exp. Sta. Research Bulletin 49 carded. The average yields of the 15 strains in the four remaining series are shown in Table 3. Work of 1920. — Wheat varieties were grown in 5-row blocks in 1919-20. Ninety-six varieties were included in the test, four replica- tions being used. Fultz wheat was grown as a check in every sixth plot. The rows were 18 feet long and were cut to 16 feet in harvest- ing. The direction of the rows was east and west. The planting plan is shown in figure 1. The wheat was sown October 15, at the rate of 6 pecks per acre. There was considerable winter injury in the plots and the condition of the wheat in early spring was rather poor. The yields obtained are shown in Table 4. Table 4. — Yields oe Wheat Varieties. In Bushels per Acre 1980. Average yield - Average yield Planting 3 Interior 5 Planting 3 Interior 5 number Variety Rows Rows number Variety Rows Rows 1 Beechwood Hybrid No. 12.. 10.8 11.1 50 Michigan Wonder No. 141 . . 10.7 10.3 2 Beechwood Hybrid No. 81.. 12.8 14.1 51 Michigan Wonder No. 155 . 10.1 9.8 3 Beechwood Hybrid No. 85.. 12.5 12.7 52 Michigan Wonder No. 209 . 12.1 12.5 4 Beechwood Hybrid No. 87.. 14.2 13.7 53 Michigan Wonder No. 211 . 9.9 9.9 5 Beechwood Hybrid No. 202. 11.9 12.5 54 Michigan Wonder No. 221 . 11.0 11.1 6 Beechwood Hybrid No. 207. 13.2 13.6 55 New York 123-32 . . . 17.2 17.5 7 C. I. 3808 16.2 16.6 56 N iagara . 13.8 13.5 8 C. I. 3846 14.2 15.6 57 Nigger . 11.8 11.8 9 C. I. 3972 14.7 15.6 58 Old Ironclad . . . . 12.5 13.2 10 C. I. 3980 16.4 17.5 59 Poole . 10.5 10.7 11 C. I. 3988 16.7 17.0 60 Poole No. 3 . . . . . 11.7 11.0 12 C. I. 4004 14.3 14.0 61 Poole B-3 . 12.5 13.3 13 Common Rye 17.3 18.5 62 Portage . 15.9 17.3 14 Dawson’s Golden Chaff .... 13.0 12.3 63 Pride of Indiana . 14.2 14.4 15 Deitz 15.1 14.6 64 Pride of Genessee . . . 15.7 18.1 16 Early Ripe 12.2 12.6 65 Reliable . 12.6 12.9 17 Early Ripe No. 26 13.2 14.0 66 Red Cross . 13.1 13.1 18 Early Red Clawson 9.9 9.5 67 Red May . 14.8 14.8 19 Farmer’s Friend 18.8 19.9 68 Red Rock (Indiana) . 18.7 19.7 20 Fulcaster 14.4 15.3 69 Red Rock (Michigan) . 7.5 6.8 21 Fultz GArchias) 12.2 13.0 70 Red Wave . 12.9 12.7 22 Gold Coin 11.7 12.2 71 Rochester Red . . . 12.7 12.9 23 Greene County 15.6 15.1 72 Rosen Rye . 20.7 24.0 24 Harvest King No. 7 13.4 14.2 73 S. P. I. 11616 . . 10.3 10.9 25 Harvest Queen 9.6 9.8 74 S. P. I. 26012 . . 13.4 12.9 26 Hickman T. 8 8(2 75 S. P. I. 26013 . 15.2 15.5 27 Illini Chief 17.5 18.7 76 S. P. I. 26014 . . 17.5 18.8 28 Jones Climax 19.1 20.7 77 S. P. I, 26015 . . 13.2 13.4 29 Kanred 21.0 22.7 78 S. P. I. 26017 . . 13.4 13.5 30 Kessinger 18.0 19.3 79 S. P. I. 26018 . . 13.6 13.6 31 Kharkov 18.9 20.1 80 S. P. I. 26019 . . 11.6 11.6 32 Leap’s Prolific 14.2 14.8 81 S. P. I. 26022 . 10.6 10.1 33 Mediterranean No. 8 9.1 9.5 82 S. P. I. 26023 . . 9.1 8.5 34 Michigan Amber 10.7 11.3 83 S. P. I. 26025 . . 12.3 13.1 35 Michigan Amber (Indiana) 17.0 17.9 84 S. P. I. 26029 . . 15.4 15.6 36 Michigan Amber No. 7 ... 10.5 10.8 85 S. P. I. 26085 . . 13.2 13.3 37 Michigan Amber No. 12 ... 9.3 9.3 86 Treadwell . 12.7 12.8 38 Michigan Wonder 10.9 11.1 87 Valley . 12.4 12.1 39 Michigan Wonder No. 4 ... 12.4 12.7 88 Velvet Chaff No. 2 . . 14.1 12.8 40 Michigan Wonder No. 8 ... 10.8 11.0 89 Velvet Chaff No. 8 .. . 9.0 9.3 41 Michigan Wonder No. 21 . . 8.2 8.7 90 Ziegler’s Fly Proof . . 10.9 11.7 42 Michigan Wonder No. 53 . . 8.5 9.6 91 13D-4a . 14.1 13.9 43 Michigan Wonder No. 54 . . 11.3 10.7 92 37a-4 . 14.6 14.7 44 Michigan Wonder No. 83 . . 13.6 13.7 93 Fulcaster (Co-op) . 17.4 18.5 45 Michigan Wonder No. 96 . . 9.7 9.7 94 Fultz (Co-op) . 15.2 15.9 46 Michigan Wonder No. 103 . 9.7 9.1 95 Kanred (Co-op) . . 19.1 20.6 47 Michigan Wonder No. 116 . 16.4 15.8 96 Poole (Co-op) . . . 19.4 21.0 48 Michigan Wonder No. 130 . 14.3 14.2 49 Michigan Wonder No. 140 . 12.3 12.7 Mean 13.4 13.8 Experiments in Fieed Plot Technic 17 * m O 49 o 2 .E^ES bo 2 g > M O i. to 3 2 o.2 jti ~ 09 (O .si H < . w to O <.2 u a E J2 3 PTf^fOfOOfs.rHCgoOoqOx^Ttov6M;Tf(^CT\ONOOO\iorOTt-CS|^-;CT\VOCNOO\OVOini-t^OOin ^ ^ '^ T t vo . t t ^^.°; M .' £ ; 0 \Tl;iO 00 00 (oO0 2 2 2 2 2 2 2 2 22 2 2 2 2 2 21 21* 12 12 2 2 2 2 22 2i 2l 22 2 22 21 2 2 21 ^ ™ ^ ^ ^ ^ w ^ ^ ^ ^ ^ u^\OTfcov£)^^rovOO\0\OfOrOTl ; ro<>jqvOTj-v£)OOOOl^roONt^voOs l ^'OONinOxOOSTfONOO t ' s ‘ vo ' t '' 0 ° o ' i S O O ,-t to On h I-H (O ^ Tj- U 1 O rH CSI rtrt rt rtOlMC0 0 O O O O O O uoSoSSoJo 'a'O'a'ri’O'O'a' § § § § § § §s G 3 GSGGGO cUrtwrtrtrtrtKj — bo bo bo bo bo bo bo 1 '* J- £ 15 15 IE IS 15 ISIS k, bo J2 .2 „ „ _ 09 09 C9 09 09 O 09 > bOT3 o o o o o o.O o O £! 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L ^ 51 ^' 2 O 3 3 3 ^ to rt rt a3 u bo ho bo bo 2 IS IS 15 IS * u. 2 . 2 . 2.2 ro VO I'tOVOOH iioooovhh 0000000 !"C' 2 r T 3 nJ F dT 3 n 3 r O' :G 5 CCCt:! cc 3 'OOOOOO OO 3 3 aJ rt bo bo ISIS 09 09 333^3333 rtajrtrtcSrtcflctJ bo bo be bo bo bo bo bo It) ve N CO Ov O n ro ro ro to tf ^ 18 Missouri Agr. Exp. Sta. Research Bulletin 49 Work of 1921. — In 1920-21 ninety-six varieties of wheat were again tested by this method. Many of the varieties were the same as those tested in the preceding year, about 20 varieties being eliminated and a corresponding number added. The planting plan was the same as that of the preceding season. Poole wheat was used as a check variety. The wheat was drilled at the rate of 5 pecks per acre, October 6, in rows running east and west. The season was favorable, but yields were reduced by the very rapid ripening of the wheat caused by the hot dry weather in the second and third weeks of June. The yields are shown in Table 5. Table 6. — Yields of Wheat Varieties and Mixtures In Bushels per Acre. 1921. Planting number Variety Average yield 3 Interior Rows 5 Rows 1 Fulcaster 17.3 18.6 2 Harvest Queen 14.2 14.5 3 Mixture No. 1 (1, 2, 4, 5) 15.9 16.2 4 Michigan Wonder 16.8 17.8 5 Nigger 10.8 10.8 6 Michigan Wonder No. 21 19.8 20.8 7 Michigan Wonder No. 54 18.9 19.3 8 Mixture No. 2 (6, 7, 9, 10) 20.8 21.3 9 Michigan Wonder No. 96 18.5 18.9 10 Michigan Wonder No. 209 21.7 22.6 11 Beechwood Hybrid No. 12 17.4 18.8 12 Beechwood Hybrid No. 85 16.5 17.3 13 Mixture No. 3 (11, 12, 14, 15) 17.6 18.4 14 Beechwood Hybrid No. 87 19.9 19.9 15 Beechwood Hybrid No. 207 17.4 17.9 16 Michigan Wonder No. 221 18.6 20.3 17 Kanred 13.6 13.8 18 Mixture No. 4 (16, 17, 19, 20) 17.8 18.0 19 New York 123-32 19.6 19.7 20 Red Rock 17.6 17.4 21 Red Hussar 16.3 17.8 22 Turkey (Kansas) 10.8 10.5 23 Mixture No. 5 (21, 22, 24, 25) 15.7 15.9 24 Michigan Amber 19.2 19.6 25 Nigger 14.1 13.4 26 Fulcaster (Co-op) 20.4 21.2 27 Fulcaster (Outl) 20.1 20.6 28 Mixture No. 6 (26, 27, 29, 30) 20.1 21.0 29 Fulcaster (Blazier) 20.6 21.5 30 Fulcaster (Cowles) 20.6 20.6 Mean 17.6 18.2 Experiments in Field Plot Technic 19 On another field in 1921, a test of mixtures of varieties and strains of wheat in comparison with their pure constituents was con- ducted. Each mixture was made up of four varieties or strains, in equal quantities of seed by weight. The composition of the mixtures and the yields obtained are shown in Table 6. The planting plan is shown in figure 4. This wheat was drilled at the rate of 5 pecks per B Cli 2 3 4 5 CK 26 27 26 29 30 CK 16 n Id 19 20 CK II 12 13 14 IS CH B B CH 6 7 a 9 10 CK 1 2 3 4- S CK 21 22 23 24 25 CK 16 17 18 19 20 CH B B CK // IZ 13 1+ 15 CK 6 r a 9 /O CK 26 27 28 29 30 CK 2 1 32 23 24 25 CH B B CK 16 17 16 19 20 CK // id 13 14 15 CK 2 3 4 S CK 36 37 28 29 30 CK B B CK 2 1 22 23 24 25 CK /6 17 18 19 20 CK 6 7 8 9 10 CK Z 3 4 S CK B B CK, 26 27 26 29 30 CK 2/ 22 23 24 25 CK II /2 13 14 15 CK 6 7 8 9 10 CK B Figure 4. — Planting Plan of Wheat Mixture Test 1921. Legend: B, border. CK, check. Numbers 1-30, planting numbers of varieties and mixtures tested, as given in Table 6. acre in rows running north and south, on October 8, 1920. This test will be referred to as the wheat mixture test. In 1921 tests of oats varieties and of oats strains were also con- ducted in 5-row blocks. Thirty-two strains of Red Rustproof, in- cluding many of those tested in 1919 and a number of others, and 32 strains of Kherson oats, obtained in the same way, were included in the oats strain test. The Kherson and Red Rustproof strains were arranged alternately, and both Kherson and Red Rustproof checks were grown, as shown in figure 5. The test of these 64 strains, in four series, occupied 16 ranges. The next eight ranges on the same plot were used for an oats variety test in which 32 varieties of oats were compared, each in four replicate plots. In this part of the field the Kherson and Red Rustproof check plots were continued. There are thus available the yields of 120 plots each of Kherson and Red Rustproof oats, or five strips of 24 plots of each arranged in pairs side by side. In both of these experiments the rows ran east and west, and were 18 feet long, cut to 16 feet in harvesting. The oats were drilled on March 12, at the rate of 10 pecks per acre. The yields of oats, particularly of the later-maturing varieties, were materially re- duced by the hot dry weather in the middle of June. The yields of the oats varieties are shown in Table 7, and those of the strains in Table 8. 20 Missouri Agr. Exp. Sta. Research Bulletin 49 B 74 R / 17 33 49 K 74 13 29 45 6/ K R 9 25 41 57 R R 5 21 37 53 K R 3 B 74 R 2 18 3.* 50 74 R 14 30 46 62 74 R 10 26 42 S7 74 R 6 22 38 54 74 R B B n R 3 19 35 5/ 74 R 15 3/ 47 63 74 R // 27 43 59 R R 7 23 39 55 74 R B B K R f ZO 36 52 K R 76 52 48 i Cb 1 K R 72 28 44 60 K R 8 24 40 56 K R B B 74 R 5 2! 37 53 74 R / 17 33 49 74 74 73 29 45 61 74 R 9 25 41 57 R R B B 74 R 6 2Z 38 54 K R 2 te 54 SO 74 74 J4 30 46 62 n R / 0 26 42 58 K R 3 B K R 7 23 39 55 K R 3 79 35 51 74 R 15 3/ 47 63 K R // 27 43 59 K R B B 74 R 8 2* 40 56 74 R 4 20 36 52 74 R 16 32 48 64 74 R J2 26 44 60 74 R B B /r R 9 25 41 57 74 R 5 2/ 37 53 74 R / 17 33 49 74 R 13 29 45 6/ 74 R 3 B /r R /O 26 42 58 K R 6 ZZ 38 54 n R 2 is 34 50 74 R 14 30 46 62 74 R 3 B K R n 27 43 59 K R 7 23 39 55 74 R 3 19 35 5/ K R 15 3/ 47 63 K R B B 74 R 12 28 44 60 K R 8 24 40 56 n R 4 20 36 52 74 R 16 32 48 64 74 R 3 B 74 R /3 29 45 61 74 R 9 25 41 57 74 R 5 2! 37 53 74 R / 77 33 49 K /? B B A R M- 30 46 62 74 R 10 26 42 58 74 R 6 21 38 54 74 R 2 18 34 so 74 R 3 B 74 R 15 3/ 47 63 74 R // 27 43 59 74 R 7 23 39 55 74 R 3 19 35 5/ 74 74 B B 74 R 16 32 48 64 R R 12 22 41 60 K R 8 Z4 40 56 K R 4 20 36 52 74 R B B 74 R 65 73 8J 89 K R 77 73 87 95 K R 69 77 85 93 74 R 67 75 83 91 K R B B n R 66 74 82 90 H R 72 80 38 96 K R 70 76 86 94 n R 68 76 84 92 n R B B 74 R 67 75 83 SI K R 65 73 81 89 74 R 7/ 79 87 95 14 R 69 77 85 93 n R B B 74 R 68 76 84 32 74 R 66 14 82 30 R R 72 80 88 96 K R 70 IQ 86 94 74 R B B n R 69 77 85 93 74 R 67 75 83 9 ! n n 65 73 81 89 K R 7/ 79 67 95 n R 3 B 74 R 70 78 86 34 K R 68 76 84 92 n R 66 74 82 90 74 R 72 80 68 96 K R B B K R 7/ 79 87 95 ff R 69 77 85 93 n R 67 75 83 5/ n R 65 73 81 89 n R B B 74 R 72 80 88 96 H R 70 78 66 94 n R 68 76 84 32 74 R 66 74 82 90 71 R 3 Figure 5. — Planting Plan of Oats Variety and Strain Tests 1921. Legend : B, border. K, Kherson check. R, Red Rustproof check. Numbers 1-64, planting numbers of oats strains, as given in Table 8. Numbers 65-96, planting numbers of oats varieties, as given in Table 7. Experiments in Field Plot Technic 21 Table 7 —Yields oe Oats Varieties. In Bushels per Acre. 1921. Planting number Variety Average yield 3 Interior Rows 5 Rows 65 Burt 49.13 51.94 66 Canadian 25.31 25.13 67 C. I. 603 22.50 23.06 68 Culberson 24.75 25.13 69 Danish Island 19.69 19.13 70 Early Dakota 21.56 21.56 71 Early Gothland 23.44 22.13 72 Garton 748 21.00 20.81 73 Green Russian 26.06 26.25 74 Irish Victor 29.81 32.06 75 Joanette 19.31 19.69 76 Fulghum 042 45.19 47.44 77 Monarch 29.63 31.88 78 Monarch Selection 35.63 36.38 79 Scottish Chief 26.63 27.38 80 Silvermine 050 31.69 32.06 81 Silvermine Selection 22.13 24.94 82 Sparrowbill (C) 15.38 14.63 83 Sterilis Selection 38.63 36.94 84 Storm King 20.06 17.63 85 Swedish Select 057 21.00 19.50 86 Fulghum 065 42.00 44.81 87 Fulghum 0113 42.00 45.38 88 Silvermine 0115 25.13 24.94 89 Silvermine 0117 21.75 22.69 90 Fulghum 0124 45.38 48.38 91 Fulghum 0145 39.19 41.81 92 Fulghum 0149 42.75 47.06 93 Fulghum 0151 39.75 43.88 94 Fulghum 0152 39.75 42.38 95 Silvermine 0165 28.31 26.81 96 Swedish Select 0165 20.81 18.56 Mean 29.85 30.70 22 Missouri Agr. Exp. Sta. Research Bulletin 49 Table 8. — Yields of Oats Strains (Red Rustproof and Kherson). In Bushels per Acre. 1921. Red Rustproof strains Average yields Planting 3 Interior 5 number Strain Rows Rows 1 066 24.00 23.25 3 067 24.00 21.75 5 068 23.25 23.44 7 069 19.31 18.00 9 072 18.38 18.75 11 074 22.31 20.63 13 075 24.19 22.13 15 0118 16.50 16.31 18 0119 22.31 21.38 20 0120 21.19 19.69 22 0122 19.13 17.81 24 0125 21.00 19.88 26 0126 25.31 22.50 28 0128 20.44 20.25 30 0129 21.94 21.56 32 0130 21.75 20.25 33 0131 24.56 23.25 35 0132 17.63 19.13 37 0133 18.94 18.75 39 0134 16.50 15.75 41 0135 17.63 15.65 43 0136* 32.44 33.19 45 0141 21.94 21.19 47 0163 13.88 12.94 50 0169 15.38 14.44 52 0181 19.88 18.00 54 0182 19.88 19.13 56 0183* 41.44 43.31 58 0383 23.63 24.00 60 0391 29.44 30.19 62 0394 22.31 21.56 64 0395 23.25 21.94 Mean 21.00 20.12 ♦Not taxonomically Red Rustproof. Ex- cluded from average. Kherson strains Average yields Planting 3 Interior 5 Number Strain Rows Rows 2 023 35.25 36.38 4 040 36.57 37.50 6 041 36.56 38.81 8 052 38.06 38.81 10 053 39.75 42.00 12 079 32.63 34.88 14 080 35.44 38.25 16 082 40.88 41.44 17 083 35.44 38.25 19 085 38.25 41.81 21 086 36.75 37.69 23 Mixture** 33.75 36.38 25 088*** 27.00 27.56 27 089 30.94 31.69 29 090 36.38 38.06 31 091 30.19 31.88 34 094 31.69 33.56 36 095 38.81 39.75 38 096 36.38 38.06 40 097 31.31 32.25 42 098 38.63 39.19 44 099 38.81 38.25 46 0100 40.13 42.75 48 0155 37.50 38.63 49 0157 43.69 45.00 51 0158 34.69 35.25 53 0159 33.38 33.94 55 0160 30.19 31.13 57 0161 34.69 36.00 59 0162 25.31 25.69 61 0167 40.69 39.94 63 0174 36.75 38.25 Mean 35.79 37.14 ♦♦Mixture of strains 082, 094, 0100, 0174. ♦♦♦Not taxonomically Kherson. Ex- eluded from average. Experiments in Field Plot Technic 23 COMPETITION AS A SOURCE OF ERROR IN PRELIMINARY TESTS. Previous Investigation. — The possibility of error from competi- tion in single-row tests was noted by Montgomery 14 in 1913, in the following passage : “In 1908 it was observed that a certain strain of early wheat in a series of row plats made a very poor appearance at harvest time, while the same strain planted in centgeners made a much better comparative showing. Ap- parently the larger and faster growing strains on each side, the rows being only 8 inches apart, exercised some competitive effect. This effect of com- petition has been noted for two years since. Also in certain variety tests of oats, grown in row plats 10 inches apart, the same effect was noted. Exact data cannot be given on this point, as the results from the series of plats planted in 1909 and in 1910 for this purpose were seriously impaired by unfavorable conditions; but Table XVIII, giving results from adjacent row plats sown at different rates, shows that the 800-seed rate made a marked increase over the 700-seed rate, while in a similar series of blocks (Table XIX), sown at the same rate, this marked increase was not noted. Since the 800-seed row was always adjacent to the 400-seed row, it may have had some advantage on this account. Danger from this source can probably be avoided if care is taken to plant only similar varieties in adjacent rows. Where the block plat is used this source of error is eliminated.” Hayes & Arny 4 found considerable competition between rod-rows grown one foot apart. Three-row plots were used in variety tests of winter wheat, spring wheat, barley, and oats, and the yields of each row determined separately, in 1916. The comparative yield of the border rows in each plot was then correlated with the comparative height and yield of the adjacent rows. There was some effect on the yield of border rows due to the height of adjacent rows in the case of barley and winter wheat. The results were variable in different plots. In the case of oats the effect of height was rather obscure, and in the case of spring wheat it was not apparent. The yield of adjacent rows appeared to be of some importance in the barley tests and in some of the spring wheat tests. These results led to the adoption of 3-row plots with discarded border rows for preliminary testing at the Minnesota Station. Love and Craig* in describing the methods used in cereal investiga- tions at the Cornell Station describe the single-row test and add : “In order to prevent any effect which may be caused by two unlike sorts growing together the different strains are arranged according to earliness and other characters so as to reduce this source of error to a minimum.” 24 Missouri Agr. Exp. Sta. Research Bulletin 49 Kiesselbach 5 ’ 7 has published rather extensive data on the compe- tition between adjacent rod-rows. In his experiments the crops were compared in alternating single-row plots and in alternating 5-row blocks, each replicated fifty times. In some cases the border rows of 5-row plots were discarded. The deviation of the result in the test in single-row plots from that of the test in 5-row plots is regarded as the measure of the effect of competition. The comparative yields of va- rieties of wheat and oats in alternating single rows and in alternating 5-row plots are shown in Table 9, from Kiesselbach 7 . Tabu; 9. — Relative Yields of Two Small Grain Varieties When Compared in Alternating Rows and in Alternating 5-Row Plats (Kiesselbach). Wheat Oats Average yield of l 50 plats Average yield of 50 plats Year and Alternating Alternating Year and Alternating Alternating variety single rows 5-row blocks* Variety single rows 5-row blocks’ 1 % % % % 1913 1913 Turkey 100 100 Kherson 100 100 Big Frame 107 97 Burt 130 112 1914 1914 Turkey 100 100 Kherson 100 100 Big Frame 85 97 Burt 139 101 1913 1913 Turkey 100 100 Kherson 100 100 Neb. No. 107 107 Swedish 82 77 28 Select 1914 1914 Turkey 100 100 Kherson 100 100 Neb. No. 63 85 Swedish 89 93 28 Select ♦Yield based on 3 inner rows of 5-row plats in 1914. Kiesselbach also submits interesting data on the competition of pure line selections of the same variety. It might be supposed that such strains, being similar in varietal characteristics, would be little affected by competition, and could therefore be safely compared in single-row plots. The average relative yields of three strains of Turkey wheat in single rows and in blocks for two seasons, however, showed that the two better strains were favored approximately 20 per cent and 15 per cent, respectively, at the expense of the poorer strain, in the single-row test. A strain which yielded 26 per cent more than an- other in the single-row test yielded only 6 per cent more in the block Experiments in Field Plot Technic 25 test. Kiesselbach has therefore adopted the practice of testing such strains in 5-row blocks replicated ten times instead of in single-row plots. Love 8 has criticized these results because in some cases at least the rows ran east and west rather than north and south. He states that in experiments at Ithaca, New York, there is little competition be- tween varieties grown in single rows, when the rows run north and south. “In order to obviate any criticism of this method,” he adds, “it might be well to follow the plan of arranging varieties so that late sorts are grown together and the earlier ones together. In other words, the different sorts could be so arranged that they grade into one another as regards yield, earliness, and the like.” To this Kiesselbach 8 replies that in some of his competition studies the rows ran north and south and in others east and west, and that striking competition occurred in both cases. He adds that although error resulting from row compe- tition would undoubtedly be reduced by grouping varieties of sim- ilar growth habits together, it appears that varieties fairly similar in growth habit may vary for some reason in relative competitive qual- ity. Experimental Results. — Some further evidence on competition as a source of error in plot experiments is afforded by a study of the relative yields of border rows and interior rows in the 5-row blocks used in these preliminary tests. It should be remembered, of course, that the effect on yield would be decidedly greater in single rows exposed to com- petition on both sides than in these border rows, which compete with another sort on only one side. The extent of the error from compe- tition in such border rows is of interest in determining whether it is necessary to discard the border rows of small blocks. When 5-row blocks are used, even if the border rows are not discarded, the relative effect of competition is greatly reduced, since only two of the five rows are subject to varietal competition and these are exposed only on one side. If this results in reducing the error from competition to a low point, or if varieties can be so arranged as to give this result, it may be advisable in practice to harvest 5-row blocks entire, thus avoiding the principal objection to the use of border rows — the loss of a considerable portion of the experimental area. Competition is particularly important as a source of error because of the fact that it tends to affect replicate plots similarly, and conse- quently does not necessarily increase plot variability. For this reason it is likely to escape detection, and, when it is involved in an experi- ment, its effect cannot be measured. There is no 'great objection to a considerable experimental error from plot variability in field experi- 26 Missouri Agr. Exp. Sta. Research Bulletin 49 ments, if the experimenter determines the extent of the error and draws his conclusions accordingly. But a preliminary variety test in which error from competition is not controlled may be very nearly worthless as an indication of the relative value of varieties for field conditions, because actually the relative values of the varieties tested may frequent- ly differ by 50 or 100 per cent from the values determined in the test, without the slightest indication in the experimental results. Illustrations of Effects of Competition . — The error from competi- tion may be illustrated by numerous examples from each of the eight tests here reported. An extreme case is the effect of competition on the relative yield of wheat and rye. Two varieties of rye, common rye and Rosen rye, were included in the wheat variety test, for com- parison with wheat. The average yields of Rosen rye and of the va- rieties of wheat adjoining it on either side, in interior rows and com- peting border rows of the four series, were as follows: Season Variety Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative i [Niagara (Wheat) 13.8 67 10.0 33 1 f30.4 100 1920< [ Rosen (Rye) 20.7 100 \ — [27.7 100 Velvet Chaff No. 2 (Wheat 14.1 68 9.8 35 1 [Red Hussar (Wheat) 14.3 80 11.6 59 1 [19.7 100 1921-j Rosen (Rye) 17.9 100 \ — [25.4 100 Poole (ck) (Wheat) 1 11,8 66 11.2 44 The disturbance of the true comparative value of the varieties by competition may be determined by comparing their relative yields in interior rows and in border rows. Thus Niagara wheat in 1920 yielded 67 per cent as much as Rosen rye in plots protected from com- petition, but only 33 per cent as much in rows not protected from competition. Similarly the yield of Velvet Chaff No. 2 wheat was re- duced from 68 per cent to 35 per cent by competition with Rosen rye. In the following season the reduction in yield of the two varieties of wheat adjoining Rosen rye (Red Hussar and Poole) was not so great, but was still decidedly significant. This clear case of compe- Experiments in Eield Plot Technic 27 tition serves to illustrate the phenomenen, although the competition between wheat and rye has little significance in itself as regards va- riety tests in general, since wheat and rye are not commonly included in the same test. Ordinarily the competition between varieties of the same crop is not so extreme. There are, however, a number of cases in which a variety of wheat or oats profited almost as extremely in competition with other varieties of the same crop as did the rye in competition with wheat in the cases cited above. The wheat variety, Michigan Wonder No. 116, which grew between two other wheat varieties, Leap's Prolific and Poole Selection, in 1921, gave the following results, as an average of the four series: Variety Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative Leap’s Prolific 14.9 91 9.9 53 f 18.8 100 Michigan Wonder No. 116 16.4 100 \ — 1 [21.7 100 Poole Selection 15.3 1 93 j 11.5 53 The effect of competition in this case is almost as pronounced as in the case of the rye, although the three wheat varieties concerned, when protected from competition, gave almost equal yields and differed little in date of heading, date of maturity, and height. In this case a small difference in actual value between the varieties, as indicated by their yields when protected from competition, is greatly increased when their yields in adjacent single rows are compared. A striking case of competition in the oats variety test of 1921 was that of the three varieties Sterilis Selection, Fulghum, and Kherson, the check variety. Their average yields were as follows : Variety Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative Sterilis Selection 38.63 99 28.50 58 f 48.75 100 Fulghum 39.19 100 \ — [ 42.94 100 Kherson (check) 40.69 104 34.18 70 28 Missouri Agr. Exp. Sta. Research Bulletin 49 These three varieties, which gave almost equal yields in rows protected from competition, differed decidedly in their yields in ad- jacent rows. Although Kherson outyielded Fulghum 4 per cent in plots protected from competition, its yield was 30 per cent less than that of Fulghum in single rows not protected from competition. Extreme effects of competition were shown in very numerous cases in the tests of Kherson and Red Rustproof strains in 1921. An example from this plot is the following: Strain Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative 0169 (Red Rustproof check) 18.38 77 22.31 119 1 [18.75 100 067 (Red Rustproof) 24.00 100 ! 1 — : 1 [17.44 100 085 (Kherson) 38.25 159 i 51.94 298 The extreme advantage of the Kherson strain in competition with the Red Rustproof, increasing its margin of superiority from 59 per cent to 198 per cent, is particularly striking. Probably even more significant is the effect of competition between the two Red Rustproof strains, resulting in the conversion of a 23 per cent loss to a 19 per cent gain. All of the cases cited above are taken from plots in which the rows ran east and west. Some examples of varietal competition from tests in rows running north and south are the following: In the barley variety test, the variety Featherston 1118 occurred between Red River 973 and Oderbrucker (the check variety). The average yields of these three varieties in the three series were as follows : Variety Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative Red River 973 27.25 96 31.20 127 f24.60 100 Featherston 1118 28.25 100 j — (25.65 100 Oderbrucker (ck) 34.87 123 42.19 164 Experiments in Field Plot Technic 29 In this case the advantage of Oderbrucker over Featherston was almost tripled by competition, and Red River, which yielded less than Featherston in the interior rows, excelled it materially in yield in the border rows. The oats varieties tested in rows running north and south in 1919 showed marked effects of competition in several cases. The follow- ing will serve as an example: Variety Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative Kherson Selection 61.3 111 84.7 [49.4 171 100 Fulghum 042 57.1 100 [50.7 100 Lincoln 50.3 88 57.5 113 In this case Lincoln, yielding 12 per cent less than Fulghum in in- terior rows, yielded 13 per cent more than Fulghum in border rows; while the advantage of Kherson Selection over Fulghum was in- creased from 11 per cent to 71 per cent. Marked competition is hardly to be expected in the oats strain test of 1919, regardless of the direction of the rows, because of the sim- ilarity of the strains in varietal characters. Three strains which proved to be taxonomically unlike Red Rustproof were included in this test, and each of these shows clearly the effects of competition. For ex- ample, strain 0124, which was classified as Fulghum, gave the follow- ing yields in comparison with the adjoining strains, 0127, classified as Kherson, and 0133, classified as Red Rustproof : Strain Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative 0127 (Kherson) 38.55 60 32.40 48 f66.83 100 0124 (Fulghum) C3.90 300 — [78.75 100 0133 (Red Rustproof) 48.00 75 43.20 55 30 Missouri Agr. Exp. Sta. Research Bulletin 49 Moreover, the Red Rustproof strains showed competitive effects among themselves to some extent, though not so conspicuously as dif- ferent varieties. For example the strains 0122 and 0123, which were taxonomically identical, yielded as follows: Strain Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative 0122 (Red Rustproof) 0123 (Red Rustproof) 47.85 53.55 89 10O 1 55.13 49.28 112 100 Strain 0122 which was apparently 11 per cent inferior to strain 0123 in the yields of interior rows, appeared to be 12 per cent superior to the same strain in the yields of their adjacent border rows. In the wheat mixture test of 1920 also the rows ran north and south. An example of competition from this test is the following: Variety Yield in interior rows Yield in competing border rows Bushels Relative Bushels Relative Poole (check) 15.1 81 14.5 60 f24.2 100 Michigan Wonder No. 221 18.6 100 \ — [21.4 100 Kanred 13.6 73 1 12.5 58 In this case also differences in yield were increased by compe- tition. The individual cases cited above will serve to show the existence of competition as a source of error in these tests. As a result of competition the differences between varieties may be increased or de- creased, and in some cases a material advantage in yield may be con- verted to a material disadvantage. The phenomenon occurs, under conditions at Columbia, whether the rows run north and south or east and west. Of course it is not true that all of the difference in yield between border rows and interior rows is necessarily caused by varietal competition. Some variation in the yield of adjacent rod- rows will occur regardless of competition. When the means of only Experiments in Field Plot Technic 31 four determinations are compared the effect of this variability may be considerable. If a field uniformly seeded to a single strain were har- vested in rod-rows and assumed to be made up of several different varieties each in four distributed plots, doubtless the average border yield would differ materially from the average interior yield in several “varieties.” It is not however, likely, that such differences as those cited above would be caused by chance variability. Nevertheless, no final conclusions regarding competition as a source of error should be drawn from such individual cases. The extent of error from compe- tition is better shown in the average differences between border yields and interior yields, and in the mean coefficients of competition for complete tests. They are given in the next section. Relation of Competition to Various Characteristics of the Com- peting Varieties. — It is essential that competition be eliminated by the use of border rows, or counteracted by some such means as grouping varieties. The latter is decidedly the preferable method, from the standpoint of economy, if satisfactory results may be obtained by its use. But competition cannot be effectively controlled by grouping varieties unless there is a close correlation between competitive value and some character like earliness or height, which may be known in advance. Determinations of the correlation between competitive ef- fects and various characteristics of the varieties have therefore been made for each of the tests. The preliminary determinations were made as follows : (1) The average yield in interior rows and the average yield in the border rows on each side for all replicate plots of each variety or strain was determined. The replicate plots thus averaged were grown between the same varieties in each series, and it may be assumed therefore that their border rows were subject to the same competition. In the following discussion of competition each individual case repre- sents the mean of all the replicate plots of the test in question. For example, when it is stated that the correlation between competition and yield is determined in a test in which one hundred cases of compe- tition are involved, each of the hundred cases represents the mean of three or four determinations in replicate plots. In most cases the number of replicate plots was four. In the barley test of 1919 only three series were grown, and in the oats variety test of 1919, though four series were grown, only three could be used because one border row of each variety in the first series was harvested for seed and laboratory material. (2) Corresponding average yields were determined for check plots, those adjoining the same variety being averaged together. For 32 Missouri Agr. Exp. Sta. Research Bulletin 49 example, in the wheat variety test diagrammed in figure 1 the four check plots which adjoined variety 1 (one in each series) were aver- aged together, the four adjoining variety 2, the four adjoining variety 3, etc. The four check plots adjoining varieties 89, 90, 91, etc. were similarly averaged. (3) The average yield of each border row for each variety was converted to the percentage of the average yield of the same variety in its interior rows. These yields of border rows in percentage will be referred to as “relative border yields.” The relative border yield gives a rough indication of the effect of competition on the variety. When it is above 100, the variety yielded more in border rows (subject to competition) that in interior rows (protected from competition). When it is below 100, the border yield was less than the interior yield, in proportion. (4) An approximate measure of the competition between each pair of adjacent varieties was obtained by dividing the higher relative border yield by the lower, in the case of their adjacent border rows, and substracting 100 from the result. When the variety on the left has a higher relative border yield, this is given a positive sign; in the reverse case a negative sign. This figure is simply the predominance of the more strongly competing variety over the other in percentage of relative border yield. It will be referred to, for convenience, as the coefficient of competition. (5) This measure of competition was correlated with various characteristics of the competing varieties, including the relative yields in interior rows, the relative grain-straw ratios, the relative dates of heading and of maturity, and the relative heights. In correlating com- petition with the relative yield of the interior rows, the relative yield was determined by dividing the higher yield by the lower, subtracting 100, and assigning a positive or negative sign, as before. The correla- tion determined, therefore, is the correlation between the percentage advantage of one variety over another in competition, and the differ- ence in yield of the two varieties, expressed in percentage, when pro- tected from competition. Relative grain-straw ratios were determined similarly, the ratios being first obtained by dividing the yield of straw by the yield of grain. Relative dates of heading and maturity and relative heights were determined simply by subtracting the value for one variety from the value for the other. In each case, of course, the sign was determined in the same way. A simple example explained in detail may serve to make this method clear. In the wheat variety test of 1921 the varieties Fultz (Bayer), Michigan Amber, and Michigan Wonder No. 211 occurred Experiments in Field Plot Technic 33 in the order named in four distributed sections of the field. The aver- age yields of these varieties in the four series, in bushels per acre, for border rows and for interior rows, are shown below, together with the average dates of heading, dates of maturity, and heights, also de- termined for the four series. 23. Fultz (Bayer) 39. Michigan Amber 55. Michigan Wonder No. 211 Row 1 Row 2, 3, 4 Row 5 Row 1 Row 2, 3, 4 Row 5 Row 1 Row 2, 3, 4 Row 5 Average yields 10.8 bu. 12.2 bu. 13.1 bu. 13.3 bu. 14.9 bu. 14.5 bu. 19.8 bu. 18.1 bu. 19.4 bu. Average date of heading 21* 21* 19* Average date of maturity 47* 48* 47* Average height 43 in. 42 in. 43 in. * Dates of heading and maturity are the numbers of days after April 30. Thus 1 is May 1, 32 is June 1, 47 is June 16, etc. Now dividing the yields in border rows by the yields of the same varieties in interior rows, we obtain the relative border yields, which are substituted in the table below for the border yields in bushels. To determine the degree of competition between the varieties Fultz and Michigan Amber we divide the larger relative border yield (107) by the smaller (89) and subtract 100, giving 20 per cent. Since in this case the relative border yield of the variety on the left is higher, the difference is given a minus sign. Similarly a value of +12 per cent is obtained for the competition between Michigan Amber and Michigan Wonder No. 211. These figures mean that the relative border yield of Fultz ex- ceeded that of Michigan Amber by 20 per cent in their competing border rows, while that of Michigan Wonder exceeded that of Michi- gan Amber by 12 per cent. The relative yields of these varieties are obtained similarly, — in the first case by dividing 14.9 by 12.2 (+22%) and in the second case by dividing 18.1 by 14.9 (+21%). Both values are positive be- cause in each case the yield of the variety on the left is higher than that of the variety on the right. The difference in dates of heading, maturity, and height are obtained simply by subtraction, being positive when the value of the variety on the right is greater and negative when 34 Missouri Agr. Exp. Sta. Research Bueletin 49 the value of the variety on the left is greater. The figures ready for correlation study will then appear as follows : 23. Fultz (Bayer) Row Row Row 1 2,3,4 5 Compe- tition data 39. Michigan Amber Row Row Row 1 2, 3, 4 5 Compe- tition data 5 5. Michigan Wonder No. 211 Row Row Row 1 2, 3, 4 5 Average yield 89 12.2 107 -20% +22% 89 14.9 97 + 12% +21% 109 18.1 107 Average date of heading 21 0 21 —2 19 Average date of maturity 47 + 1 48 0 47 Average height 43 — 1 42 + 1 43 The columns headed “competition data” show the relation of the effect of competition to the yield, earliness, and height of the competing varieties. For example, Michigan Amber was at a disadvantage of 20 per cent in competition with Fultz, though it was 22 per cent superior in yield when protected from competition. It headed the same day, matured one day later, and was one inch shorter. After correspond- ing data had been prepared for all the 96 varieties in this test, correla- tion tables with the coefficient of competition as subject and relative yield, date of heading, date of maturity, and height as relative were constructed. Correlations were determined similarly in the other tests. One of these correlation tables is shown in figure 6. In general, merely o o o o o o o o o o o CM o 00 CM CM to 00 © 7 1— 1 1 1 1 1 | rH o o o o o O O o o o o o o o o o o o o o o O 4-» /— s o 1— 1 1 00 1 o 1 1 1 CM tO 00 W H —40 to —60 1 1 —20 to —40 2 1 1 2 4 2 2 14 0 to —20 1 1 7 10 5 2 26 0 to 20 1 2 6 10 8 3 5 35 20 to 40 2 3 4 11 2 1 1 24 40 to 60 1 1 3 5 60 to 80 1 1 1 3 Total 2 1 3 5 20 26 21 16 8 5 1 112 Figure 6.— Correlation Between Coefficient of Competition and Rela- tive Yield, in Wheat Variety Test 1920. r= +.582 ± .043. Experiments in Field Plot Technic 35 the coefficient of correlation and its probable error are given, for lack of space. In the barley variety test, 1919, the effect of competition was quite marked. The average yield of border rows differed from the average yield of interior rows by 11.13 per cent, and the mean coefficient of competition was 21.30 per cent. Attempts were made to correlate competition with relative date of heading, date of maturity, grain-straw ratio, and yield. The correlation coefficients determined are shown in Table 10, together with the mean differences between competing Table 10. — Correlation of Competition With Various Characteristics in Barley Varety Test 1919. Character Date of heading Date of maturity Grain-straw ratio Yield Mean difference be- Coefficient of correlation tween competing with competition varieties 4.0 days 2.6 days 38.0% 52.3% —.153 ±.120 —.063 ±.123 + .072 ±.122 + .442 ±.099 varieties in the characters whose relation to competition was studied. Although none of these correlations is statistically significant, in the strictest sense, it is noticeable that the correlation between compe- tition and yield is much greater than any of the others, and is equal to about four and one-half times its probable error. There was ap- parently some tendency for the better yielding varieties to profit by competition with the poorer yielders. On account of the relatively small number of cases involved in this and the other 1919 tests, the probable errors are high, and a fairly high coefficient of correlation Table 11. — Correlation of Competition With Various Characteristics in Oats Variety Test 1919. Character Date of maturity Grain-straw ratio Yield Mean difference be- Coefficient of correlation tween competing with competition varieties 3.56 days —.456 ±.103 50.2% —.091 ±.129 53.5% +.314 ±.117 may consequently fail to attain statistical significance. Such a co- efficient, while not establishing the correlation, by no means indicates that the correlation does not exist. 36 Missouri Agr. Exp. Sta. Research Bulletin 49 The oats variety test of 1919 also showed distinctly the effects of competition. The border rows in this test differed in yield from the interior rows by 1278 per cent, on the average, and the mean coefficient of competition was 27.67 per cent. Correlations were de- termined for competition and relative yield, date of maturity, and grain-straw ratio. Unfortunately the dates of heading are not avail- able for all varieties in this test. The correlation coefficients are shown in Table 11. Again no correlations of statistical significance are found, but the relation of yield and earliness of maturity to competing strength is at least suggestive. There was a tendency for early and high-yielding varieties to profit by competition at the expense of later and lower-yield- ing varieties, but the number of varieties was too small to permit the drawing of positive conclusions. The oats strains grown on the same field showed much less strik- ingly the effects of competition. The mean difference in yield be- tween border rows and interior rows in these 15 strains was only 6.50 per cent and the mean coefficient of competition only 13.11 per cent. This is undoubtedly accounted for by the fact that the differences be- tween competing strains were so much less than in the oats variety test. When the three strains taxonomically unlike Red Rustproof are eliminated, leaving 12 strains of the same variety, the average devia- tion of border yields from interior yields is reduced to 4.69 per cent and the average coefficient of competition to 8.69 per cent. It is note- worthy that the competition between these strains of the same va- riety is decidedly less than that between different varieties. No sig- Tabee 12. — Correlation oe Competition With Various Characteristics in Oats Strain Test 1919. Character Date of heading Date of maturity Grain-straw ratio Yield Mean difference be- Coefficient of correlation tween competing with competition strains 2.67 days 1.56 days 14.2% 17.1% —.376 ±.136 — .244 ±.149 + .012 ±.159 +.316 ±.143 nificant correlation was found between these minor effects of compe- tition (for the 15 strains) and the relative time of heading, time of maturity, grain-straw ratio or yield, as is shown in Table 12, though in this case again the early strains and the high-yielding strains showed some tendency to profit by competition. Experiments in Field Plot Technic 37 In the wheat variety test of 1920 the average yield of border rows differed from the average yield of interior rows by 12.30 per cent and the mean coefficient of competition was 19.79 per cent. These figures represent the average determinations when the two varieties of rye and the border yields of the varieties of wheat adjoining them were eliminated. The correlation between competition and relative yield, date of heading, and date of maturity were determined for this test and the coefficients of correlation are shown in Table 13. Table 13. — Correlation of Competition With Various Characteristics in Wheat Variety Test 1920. Character Mean difference be- Coefficient of correlation tween competing with competition varieties Date of heading 2.3 days — .515 ±.048 Date of maturity 2.7 days — .552 ±.045 Yield 28.9% +.582 ±.043 Competition in this test was negatively correlated with earliness of heading and maturity and positively with yield. All of the correla- tion coefficients are clearly significant. In other words, there was a rather pronounced tendency for the early and high-yielding varieties to profit in competition. To a considerable extent the early varieties were the high yielding varieties in this test, as indicated by the fact that the correlation coefficient for date of heading and yield was —.511 ±.051, and that for date of maturity and yield was —.642 ±.041. Al- though it is clear from these results that early, high-yielding varieties excelled in competition, it is not clear whether they did so chiefly as a result of their earliness or chiefly as a result of their yield. Table 14. — Correlation of Competition With Various Characteristics in Wheat Variety Test 1921. Character Mean difference be- Coefficient of correlation tween competing with competition varieties Date of heading Date of maturity Height Yield 2.1 days 1.6 days 3.3 inches 19.5% —.271 ±.060 —.222 ±.062 + .347 ±.057 + .294 ±.059 Similar results were obtained in the wheat variety test of 1921 in which the difference between the average yield of border and in- terior rows was 12.89 per cent and the mean coefficient of competition 38 Missouri Agr. Exp. Sta. Research Bulletin 49 was 18.85 per cent. Correlations were determined for competition and relative yield, date of maturity, date of heading, and height in this test. The coefficients of correlation thus determined are shown in Table 14. In this case, as in the wheat variety test of the preceding season, dates of heading and maturity were correlated negatively and yield was correlated positively with competition. The coefficients of correla- tion were materially lower, and in fact are hardly significant. It is in- teresting that in this case height was correlated more closely with competition than were either earliness or yield. In this season again earliness was correlated to some extent with yield, the coefficients of correlation, for date of heading and yield being — .331 ±.062 and for date of maturity and yield —.419 ±.057. In the wheat mixture test of 1921 the varieties were grouped roughly in respect to earliness, and in only three cases was there a greater difference than two days in heading or maturity between ad- jacent varieties. The rows in this test ran north and south. The conditions may be considered favorable in this test for the reduction of competition. Nevertheless the average yield of border rows dif- fered from that of interior rows by 10.07 per cent and the mean co- efficient of competition was 14.28 per cent. The coefficients of correla- tion determined for competition and date of heading, date of maturity, and yield, are shown in Table 15. Table 15. — Correlation of Competition With Various Characteristics in Wheat Mixture Test 1921. Character Mean difference be- Coefficient of correlation tween competing with competition varieties Date of heading 1.2 days — .514 ±.083 Date of maturity 0.8 days — .613 ±.070 Yield 19.2% +.554 ±.078 In this test significant negative correlations between competition and dates of heading and maturity and a significant positive correla- tion between competition and yield are shown. The tendency for early, high-yielding varieties to profit by competition was about as strong as in the wheat variety test of the preceding season, though the extent of competitive effect was considerably reduced. The effects of competition in the oats variety test in 1921 were extreme. The yields of border rows differed by 16.74 per cent, on the average, from the yields of interior rows, and the mean coefficient of competition was 39.15 per cent. The extreme effects of compe- Experiments in Field Plot Technic 39 tition in this test are probably accounted for by the fact that the va- rieties differed very widely in varietal type and in yield. Differences of as much as 17 days in date of heading, 13 days in date of maturity, and almost 200 per cent in yield, were involved. The correlations de- termined between competition and relative date of heading, date of maturity and yield, are shown in Table 16. Table 16. — Correlation of Competition With Various Characteristics in Oats Variety Test 1921. Character Mean difference be- Coefficient of correlation tween competing with competition varieties Date of heading 4.8 days — .648 ±.060 Date of maturity 4.1 days — .860 ±.028 Yield 51.1% +.484 ±.082 A remarkably high negative correlation between date of maturity and competition is shown. The negative correlation between date of heading and yield is also quite high, while the positive correlation be- tween yield and competition is barely significant. In this test, in which extreme differences in time of maturity occurred, the early-maturing varieties had a very distinct advantage in competition with the later varieties. Earliness was very closely correlated with yield in the oats variety test of this season, the coefficient of correlation for date of heading and yield being —.750 =+052 and that for date of maturity and yield being —.894 ±.024. Considering the close correlation of earliness and yield, and the relatively low correlation of yield and competition, it would seem that the latter may be merely a by-product of the rela- tion of earliness to competition. Since the early varieties were the leaders both in competition and in yield, some correlation of yield and competition is inevitable. In the oats strains test of 1921 Kherson and Red Rustproof strains were alternated and both a Kherson and a Red Rustproof check were grown. In most cases therefore the competing border rows repre- sented these two varieties, though in some cases two Red Rustproof or two Kherson plots occurred together, as is shown in the planting plan in figure 5. The effects of competition in this plot were quite distinct, as is to be expected, though they were not so extreme as in the oats variety test discussed above, which was located on the same field. The average yield of border rows differed from the average yield of interior rows by 11.76 per cent. The mean coefficient of compe- tition was 23.85 per cent. When we exclude the competition between the three strains not true to name and the strains adjacent to each, that between the Kher- 40 Missouri Agr. Exp. Sta. Research Bulletin 49 son and Red Rustproof check plots, and that between adjacent strains of the same variety, 58 cases of competition between different strains of Kherson and Red Rustproof remain. In these the mean yield of border rows differed from that of interior rows by 14.06 per cent and the mean coefficient of competition was 30.86 per cent. In every case the Kherson strain outyielded the adjacent Red Rustproof strain, though the advantage in yield varied from 27 per cent to 165 per cent. Similarly, the Kherson strains were earlier in maturity and heading, and taller, in each case, but with a rather wide variation in the extent of their advantage. In all but three of the 58 cases the Kherson strains showed a greater advantage in yield over the adjacent Red Rustproof strains in their competing border rows than in their interior rows. The average yields of the 30 Red Rustproof strains and 29 Kherson strains, in interior rows and competing border rows, were as follows : Average yield in Average yield in interior rows competing border rows Bushels Relative Bushels Relative Red Rustproof strains 21.00 100 18.59 100 Kherson strains j 35.79 170 41.00 222 The Kherson strains outyielded the Red Rustproof strains by 70 per cent in their interior rows and by 122 per cent in their competing border rows. The coefficients of competition, like the relative yield, earliness, and height, varied rather widely. Correlations were there- fore measured for the advantage of the Kherson strain of each adjacent pair in competition and its advantages in yield, date of heading, date of maturity, and height. The coefficient of correlation in each case was insignificant. Discussion . — In each of these tests, with the exception of the oats strain test of 1919, in which most of the strains compared be- longed to the same variety, border rows differed from interior rows in yield by more than 10 per cent. Differences as great as this will change materially the relative standing of varieties. In single-row tests the effects of competition would be considerably greater than in these border rows, affected by competition on only one side. Furthermore, in each test, of course, there were many cases in which competition caused much larger differences in yield than are shown by average figures. Experiments in Field Plot Technic 41 The relation of the direction of rows to the effects of varietal com- petition is not clearly shown by these experiments. The tests which showed least the effect of competition, the oats strain test of 1919 and the wheat mixture test of 1921, were in rows running north and south. But relatively little effect from competition is to be expected in these tests, regardless of the direction of the rows, because of the similarity of adjacent strains. In the oats strain test 12 of the 15 strains were taxonomically identical, and it has been shown that the effects of competition among these was much less than among the strains of different varieties. In the wheat mixture test the varieties making up each mixture, which were grown side by side in the test, were chosen partly for their similarity in time of maturity, and the differences between adjacent varieties were therefore considerably less than in the wheat variety test of the same season. It cannot be stated definitely, therefore, from the results of these tests, that tests in rows running north and south are either more or less subject to error from competition than tests in rows running east and west. It is clear, however, that a considerable error from varietal competition may occur in tests in which the rows run north and south, as is evi- denced particularly by the barley and oats variety tests of 1919. The relation of competition to relative date of heading, date of maturity, grain-straw ratio, height, and yield, insofar as it was in- vestigated in these experiments, is shown in summary form in Table 17. Although none of these characteristics shows a significant rela- Tabee 17. — Summary of Effects of Competition in Aee Tests. Test Season .... Number of varieties or strains j Mean coeffi- cient of competition. . . Date of Heading. Coefficient of Correlation between Competition and — Date of Maturity. Grain- Straw Ratio. Height. Yield. Barley variety 1919 | 27 21.30 — .153 ± . 120 — .063±.123 + .072 + .122 + .442±.099 Oats variety 1919 24 27.67 — .456±.103 — .091 + . 129 -f-.314-H.H7 Oats strain 1919 15 13.11 -.376±.136 — .244±.157 + .012 + .159 + .316+.143 Wheat variety 1920 1 94 19.79 — .5l5±.048 — .552 + . 045 + .582 + . 043 Wheat variety 1921 94 18.85 — .271±.060 — .222 + . 062 + .347±.057 + .294 + .059 Wheat mixture 1921 1 30 14.28 — .514 + . 083 — .613±.070 + .554±.078 Oats variety 1921 32 39.15 — .648 + .060 — .860±.028 +.484 + .082 tion to competition in every case, the results of the tests are fairly consistent. The correlation of competition with yield is always posi- tive, and is fairly high in every case, the lowest coefficient being -(—.294 —.059. From these results there can be no doubt that the higher yield- ing varieties are those which in general have profited by competition. The date of heading and the date of maturity show a negative correla- 42 Missouri Agr. Exp. Sta. Research Bulletin 49 tion with competition in each case, though some of the coefficients are insignificant. It is clear therefore that early varieties are, in gen- eral, able to compete more strongly, but the extent of this relation is quite variable. The grain-straw ratio showed no significant relation to competition in any of the experiments of 1919, and was not deter- mined for the succeeding tests. Height was correlated positively with competition in the one test in which height was determined, the wheat variety test of 1921. In this test height was more closely related to competition than were date of heading, date of maturity, or yield. In the oats variety tests, the relation of early maturity to compe- tion is particularly marked, the coefficients of correlation in both oats variety tests being distinctly greater for date of maturity and compe- tition than for yield and competition. In the wheat tests there was little difference in the degree of relation to competition between earli- ness and yield. In the one test of barley varieties conducted, yield was more closely correlated with competition than was either the date of heading or date of maturity, but none of the three showed a clearly significant correlation. It is clear that in these trials the early, high-yielding varieties profited by competition. To a considerable extent these may be the same varieties, for the correlation of earliness and yield was high in most of the tests conducted. The relation of earliness and other characters to yield under Missouri conditions will be considered more fully in another paper, but data of interest in this connection are ap- propriate here. The coefficients of correlation of yield with date of heading and date of maturity in the variety tests discussed in this paper are shown in Table 18. Table 18. — Correlation of Yield With Dates of Heading and Maturity in Variety Tests of Barley, Oats, and Wheat Coefficient of correlation of Number of yield with — Crop Season varieties Date of heading Date of maturity Barley 1919 27 —.281 ±.120 —.271 ±.120 Oats 1919 40 —.627 ±.065 Oats 1921 32 — .750 ±.052 — .894 ±.024 Wheat 1920 94 — .511 ±.051 — .642 ±.041 Wheat 1921 94 —.331 ±.062 —.419 ±.057 When a very high correlation exists between earliness and yield it is likely that a character closely correlated with one may show a high degree of correlation with the other, which might not be shown were it not for the first correlation. For example, suppose earliness Experiments in Field Plot Technic 43 of maturity is largely responsible for strong competitive value. Then in a season when earliness is closely correlated with yield a close cor- relation of competition and yield is likely to be found, not because high yield makes for strong competition but because the high-yielding va- rieties are early. Conversely, the competing value may be dependent on the yield and the correlation with earliness may be incidental, under the same conditions. If the relation of earliness and yield were con- stant, such a question would have little practical importance, but when the relation is reversed, as it may be in different localities and even in different seasons in the same locality, the relation of competition to the two characteristics may be very different. The relation of compe- tition to earliness and yield in these tests, therefore, may be due pri- marily to the predominating influence of either of these two charac- teristics, or to the influence of both. General conclusions regarding competition should not be drawn from these tests. The problem of competition is complicated by many factors, and will require numerous and extensive investigations for its solution. These results, however, indicate that gross errors from this source are commonly involved in variety tests, that such errors occur both in rows running east and west and in rows running north and south, that the error is less when the varieties and strains com- pared are structurally similar than when they are widely different, and that the error may be reducible to some extent by the grouping of va- rieties according to the time of maturity and possibly other characters, when the relation of such characters to competition is more fully studied. In the present state of knowledge regarding the relation of competition to the characteristics of the varieties compared, the use of border rows is highly desirable, since by their use the error from competition can be practically eliminated. SIZE AND REPLICATION OF PLOTS. Previous Investigation. — Most of the direct evidence reported on replication and size of plots has been obtained in experiments in which a field of a uniformly handled crop is harvested in a large num- ber of small sections. These sections are grouped to form plots of different shapes and sizes, and systematically distributed sections are averaged to represent replicate plots. The relative variability of the yields determined by each plot arrangement is the criterion of expe- rimental accuracy. Such experiments have been reported by Morgan 15 with wheat and fodder corn, Wood and Stratton 18 with mangels, Mer- cer and Hall 12 with wheat and mangels, Hall and Russell 3 with wheat, 44 Missouri Agr. Exp. Sta. Research Bulletin 49 Montgomery 13 ’ 14 with wheat, Kiesselbach 5 with oats, and Day 1 with wheat. The general conclusions drawn from these experiments are in harmony, though the specific size and shape of plot and number of replications found most desirable vary rather widely. In general, plot variability was reduced by increasing the size of the individual plot, up to a certain limit, but it was reduced much more effectively by rep- lication of plots. For a given area a large number of small plots was always found more accurate than a small number of large plots. But the size of the plot cannot be reduced indefinitely for several reasons. As the plot becomes smaller the proportion subject to “bor- der effect” rapidly becomes greater. This border effect may be due to the modified growth of plants adjoining an alley or to the in- fluence of the competition of different varieties in adjacent rows. If the borders are not discarded an important systematic error is involved ; if they are discarded a considerable portion of the land and labor is lost. In either case the disadvantage is increased as the size of the plot is decreased. When single rod-row plots ar,e used the whole plot is subject to border effect. The importance of this error has already been discussed. Another disadvantage of the extremely small plot is that slight differences in stand and small mechanical er- rors have a marked effect on the yields. The increased labor involved in handling a large number of small plots rather than a small number of large plots is also an important disadvantage. The length of the so-called rod-row has usually been determined by convenience. Commonly used lengths when the rows are a foot apart are 16 feet for wheat, 20 feet for barley, and 15 feet for oats, since with these lengths yields in grams per row may easily be con- verted to bushels per acre. In other cases the most convenient length is determined by the dimensions of experiment fields. Although in- creasing the length of the row would doubtless reduce variability, a greater gain could be made on the same area by further replication. Ordinarily it is preferable, therefore, to retain the most convenient length and to make any desired increase in size of plot in the width, for widening the plots will rapidly reduce the proportion subject to border effect. Experimental Results. —Size of Plots . — By comparing the stand- ard deviations of single rows and blocks consisting of three and five rows each, in the check plots, it is possible to determine the relative value of plots of the three sizes in counteracting plot variability. In this comparison the single-row and three-row plots correspond respec- tively to 3-row and 5-row plots in which the border rows are dis- Experiments in Field Plot Technic 45 carded, since they are made up of rows protected from varietal compe- tition by border rows. In each of the computations summarized be- low each check plot is represented by only one yield. For example, in determining the yield and standard deviation of single rows in the 20 check plots of the oats variety test of 1919, the constants for single rows are the average of determinations made independently for Row 2 of each of the 20 plots, for Row 3, and for Row 4. The determina- tions for 3-row plots are similarly made from the computed yields of the three interior rows of each check plot, and those for 5-row plots from the computed yields of the entire plots. Thus each determination represents the same number of plots and the same area, the only dif- ference being in the size of the individual plot. It would be possible, of course, to test 40 per cent more varieties with the same number of replications or to increase the number of replications by 40 per cent for the same number of varieties on the same area, if 3-row blocks were used rather than 5-row blocks. The yield and variability of check plots of different sizes in the barley variety test of 1919 are shown in Table 19. The variety grown in these check plots was Oderbrucker, seeded at the rate of 8 pecks per acre. The check variety was grown in every sixth plot. Table 19. — Yield and Variability of Check Plots. Single-row, Three-row, and Five-row — Barley Variety Test 1919. Size of plot Number of plots Yield per acre Standard deviation bu. bu. % Single-row Row 1 21 41.26 7.95 19.26 Row 2 21 36.71 8.30 22.61 Row 3 21 37.15 8.48 22.84 Row 4 21 35.82 10.37 28.96 Row 5 21 42.12 11.86 28.16 Mean of three interior rows 21 36.56 9.05 24.80 Mean of five rows 21 38.61 9.39 24.37 Three-row Plot (Interior rows) 21 36.56 8.11 22.18 Five-row Plot 21 38.61 8.29 21.47 The variability of the single-row plots is 12 per cent higher on the average than that of the 3-row plots. That is, 3-row plots with 46 Missouri Agr. Exp. Sta. Research Bulletin 49 borders discarded would have given in this case somewhat more va- riable results than 5-row blocks with borders discarded. The same 5- row blocks harvested entire (with borders retained) gave slightly less variable yields than when the borders were discarded. The same comparison may be made in the check plots of the oats variety test of 1919. The check variety was Red Rustproof, drilled at the rate of 10 pecks per acre in every ninth plot. The re- sults are shown in Table 20. Table 20. — Yield and Variability of Check Plots. Single-row, Three-row, and Five-row — Oats Variety Test 1919. Size of plot Number of plots Yield per acre Standard deviation bu. bu. % Single-row Row 1 20 44.64 10.37 23.23 Row 2 20 47.97 9.81 20.45 Row 3 20 46.56 11.42 24.53 Row 4 20 46.95 13.81 29.41 Row 5 20 42.09 11.58 27.51 Mean of three interior rows 20 47.16 11.68 24.80 Mean of five rows 20 45.64 11.40 25.03 Three-row Plot (Interior rows) 20 47.16 10.62 22.59 Five-row Plot 20 45.64 9.72 21.30 The results in this case are practically identical with those of the barley variety test. Protected single rows were 10 per cent more va- riable than protected 3-row blocks, while the latter were only 6 per cent more variable than unprotected 5-row blocks. In the test of strains of Red Rustproof oats, conducted on the same field in 1919, adjoining the oats variety test, the same variety was used as check, and the crop was seeded on the same day with the same machine, but the check plots were in every sixth instead of every ninth plot. The corresponding data for these check plots are given in Table 21. Although the variability of these plots is lower, the relative va- riability of plots of different sizes is similar to that of the variety test. The single interior rows are on the average 24 per cent more variable than the 3-row block. The 3-row plot is only very slightly more va- riable than the 5-row plot. Experiments in Fieed Plot Technic 47 Table 21. — Yield and Variability of Check Plots. Single-row, Three-row, and Five-row. — Oats Strain Test 1919. Size of plot Number of plots Yield per acre Standard deviation bu. bu. % Single-row Row 1 18 41.87 6.35 15.15 Row 2 18 40.88 5.52 13.51 Row 3 18 43.50 5.81 13.37 Row 4 18 45.00 7.31 16.25 Row 5 18 41.50 6.37 15.35 Mean of three interior rows 18 43.13 6.21 14.38 Mean of five rows 18 42.55 6.27 14.73 Three-row Plot (Interior rows) 18 43.13 5.04 11.68 Five-row Plot 18 42.55 4.86 11.41 In the wheat variety test of 1920 the check variety was Fultz, which was seeded at the rate of six pecks per acre in every seventh plot. The results of interest in this connection are shown in Table 22. Table 22. — Yield and Variability of Checks Plots. Single-row, Three-row, and Five-row. — Wheat Variety Test 1920 Size of plot Number of plots Yield per acre Standard deviation bu. bu. % Single-row Row 1 80 20.74 6.58 31.72 Row 2 80 17.28 5.02 29.05 Row 3 80 18.34 4.50 24.52 Row 4 80 17.29 5.10 29.48 Row 5 80 19.37 6.00 30.97 Mean of three interior rows 80 17.64 4.87 27.68 Mean of five rows 80 18.60 5.44 29.15 Three-row Plot (Interior rows) 80 17.64 4.43 25.11 Five-row Plot 80 18.63 4.77 25.60 48 Missouri Agr. Exp. Sta. Research Bulletin 49 Again the single rows are distinctly more variable than the 3-row plot, in this case to the extent of 10 per cent. The 5-row and the 3-row plots are about equally variable, the slight advantage in this case being in favor of the latter. To summarize, it is evident that the protected 3-row plot is some- what less subject to plot variability than the protected single-row, but the relative value of the 5-row plot harvested entire and the same plot harvested as a protected 3-row block is not clear. Some further comparison of these two methods was made in 1921. The variability of the check plots in both the wheat and oats tests was computed as protected 3-row and as unprotected 5-row plots. In the wheat tests the check variety was Poole, seeded at 5 pecks per acre in every seventh plot in the variety test, and in every sixth plot in the mixture test. In the oats tests the check variety was Kherson, seeded at 10 pecks per acre in every sixth plot. The results are shown in Table 23. Table 23. — Yield and Variability of Check Plots. Three-row and Five-row. — Wheat and Oats Tests, 1921. Size of plot Number of plots Yield j per acre j Standard deviation Wheat Variety Test Three-row Plots 80 bu. 14.89 bu. 2.16 % 14.50 (Interior rows) Five-row Plots 80 13.98 1.90 13.61 Wheat Mixture Test Three-row Plots 30 15.48 3.25 20.98 (Interior rows) Five-row Plots 30 15.78 3.55 22.49 Oats Variety and Strain Tests Three-row Plots 120 37.95 4.61 12.15 (Interior rows) Five-row Plots 120 38.37 4.70 12.25 In no case are the differences very great. The variability of 3-row blocks is slightly greater in the mixture test and that of 5-row blocks in the variety test of wheat. There is practically no difference between the two in the oats tests. Apparently there is no constant material gain in plot uniformity obtained by the inclusion of the border rows of the 5-row plot, even though the size of the plot is materially increased by this procedure. Even if variability were decreased by their inclusion, the practice would be of doubtful value in most tests, for the reasons given in the last section ; but with practically no decrease in variability there is left no Experiments in Field Plot Technic 49 reason for the harvesting of these rows. They are not wasted because they are not harvested, for they serve a valuable purpose; the waste would be involved rather in harvesting them, for the added labor and expense would contribute nothing to the accuracy of the experiment. Although protected 3-row plots are less variable than protected single-row plots, they are not necessarily preferable. Three protected 3-row plots require the same area as five protected single-row plots, and the harvesting of almost twice as large a crop (nine rows in the first case for every five in the second). If the mean yield of five single rows has as low a probable error as the mean yield of three 3-row plots, the protected single-row plot will ordinarily be pre- ferable, because of the reduction of labor in harvesting and thresh- ing. When the standard deviation of the check plot yields is known, the probable error of the mean of any number of replicate plots can be computed and the number of replications for any given degree of accuracy determined. If single-row plots were 29 per cent more variable than 3-row plots, the probable errors of the mean of three 3-row plots and of five single-row plots would be equal, since the prob- able error of the mean is equal to the probable error of a single deter- mination divided by the square root of the number of determinations, and since the square root of 5 is 29 per cent greater than the square root of 3. In the cases herein cited the advantage of the 3-row plots was considerably less than 29 per cent in every case, and we may con- fidently expect therefore that protected single-row plots repeated five times will be less variable than protected three-row plots repeated three times, which would require the same area and more labor. Some further evidence on the relative variability of the protected 3-row plot and the unprotected 5-row plot, or, in other words, of 5-row plots, harvested with and without their border rows, may be ob- tained from the yields of the tested varieties and strains. Since the number of replications of each strain is small, average deviations are given instead of standard deviations. The inclusion of border rows in the 5-row plots should not increase variability, since the adjacent va- rieties are the same in each series, and the competitive effect should be no more variable than would be that of the same variety. A clear-cut comparison of 5-row and 3-row plots is therfore available in this case. In the case of the check plots this comparison was somewhat obscured by the competitive effect of different varieties on the border rows, which might be expected to increase variability and thus to conceal a possible advantage of the 5-row plot. The average variability of 3-row and 5-row plots in the strains tested in these experiments is shown in Table 24. Tn each case the 50 Missouri Agr. Exp. Sta. Research Bulletin 49 figure given is the mean of the average variabilities determined for all of the varieties or strains in the experiment. Table 24. — Yield and Variability oe Test Plots. Three-row and Five-row. Test Season Number of vari- eties I Number 1 of Repli- j cations - Three-row Plots 1 Five-row Plots Yield bu. per acre Average ! Devia- tion % Yield bu. per acre ! Aver- i age ; Devia- i tion % Barley varieties 1919 27 3 22.06 15.35 21.95 15.44 Oats strains 1919 15 4 50.07 5.96 50.20 ! 5.10 Wheat varieties 1920 | 96 4 13.39 24.27 13.78 | 24.36 Wheat varieties 1921 96 i 4 15.42 10.30 15.57 9.74 Wheat mixtures 1921 i 30 ! 4 17.62 9.84 18.15 10.03 Oats varieties 1921 32 i 4 29.85 10.86 30.70 i i 10.14 Oats strains 1921 64 i 4 28.40 10.82 j 28.63 10.58 There is no consistent difference in variability between the 3-row plots and the 5-row plots. In some cases the former are more va- riable; in others the latter; and in no case is the difference in varia- bility great. These results are contrary to the general impression that variability decreases with increase in size of plots. Apparently, in tests of this kind, the 3-row plot is la^ge enough to give a fair sample and nothing is gained by adding the other two rows. When it is con- sidered that the addition of these two rows undoubtedly introduces systematic error from competition to a greater or less extent, and involves a very considerable increase in the labor of harvesting and threshing, there remains little doubt that the border rows of 5-row plots are best discarded in experiments of this sort. Replication of Plots . — It is generally considered that the error from soil variability may be reduced to any desired point by replica- tion in sufficient degree. For any given degree of precision the num- ber of replications required is dependent on the variability of the replicate plots. When every plot in a single-row test is provided with two border rows the area required for the test is tripled, the replicate plots are separated more widely, and variability is usually increased, since the range of soil variability will usually be greater when a larger area is included. The removal of border effect from the rows harvested for yield may in some cases reduce variability more than enough to balance this increase, but when the unprotected single rows are grown in the same order in each series, variability will not be much affected by competi- tion, as before stated. Consequently more replications of single-row Experiments in Field Plot Technic 51 plots protected by borders than of the single-row plots not so pro- tected may actually be required for a given degree of plot variability. Similarly, more replications may be required in a test of a large num- ber of strains than in a test of a small number, as Montgomery 14 has suggested. The number of replications required may be determined with a fair degree of accuracy from the variability of the check plots. The variability of the check plots in parts of the large fields used as com- pared with the variability of the check plots in the whole fields shows the importance of this point. In Table 25 are given the standard de- Table 25. — Relation of Plot Variability to Size of Experiment Field. Check Plots in Wheat Variety Test 1920. Size of field No. of Plots Yield bu. per acre Standard deviation bu. % Four ranges (1st)* 20 14.79 3.789 25.62 Four ranges (2nd) 20 18.35 4.073 22.20 Four ranges (3rd) 20 16.67 3.659 21.94 Four ranges (4th) 20 20.74 3.876 18.69 Mean 20 17.64 3.849 22.11 Eight ranges (1st) 40 16.57 4.316 26.05 Eight ranges (2nd) 40 18.71 4.285 22.90 Mean 40 17.64 4.302 24.48 Sixteen ranges 80 17.64 4.430 25.11 ♦The four-range and eight-range sections are in order from west to east. viations of the yields of the check plots in the wheat variety test of 1920. The yields of the three interior rows of the check plots were used in computing these constants. Twenty-four varieties could have been replicated four times in the four ranges comprising any quarter of the field. As the probable error of a single plot yield is 14.92 per cent we may conclude that the probable error of the mean of four such yields would be about 7.46 per cent. But when 96 varieties must be tested, as they were in this test, four replications require 16 ranges, and the probable error of the mean yield becomes 8.47 per cent. A degree of precision which could be attained with four replications in a test covering four ranges could hardly be attained with five replications in a test covering sixteen ranges. Corresponding data for the wheat variety test of 1921 are given in Table 26. Although the variability in this experiment was much lower, the relative variability of large and small experiment fields was 52 Missouri Agr. Exp. Sta. Research Bulletin 49 Table 26. — Relation oe Plot Variability to Size oe Experiment Field. Check Plots in Wheat Variety Test 1921. Size of field No. of Plots Yield bu. per acre Standard Deviation bu. % Four ranges (1st)* 16 15.78 1.584 10.04 Four ranges (2nd) 16 15.48 1.586 10.25 Four ranges (3rd) 16 15.28 2.099 13.74 Rour ranges (4th) 16 13.01 2.091 16.07 Mean 16 14.89 1.840 12.53 Eight ranges (1st) 32 15.63 1.592 10.19 Eight ranges (2nd) 32 14.14 2.383 16.85 Mean 32 14.89 1.988 13.52 Sixteen ranges 64 14.89 2.159 14.50 *The four-range and eight-range sections are in order from west to east. similar. Again the degree of accuracy obtained with four replications in four ranges would have been unattainable with five replications in 16 ranges. The oats variety test and strain test in 1921 were contiguous, oc- cupying 24 ranges, with 120 check plots of Kherson oats, or one in every sixth plot. The variability of these check plots in sections of Table 27. — Relation oe Plot Variability to Size oe Experiment Field. Check Plots in Oats Variety and Strain Test, 1921. No. of Size of field plots Yield Standard deviation bu. per acre bu. % Four ranges (1st) Four ranges (2nd) Four ranges (3rd) Four ranges (4th) Four ranges (5th) Four ranges (6th) Mean Eight ranges (1st) Eight ranges (2nd) Eight ranges (3rd) Mean Twelve ranges (1st) Twelve ranges (2nd) Mean Twenty- four ranges 20 35.81 20 34.95 20 38.14 20 39.60 20 38.91 20 40.31 20 37.95 40 35.38 40 38.87 40 39.60 40 37.95 60 36.30 60 39.60 60 37.95 120 37.95 4.75 13.26 2.90 8.30 4.21 11.04 4.66 11.77 4.14 10.65 4.14 10.27 4.13 10.88 3.96 11.19 4.50 11.58 4.21 10.62 4.22 11.13 4.25 11.71 4.36 11.01 4.31 11.36 4.61 12.15 Experiments in Field Plot Technic 53 four, eight, and twelve ranges, and in the whole field of 24 ranges, is shown in Table 27. The variability of the whole field of 24 ranges was 12 per cent greater than the average variability of sections of four ranges each. In this case again, five replications in the larger field would have given less accurate results than four replications in the smaller. In each of the cases cited above a steady increase in variability is apparent as the size of the experiment field is increased. It is obvious that the substitution of 3-row plots with discarded borders for single rows will result in greater variability, and will require increased rep- lication for the same degree of accuracy. From the foregoing statements it will be clear that the number of replications necessary for a given degree of accuracy may vary con- siderably with conditions. The number to be used in any specific ex- periment should be determined from the variability of the field in question and the degree of accuracy required. The variability of the check plots is usually considered a measure of the variability of the field. But when the number of replications to be used or the extent of experimental error is determined from the variability of the check plots, it is assumed that the variability of different varieties of the same crop is approximately the same under the same conditions. This of course is not strictly true. The yield of two varieties may be deter- mined by very different factors, as has been stated, and their relative variability may also be quite different. The variability of 120 plots Table 28. — Soil Heterogeneity oe an Experiment Field as Determined From Yields oe Two Check Varieties. Oats Variety and Strain Tests. 1921. Number Average Probable error of a Check variety of plots yield bu. Standard deviation bu. % single yield determination bu. % Kherson 120 37.95 4.61 12.15 3.11 8.20 Red Rustproof 120 22.44 3.99 17.78 2.69 11.99 each of Kherson and Red Rustproof oats, grown side by side as check plots in the oats variety and strain test of 1921, illustrate the possibil- ity of a serious error in the use of the standard deviation of check plots as a measure of the variability of an experiment field. These determinations are shown in Table 28. The field would have been considered decidedly less variable if Kherson had been used as the check variety than if Red Rustproof had 54 Missouri Agr. Exp. Sta. Research Bulletin 49 been used. Both of these are standard recommended varieties for the region, though they differ decidedly in their characteristics. Both have been used frequently as check varieties at the Missouri station in past seasons. From the variability of the Kherson check plots the mean yield of four replicate plots in this experiment would be considered to have a probable error of 4.10 per cent ; from the Red Rustproof plots the same determination would be given a probable error of 6.00 per cent. A degree of precision for which we would assume four replica- tions necessary, judging from the Kherson check, would require nine replications according to the yields of the Red Rustproof check. The importance of choosing a check variety typical of the va- rieties tested, if its variability is to be considered a criterion of the variability of the field, is obvious. Whether it is possble to choose a “typical variety” for the purpose, in the case of ordinary variety tests, remains to be seen. ADJUSTMENT OF YIELDS BY MEANS OF CHECK PLOTS Adjustment of plot yields by the use of check plots has been a common practice in field experiments during recent years. It is recognized that no experiment field is perfectly uniform in produc- tivity, and the attempt is made, by means of the check plot adjustment, to compensate the varieties or treatments which chance to be located on the less productive plots for the resulting loss in yield. The com- mon method, in variety tests, is to distribute over the field, as fre- quently as practicable, check plots planted to the same variety and similarly handled in every way. The variation in yield among these check plots is then considered a measure of the productivity of the soil. By various methods, differing only in detail, the yields of the test plots in parts of the field giving high check yields are reduced, and those of test plots in parts giving low check yields are increased, in proportion to the productivity of the soil, as indicated by the yields of neighboring check plots. Previous Investigation. — Several investigations of the effect of such adjustment on the variability of replicate plots have been re- ported. The majority of these have been conducted in connection with experiments of the type discussed in the preceding section, in which uniformly handled fields have been harvested in small sections. Cer- tain of these sections, or plots, have been considered check plots, and on the basis of their yields the yields of the remaining plots have been Experiments in Field Plot Technic 55 adjusted. The reduction of variability of the adjusted plot yields is the measure of the efficiency of the method. Morgan 13 reports an experiment of this sort, in which 63 plots, planted first to wheat and then to fodder corn, in the same season, were used. The variability of the plot yields was steadily reduced as the number of check plots was increased. In a similar experiment reported by Lyon 11 , in which 37 replicate 1/100 acre plots of corn were harvested, the use of checks in every second or third plot was found to reduce variability, but they were of little value when farther apart. Montgomery 14 states that alternating check plots with test plots gives a high degree of accuracy, but the total number of plots required when this method is used is greater than when the same degree of ac- curacy is attained by the use of replication. Kiesselbach 5 reports a comprehensive trial of three methods of adjusting yields by means of check plots in a uniform field of 207 1/30-acre plots of Kherson oats. The effect on plot variability is shown in Table 29. Table 29. — Effect on Plot Variability of Adjusting Yields by Check Plots (Kiesselbach). Coefficient Method of of variability adjustment Actual Adjusted yields yields Alternate check plots. Correction based on average of two ad- jacent checks 7.85 7.01 Checks every third plot. Correction based on one adjacent check plot 7.79 7.35 Checks every third plot. Correction by progres- sive method, based on two nearest checks 7.87 6.57 From these results Kiesselbach concludes “The yield of system- atically distributed check plats cannot be regarded as a reliable meas- ure for correcting and establishing correct theoretical or normal yields for the intervening plats.” It should be noted at this point that even if adjustment by check yields were found invariably effective in experiments of this sort, 56 Missouri Agr. Exp. Sta. Research Bulletin 49 its value in ordinary variety testing would not be definitely estab- lished. The practice involves not only the assumption that the yields of the check plots are a fair indication of the productivity of the in- tervening plots for the check variety, but the further assumption that different varieties respond similarly to differing growing conditions. Adjustment of yields should therefore give better results in such ex- periments as those cited above than it could be expected to give in actual variety tests. This point is well illustrated by observations reported by Salmon 18 . Two varieties of barley, Gatami and Odessa, were grown side by side in fiftieth-acre plots in five distributed portions of a field. Gatami gave an average yield of 18.3 bushels per acre, with quite uniform yields in the five plots, as evidenced by their probable error of 0.68 bushel, while Odessa yielded 13.3 bushels per acre in the first plot, 6.35 bushels per acre in the second, and a negligible yield in the other three. Ob- viously the adjustment of the yield of either of these varieties on the basis of the other variety as a check, would enormously increase rather than decrease the experimental error. As Salmon points out, an error similar in kind though less in degree may occur commonly in variety tests, when the yields of varieties are determined by dif- ferent limiting factors. And if this is generally the case, adjustment by check yields will be of doubtful value, even if it were found to eliminate variability completely in uniform plot tests. There is a growing tendency, consequently, to discontinue the use of check plots for adjusting yields in variety tests, and to use them only to measure soil variability and to indicate the degree of error in yield determinations of the tested varieties. Adjustment of yields has never been as common in preliminary tests as in tests on larger plots, principally because of the great amount of computation neces- sary in adjusting the yields of ten or twenty replicate rod-rows of a large number of varieties, and because the yield of a single rod-row, exposed to varying competition and materially affected by small me- chanical errors, is at best a very unreliable measure of productivity on which to base the adjustment of the yields of several other plots. Experimental Results. — It would of course be very desirable to use check plots for reducing plot variability, if the method could be relied on, because of the economy of the practice. The only certain method of reducing plot variability is by means of replication, and it may be considered a fairly general rule that the variability of plots on a given field, as measured by the standard deviation or the prob- able error, will in general be reduced by replication in proportion to the square root of the number of replications. In other words, the Experiments in Fieed Plot Technic 57 variability of the mean of 16 replicate plots will be about half that of the mean of 4 replicate plots. Now the maximum use of check plots, that is, the practice of alternating check plots and test plots, requires the same land and labor as would be required by doubling the num- ber of replications, if no check plots were used. As doubling the number of replications will in general give a standard deviation about equal to the original standard deviation divided by the square root of 2, it will reduce variability about 30 % = -7071 ^ . If alternat- ing with check plots will consistently reduce variability more than 30 per cent it will be generally a more economical way to control error. Similarly, the use of check plots in every third plot requires as much land as would be required by increasing the number of replications by 50 per cent (using three replications instead of two, or fifteen instead of ten). From this relation the reduction of variability necessary if this practice is to equal replication in effectiveness can be easily com- puted. Such determinations for check plots at various intervals are shown in Table 30. Table 30. — Reduction of Variability by the Use of Check Plots Equivalent to That Probably Attainable With the Same Number of Plots by Replication. Distribution of check plots Equivalent increase in number of replications % Reduction in standard deviation to be expected by such increase in replication % Alternate plots 100.00 29.29 Every third plot 50.00 18.35 Every fourth plot 33.33 13.50 Every fifth plot 25.00 10.55 Every sixth plot 20.00 8.71 Every seventh plot 16.67 7.41 Every eighth plot 14.29 6.47 Every ninth plot 12.50 5.75 Every tenth plot 11.11 5.12 If protected single-row or 3-row plots are used in preliminary experiments a more reliable measure of soil productivity is available, and consequently the adjustment of yields is more likely to be of value, than when unprotected single-row plots are used. By the use of planting plans of the sort employed in these experiments, it is pos- 58 Missouri Agr. Exp. Sta. Research Bulletin 49 sible to adjust the yields by a somewhat shortened method. If adjust- ment of yield is effective in reducing plot variability in this sort of test it can be accomplished with but little increase in labor. In each of the tests reported in this paper a trial of the effectiveness of adjust- ing yields by means of check plots was made, the criterion of accuracy being in each case the variability of the yields of the replicate plots of each variety. Since the number of replicate plots was only three or four the average deviation was determined instead of the stand- ard deviation. Method Used in Adjusting Yields . — The method employed in ad- justing yields may be described as follows: The average yield of all check plots and the relative yield of each check plot in terms of this average (that is, the quotient obtained by dividing the yield of the in- dividual check plot by the average yield of all check plots) were de- termined. The relative yield of each check plot, expressed in per- centage of the mean check yield, is designated hereafter as the “plot value” of that check plot. When the average yield of all check plots is 25 bushels per acre, the plot value of a check plot yielding 30 bushels per acre is 120 per cent — in other words it is 20 per cent more pro- ductive than the average. Now assuming gradual change in the pro- ductivity of the soil between check plots, each test plot is assigned a plot value by interpolation. The adjusted yield of each plot is then determined by dividing the actual yield by the plot value. The short method for adjusting yields, referred to above, is based on the fact that the varieties occur in the same order in each series. Thus in the field diagrammed in figure 1, the following se- quence of plots occurs in each of the four series : ck 1 17 33 49 65 81 ck Now if the average yield of the four check plots adjoining variety 1, and the average yield of the four check plots adjoining variety 81 are each given a plot value, corresponding plot values for the mean yields of varieties 1, 17, 33, 49, 65, and 81 may be interpolated, and the mean yields may be adjusted in one operation. The same method may be used, of course, regardless of the number of replications. The result will not be exactly the same as that of averaging the adjusted yields determined individually, but will in most cases approximate it closely, the slight difference being caused by the disproportion of yield and plot value in the plots averaged. It is doubtful that either meth- od is consistently more accurate than the other. When the check plot yield is used in the adjustment of the yields of other plots it is of course essential that it should be a reliable de- termination, not unduly affected by factors not affecting the neighbor- Experiments in Field Plot Technic 59 ing plots. For example if the yield of a check plot is reduced 20 per cent by a poor stand, the adjusted yields of neighboring plots will be increased to the same extent as if the check plot yield had been low be- cause of poor soil, and will consequently be considerably higher than they should be. It is important therefore that conditions be made as fa- vorable as possible for accurate yield testing when this method is used. One cause for poor results in the adjustment of yield in some of the experiments reported in this paper was failure to protect the outside strip of check plots by means of border rows, in a few of the tests, be- Table 31. — Relative Variability of Actual and Adjusted Yields. Average Deviation in Percentage of Yield. — Barley Variety Test 1919. Planting number Variety 3 Average deviation Actual yields Adj usted interior rows 5 rows 3 interior rows % % % i yields 5 rows % 1 Hanna 906 19.81 17.82 13.15 10.80 2 Steigum 907 15.17 18.79 13.48 8.40 3 Luth 908 29.97 28.17 5.51 4.62 4 Eagle 913 26.14 29.79 9.71 12.98 6 Servian 915 18.37 17.97 7.36 5.59 7 Odessa 916 2.31 4.33 23.99 17.01 8 Lion 923 14.65 12.10 11.37 . 11.03 10 Horn 926 4.08 2.00 16.45 12.74 11 Odessa 927 13.62 9.16 21.62 13.76 12 Summit 929 5.28 6.45 14.23 11.59 13 Mariout 932 11.02 11.57 18.68 14.31 14 Odessa 934 13.73 13.91 11.18 10.31 15 Peruvian 935 13.25 17.87 12.42 16.82 16 Trebi 936 11.27 12.53 18.78 18.28 18 Oderbrucker 940 10.77 14.46 13.60 15.98 19 Frankish 953 20.53 19.65 22.63 18.49 20 Manchuria 956 6.88 6.33 13.89 10.68 21 Oderbrucker 957 17.88 13.62 1.97 3.27 22 Manchuria x Champi of Vermont on 39.47 39.19 21.93 20.35 23 Luth 972 16.77 18.61 7.05 7.48 24 Red River 973 13.94 11.02 12.37 12.33 25 Featherston 1118 21.40 20.89 8.47 13.39 26 Featherston 1119 16.59 15.25 4.78 12.26 27 Featherston 1120 15.91 13.64 2.48 6.44 28 Hanna x Champion of Vermont 1121 16.00 16.81 28.36 28.50 29 Manchuria 1125 6.79 14.42 7.78 1.55 30 Malting 1129 12.86 10.40 5.40 9.02 Mean 15.35 15.44 12.91 12.15 60 Missouri Agr. Exp. Sta. Research Bulletin 49 cause of lack of space. The check plots growing on the border of the field were materially reduced in yield, in some cases, notably the oats strain test of 1919 and the wheat variety test of 1921. In these cases the variability of the actual and adjusted yields has been computed both for all series and for the remaining series when the one affected by an unreliable check is discarded. Relative Variability of Actual and Adjusted Yields . — The relative variability of actual and adjusted yields of both 3-row and 5-row plots in the barley variety test is shown in Table 31. In this test there were three replications, and the check variety was Oderbrucker, in every Table 32. — Relative Variability oe Actual and Adjusted Yields. Average Deviation in Percentage of Yield Oats Variety Test 1919. Planting number Variety Average Deviation 3 Series 4 Series (3 interior rows) (3 interior rows) Actual Adjusted Actual Adjusted yields yields yields yields % % % % 1 A. sterilis nigra 4.32 1.46 9.18 5.02 2 Black Mesdag 9.03 9.69 7.29 12.90 3 C. I. 602 13.72 16.00 16.30 13.63 3 C. I. 603 4.72 3.09 5.84 3.88 5 C. I. 620 4.73 10.14 11.24 13.67 6 Early Champion 18.63 15.18 14.97 14.65 7 Early Gothland 14.20 4.67 11.55 4.18 8 Garton 473 5.99 6.25 8.42 9.42 9 Garton 585 14.08 19.44 16.92 19.95 10 Golden Giant 9.44 14.31 14.40 15.09 11 Irish Victor 9.69 3.29 7.72 16.40 12 Japanese Selection 6.87 4.71 11.85 5.20 13 June 18.37 11.19 17.53 10.37 14 Kherson Selection 17.01 9.20 15.06 20.36 15 Fulghum 9.69 11.36 13.06 17.32 16 Lincoln 21.07 12.54 16.56 11.83 17 Monarch 6.12 4.55 9.06 33.36 18 North Finnish 8.69 5.17 7.84 27.03 19 Scottish Chief 5.05 4.28 5.10 15.42 20 Sparrow bill (Missouri) 10.98 10.82 12.38 13.15 21 Sparrow bill (Cornell) 4.45 3.25 12.11 3.85 22 Tobolsk 1 6.17 3.85 13.92 5.38 23 Tobolsk 2 11.56 9.24 20.35 13.96 24 White Tartar 10.94 4.75 9.51 4.54 Mean 10.23 8.27 12.01 12.94 Experiments in Field Plot Technic 61 sixth plot. As a result of the adjustment of yields, the average devia- tion of 3-row plots was reduced from 15.35 per cent to 12.91 per cent, a reduction of 16 per cent, and that of 5-row plots from 15.44 per cent to 12.15 per cent, a reduction of 21 per cent. The relative variability of actual and adjusted yields in the oats variety test of 1919 is shown in Table 32. In this field the check, Red Rustproof, was in every ninth plot. When the series affected by the faulty check yields of the border plots is included the variability of the adjusted yields is slightly higher than that of the actual yields, but when this series is discarded the average variability as measured by the mean deviation is reduced 19 per cent. It might be expected that the oats strains grown on the same field would show a greater reduction of variability than the varieties, since practically all of them were of the same variety as the check, and since Table 33. — Relative Variability oe Actual and Adjusted Yields. Average Deviation in Percentage of Yield. Oats Strains Test 1919. Planting number Accession number Average deviation Actual yields Adjusted yields 3 interior rows 5 rows 3 interior rows 5 rows % % % % 1 0119 10.30 7.75 11.22 8.81 2 0120 4.70 6.58 3.01 1.76 3 0121* 5.76 3.25 4.57 4.14 4 0122 4.62 3.52 6.46 3.82 5 0123 9.91 8.25 11.47 9.62 6 0125 3.18 3.34 5.39 6.60 7 0126 7.62 5.95 10.56 16.58 8 0127* 6.76 5.09 6.92 9.85 9 0124* 6.13 6.34 4.92 4.59 10 0133 7.07 4.77 3.92 5.36 11 0128 4.17 3.56 3.58 3.36 12 0129 5.02 7.07 5.94 6.35 13 0130 4.20 2.62 6.74 9.72 14 0131 2.59 2.59 4.98 2.94 15 0132 7.38 5.81 12.38 12.08 Mean 5.96 5.10 6.80 7.04 * Not taxonomically Red Rustproof. the check plots were more frequent, being in every sixth plot. The results of adjusting yields in this test, both for protected 3-row plots and for unprotected 5-row plots in four series are shown in Table 33. Contrary to expectation, the variability was not reduced by adjustment Table 34. — Relative Variability oe Actual and Adjusted Yields. Average Deviation in Percentage of Yield. — Wheat Variety Test 1920. 62 •2s«3 • +-> .—,+j m . 2-0 c jg > £ % >T3 ^ 3 An!? bo v. •£i! Missouri Agr. Exp. Sta. Research Bulletin 49 D^OTfTflOCMONCOr'.TrOvt'.l^CMlN.rfOvOOro'OCM© ^'M-^’ooTr'cM^.^’cMCMCM^-vd^co.-H’idTi-'oidvdou-iTfvdol HHMhN CM rl r( —I —< .-H 1-1 CM 5 rf co vo On \VOOrtaO\OONNNOOOi^\£^r^rH Tj-oOOOt-^iO ^NONOPOrtOONOodiMrHcg^odrHiri^doNO O'ln^rtNCiO'tr.-tCMC'OvO'tOOCTfCOoC'OONN'OrO'ONinNrtOiOMi i CO CM tJ" co On CM Cn ,N<»3\0'OOr.VO^N'Ol ) t>. O' 3 " t}- t'* '-I ' O Tf Os 00 i-i CM ■ . \C O T-I o (NHOOino\ OOCI'OrfJCM'-iPONN't'OCIw'OONfO'OCMNMO'OO'tOO'HCJOinrHroCMNrOr* h ! rd rd Cn! 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CMCMCMrtnrtrO^,rNr.rt 5 \D O <^5 Tj- 00 O N'OdO'doO'trOTf CMCMM-M-CMCMCMr-c> d cd o 6 d ci d o d o oi n r- oo d d CM'-'CM-i'trO'I'rtfOM^CMCMCVlCMCMfO'-i i-^CMCMCMCMCMCMcorOPOCMCMco^-POCM^cOCM 00O00 ih(MCM 00 iO 0\CMM'0 '-iO'-i'O 0\“Ot^. T tOOCM^t^M-Ot^OO'CM , M-rJ--^-< v, 5ir)OOCsOOCT\CMcOOOvOUO-. OliHinNOO 00 00 00 CM CM f^vOOl rt 512 .T S o ~ ^ r ■ 00 CM uo VO 00 c^ , o o' o' o* o o' >v kkxkke 1 T3 T) T3 °0 VO (M 1 y o fi® o' » c 1 °3 zz ooooooo d co yyyyyyyyyyy c c c c O n 5 o nO^NMCOO OOoOOo 00 «= 000 ' a '°c' illll . . . . . .|| M >«>«>. sacs > s * zz~~ £ii ^2 ^ |i5 ,P ,9 42 S y y y y O y y y y y y y y y y V •-.S 1 « . y- & & & *j" « i: o — . U 2 2 8 t 3 .s 6 « a 6 ° cccccccc r 2 ri rs rt Cw Rl J — be S Crtrtrtrtrtrtrtcartra«rtrtcapis 3 ?? £ “aS IS IS IS IS IS IS IS IS IS IS IS IS IS IS IS yyyy c-' (U v.v-uvh— .- iy 33 y. 3 ^ : 3 fl ° f> riNfOM-iOvONMONO HCM'O'tiO'ONI Experiments in Fieed Plot Technic 63 of yield. A possible explanation is the extremely low variability of the actual yields, indicating that the field, which was quite small, was re- latively uniform. Any gain in uniformity from a check adjustment of yields would of course be expected to be greater in highly variable than in more uniform fields. The relative uniformity of this field is indicated not only by the low mean deviation of the test plots, but also by the low standard deviation of the check plots, which was only 11.68 per cent, as compared with a standard deviation of 22.59 per cent in the check plots of the adjoining oats variety test. The effect of adjusting yields on the variability of 3-row and 5-row plots in the wheat variety test of 1920 is shown in Table 34. In this test the check variety, Fultz, was grown in every seventh plot. There were four series of the ninety-six varieties. The reduction in variability was very marked, being 37 per cent for 3-row plots and 42 per cent for 5-row plots. The variability of almost every variety was reduced, and the reliability of the results was undoubtedly much increased. The wheat variety test of 1921, occupying an equal area on a neighboring field, and with similar varieties and the same planting plan, gave decidedly different results. In this field the check va- riety was Poole. Several check plots on the border were abnormal, and the computations are therefore given both for three series and for four, the series affected by the abnormal check yields being dis- carded in the former case. The relative variability of actual and ad- justed yields is shown in Table 35. Although the check yields are somewhat less variable for three series than for four, the adjustment was not effective in either case in reducing variability. The adjusted yields are 10 per cent more va- riable than the actual yields for the three series and 34 per cent higher for the four. Similar results were obtained in the wheat mixture test of the same season, in which several of the same varieties were included, and the same check variety was used. In this test the check variety was in every sixth plot, and four replications were used. The results of adjusting yields are shown in Table 36. Variability was increased from 9.84 per cent to 13.81 per cent, an increase of 40 per cent. Thus the results of adjusting yields of wheat varieties in 1921 are directly contrary to the results of the same practice in 1920. Difference in Results Obtained by Adjustment with Different Check Varieties . — In the oats variety and strain tests of 1921, two check varieties, Kherson and Red Rustproof, were grown. In these tests 96 strains were included, 32 of Kherson, 32 of Red Rustproof, and Table 35. — Relative Variability of Actual and Adjusted Yields. Average Deviation in Percentage of Yield. — Wheat Variety Test 1921, 64 Missouri Agr. Exp. Sta. 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S’ S’.SP.^f.SP.^.S 1 .^ >•3 £ £ ! = 'S’5 i „ ^ „ — — jHpanewrtca^.^oooooooooooooc o O'- rtrtrtfl33333333 3,3 3 3 «3 S « « ox *j-; - -55SSS5SSSIP3 : ' PPOCJUCJOCjQQ WPJWpHpHtapHpHpHpHpHpnfe^PpH Ph ffi I— I ►— >M -CMCO7UNSON00ON OHCMfONf-iovONOOCNOrHCMCiTt lONOCsOOCOrHCM'OHtmNONOOONOrHCMfO^iONONXO rHHrtrHnHrHrHrHrHHCaCMCMCMCMCMN(NlCMCMCOP)co O 4-> O O O O ■*-> O O O O O H-» O O o o o O O O o O O O O o O © © '4—* CO ■'f © 50 CO © o 03 00 © © O T_l T “ l 1-1 rH 03 03 03 03 03 03 03 03 H-i 60 to 70 1 1 70 to 80 1 1 1 3 80 to 90 1 1 1 1 1 1 6 90 to 100 2 4 1 1 1 1 10 100 to 110 1 2 2 2 4 3 2 3 1 20 110 to 120 1 1 4 3 2 2 2 2 1 18 120 to 130 1 3 4 1 4 4 2 3 22 130 to 140 1 5 1 2 3 3 1 1 1 1 19 140 to 150 1 1 1 3 2 2 1 11 150 to 160 2 2 4 160 to 170 1 2 1 1 5 170 to 180 1 1 Total 1 2 4 2 10 20 11 20 19 16 9 3 2 0 1 120 Figure 7— Correlation Between Yields of Kherson Check Plots and Yields of Adjacent Red Rustproof Check Plots, in Oats Variety and Strain Tests 1921. r= +.162 ± .060. 70 Missouri Agr. Exp. Sta. Research Bulletin 49 ductivity of the same portion of the field as determined by an adjacent Red Rustproof check plot. This correlation is shown in figure 7. The coefficient of correlation is less than three times its probable error — the correlation has not even statistical significance ! The relative pro- ductivity of different portions of the field, as indicated by the two check varieties, is shown in figure 8. If Kherson had been used as a check Figure 8. — Relative Variability of Different Parts of an Experiment Field, as indicated by the Yields of Adjacent Check Plots of Kherson and Red Rustproof Oats. Oats Variety and Strain Tests 1921. In the diagram on the left, points of equal productivity, as indicated by the yields of the Kherson check plots, are connected by lines (as points of equal elevation are connected by lines on a contour map). In the diagram on the right, the same field is similarly mapped according to the yields of the Red Rustproof check plots. The numbers indicate the plot values of the points concerned. variety for adjusting yields, the yields of certain plots would have been increased to compensate for the low productivity of the soil ; if Red Rustproof had been used the yields of the same plots would have been decreased to compensate for the high productivity of the same soil. The fact is that certain parts of the field were actually more productive than the average for Kherson oats and less productive for Red Rustproof, as is indicated by the fact that each variety of check was considerably more effective in the adjustment of the yields of strains of the same variety than of strains of the other. But neither check was a very accurate measure of the productivity of the soil, even for its own variety, as indicated by the failure of adjustment to reduce variability consistently even when Kherson strains were ad- Experiments in Field Plot Technic 71 justed according to the Kherson check and Red Rustproof strains according to the Red Rustproof check. Value and Limitations of Adjusting Yields by Means of Check Plots . — The effect on plots variability of adjusting yields by means of check plots in all of the tests is shown in summary form in Table 40. The variability of the test plots was reduced by adjustment in three tests and was increased in the other five. It is noteworthy that the three tests in which plot variability was reduced by adjustment were characterized by high plot variability, as indicated by the standard deviation of check plots, while the tests in which adjustment was not Table 40. — Summary of Relative Variability of Actual and Adjusted Yields of Interior Rows in All Tests. Test Season Number of var- eties or strains Number or rep- lica- tions Average Actual yields % deviation Adjusted yields % Barley Variety 1919 27 3 15.35 12.91 Oats Variety 1919 24 3 10.23 8.27 Oats Strain 1919 15 4 5.96 6.80 Wheat Variety 1920 94 4 24.27 15.32 Wheat Variety 1921 94 Q o 10.45 11.52 Wheat Mixture 1921 30 4 9.84 13.81 Oats Variety 1921 32 4 10.86 11.43* Oats Strain 1921 64 4 10.82 11.07* * Adjustment by Kherson check. effective were in general low in plot variability. In 1919 adjustment was quite effective in reducing variability in the oats variety test, while it increased variability in the oats strain test, which was conducted on the same field and similarly handled in every way. In fact, conditions were considered more favorable for the effectiveness of the practice in the strain test than in the variety test, for the check plots were closer together and 12 of the 15 strains tested were taxonomically identical with the check. But the standard deviation of check plots on the part of the field on which varieties were grown was almost twice as great as on the part of the field on which the strains were grown. Ap- parently the high variability of the plots in the variety test was caused in large part by differences in actual soil productivity which were largely counteracted by the adjustment of yields, while there was little variation in soil productivity in the strain test and such plot variability as occurred was largely due to other factors. In general therefore the adjustment of yields will probably be found more ef- 72 Missouri Agr. Exp. Sta. Research Bulletin 49 fective on fields highly variable in soil productivity than on more uniform fields, and for similar reasons the method will probably be found more effective in tests covering a rather large area than in tests covering a smaller area. It is clear that the adjustment of yields by means of check plots entails several serious disadvantages, and may increase experimental error considerably. Not only is the yield of the check plot a far from perfect measure of soil productivity for the check variety, but the pro- ductivity of the same soil for other varieties may be decidedly dif- ferent. The method is therefore more effective in tests of strains of the same variety as the check, than in tests of different varieties. When the yields of check plots are materially affected by factors not similarly affecting the neighboring test plots, adjustment of yields will increase experimental error. The check plots must therefore be effectively protected from competition, border effect, mechanical errors, and the like. Moreover, it is to be expected that the effectiveness of adjusting yields will vary with the season, since the relative influence of soil productivity on yield varies with the season. For example, in a season in which winter injury is exceptionally severe, actual soil fertility may have comparatively little to do with plot yields. Now, if the check variety is hardy, its yields may vary with the soil fertility, but when corresponding adjustments are made on the yields of tested varieties limited in yield by winter injury, a decrease in the variability of repli- cate plots is hardly to be expected. The same considerations apply of course to yields limited by many other factors. But, although a multitude of objections may be made to the theoretical bases of the practice of adjusting yields in variety tests, and although in many cases it undoubtedly results in an increase rather than a decrease in experimental error, the practice offers promise of value and is worthy of further investigation. The effectiveness of the adjustment of yields in the wheat variety test of 1920, in which the variability of replicate test plots was reduced about 40 per cent, is a demonstration of the possibilities of the method. An increase in replication of plots involving the same increase in land and labor would probably have reduced plot variability only about 7 per cent. A thor- ough knowledge of the value and limitations of yield-adjustment by means of check plots might enable us to reduce variability, at least in some types of plot tests, much more effectively by this means than by replication. The saving in area required is of particular significance in preliminary tests if border rows must be used for the elimination of competition, since in this case the area required for a large number of replications is in many cases prohibitive. Experiments in Fieed Plot Technic 73 CONCLUDING REMARKS The best method for preliminary variety testing is one which will permit the accurate determination of the relative value of the va- rieties under field conditions, with the use of a small area of land for each variety. Some precision must be sacrificed to save land, and in so far as the errors involved are of such nature that their extent can be approximately determined, and conclusions drawn accordingly, this sacrifice of precision is permissible. In many cases it is advisable, for example, to reduce the number of replications and to increase the least difference in yield regarded significant to a sufficient degree to com- pensate for the decrease in precision. But these considerations do not apply to systematic errors, which, since they affect the yields of replicate plots similarly, and consequently have little effect on plot variability, cannot be accurately measured. Typical systematic errors commonly involved in preliminary testing are (1) modification of growing conditions favoring some varieties more than others, such as hand planting or wide spacing between rows, and (2) competition between varieties of different type, resulting from the use of single-row plots. The relative value of varieties under such conditions may be vastly different from their relative value un- der typical field conditions. Even should measurable experimental error be reduced to the absolute minimum, such a variety test might give results entirely misleading. The error cannot be counteracted, as can non-systematic errors, by increasing the least difference con- sidered significant, nor can the extent of error of this sort be measured or estimated by a study of the experimental results. Systematic error must therefore be reduced by every practicable means. Growing conditions in the preliminary test- should be made as similar to ordinary field conditions as possible. The effect of varietal competition must be reduced to the minimum. If this can be ac- complished without increasing the size of plots, it is desirable to do so. On the other hand, if larger plots are necessary for the control of competition, larger plots should be used. If the area to be used for preliminary testing cannot be correspondingly increased, the num- ber of replications can be reduced sufficiently to permit the use of the larger plots required on the area available. This will necessitate a decrease in the degree of precision of the test, and will reduce the rapidity of elimination of the less valuable varieties. But is it not better to eliminate the undesirable varieties slowly than to risk the elimination of desirable ones by a more rapid analysis? 74 Missouri Agr. Exp. Sta. Research Bulletin 49 The error from competition is greater when different varieties are compared than when different strains of the same variety are compared, and the extent of error is roughly in proportion to the de- gree of difference in type of the varieties tested. Competition was not found to be correlated closely enough with earliness of heading, earliness of maturity, height, or grain-straw ratio in these experi- ments to permit its control by grouping varieties in respect to these characters. The factor found most closely correlated with competitive value was yield, but the correlation even in this case was not close enough to permit of effective control by grouping varieties. Moreover, it would be impossible in practice to group varieties with regard to yield, since the relative yield of varieties varies so widely with the season. The variety expected to yield poorly is not ordinarily included in the variety test. When different strains of the same variety are grown, the error from competition, in some cases at least, may be slight enough to justify the use of single-row plots. However, competition in such cases is not wholly absent, and may occasionally be quite marked. The importance of competition as a source of error in tests of pure line selections of the same variety merits detailed investigation. If it is found that the effects of competition between pure lines is slight it may be practicable to use single-row plots, or at any rate to use 3-row plots without discarding border rows. The latter method will reduce the error from competition materially, without necessitating the loss of any of the experimental area. When the same total area is used, however, single row plots are somewhat more reliable than 3-row plots, because more replications can be used. The best size of plot for ordinary variety testing, as indicated by this investigation, is probably the 3-row plot with border rows discarded. The length of the plot as harvested is assumed to be 16 feet, but the same considerations will apply for any other convenient length. The number of replications will vary with the heterogeneity of the field and the degree of precision required (and, to some extent, with the season and the variety). Check plots have been used in preliminary variety tests mainly for the following purposes: (1) For the adjustment of the yields of the test plots, and (2) To provide a measure of plot variability for the field used, and thus to determine the degree of precision of the experimental re- sults, or the number of replications which would be required for a given degree of precision. In both cases the behavior of the check variety is the basis for conclusions regarding the tested varieties. This involves the as- sumption that different varieties of the same crop respond similarly Experiments in Field Plot Technic 75 to varying conditions. In one case, reported in this paper, two stand- ard varieties, used as duplicate checks, and grown side by side in 120 distributed sections of a field, showed no significant correlation in relative yield of adjoining plots, and differed so widely in plot varia- bility that the number of replications necessary for a given degree of accuracy was more than twice as great for one check variety as for the other. Further investigation is necessary to determine how generally such cases may occur, but this single case indicates at least a possible source of extreme error in the use of check plots, either for adjust- ment of yield or for the determination of the probable error of the experimental results. For this and various other reasons the adjustment of yields by means of check plots is at present of doubtful value as a general prac- tice. In some cases, however, such adjustment accomplishes a great improvement in the precision of an experiment, with a relatively slight increase in expense. The practice is more promising for tests of strains or selections of the same variety than for tests of different types. A thorough study of the use of check plots in variety and strain testing may discover methods of overcoming the disadvantages, and thus make available an economical and effective method of increasing precision. Meanwhile, check plots should be used cautiously. Meth- ods for adjusting yields and for determining the extent of plot varia- bility without the use of check plots are available 17 ’ 18 , and check plots must demonstrate actual value if they are to continue in use in variety tests. SUMMARY 1. In variety and strain tests of barley, oats, and wheat, in five- row blocks, the competing border rows of adjacent sorts gave relative yields often widely different from those of the interior rows of the same plots. 2. Such competitive effects were much more extreme between different varieties than between different commercial strains of the same variety. 3. A considerable error from competition affected tests in rows running north and south, as well as those in rows running east and west. 4. Although in general the higher yielding varieties were favored in competition, the reverse frequently occurred. In some cases a ma- terial advantage in yield in the interior rows was converted to a material disadvantage in yield in the border rows. 76 Missouri Agr. Exp. Sta. Research Bulletin 49 5. Competing quality was correlated fairly consistently with yield and with earliness of heading and maturity. No relation to grain-straw ratio was found in the one season in which this charac- ter was determined. A significant correlation between competition and height was found in the wheat variety test of 1921, but the rela- tion of competition to height was not determined in the other tests. 6. In the oats tests competition was most closely related to earli- ness of heading and maturity, but was also related to yield. In the wheat, competition was related fairly closely to both yield and earliness. In the barley it was not significantly correlated with any of the char- acteristics studied, though the relation to yield was considerably closer than the relation to any of the other characteristics. 7. In the wheat and oats tests in which earliness and yield were correlated with competition, earliness and yield were correlated quite closely with one another. 8. Single-row plots, protected from competition by border rows discarded at harvesting, were somewhat more variable in yield than 3-row plots similarly protected, but the difference was not great enough to outweigh their advantage in size. The mean yield of five replicate protected single-row plots is therefore more reliable, under the conditions of these tests, than the mean yield of three replicate protected 3-row plots, which would occupy the same area and require considerably more labor in harvesting and threshing. 9. There was no consistent difference in variability between 3- row and 5-row plots. 10. Plot variability was increased with increase in the size of the experiment field. The number of replications required for a given degree of precision, as measured by the variability of plot yields, is therefore increased somewhat when border rows are added for the control of competition. 11. The variability of 120 distributed check plots of Kherson oats differed widely from that of 120 distributed plots of Red Rustproof oats, adjacent to them. If the variability of the check yields were con- sidered a measure of the precision of the test, entirely different con- clusions would be drawn on the basis of the yields of these two check varieties. 12. Adjustment of plot yields on the basis of the yields of check plots resulted in a decrease in plot variability in three tests and in an increase in five tests. In general the practice was effective on fields of high plot variability, and was ineffective on fields of low plot varia- bility. Experiments in Field Plot Technic 77 13. In the oats strain test in which both Kherson and Red Rust- proof check plots were included, the Kherson check was more effect- ive than the Red Rustproof check as a basis for adjusting the yields of the Kherson strains, while the Red Rustproof check was more ef- fective as a basis for adjusting the yields of the Red Rustproof strains. 14. The correlation between the yields of adjacent Kherson and Red Rustproof check plots was not statistically significant. Adjust- ment of the yields of the Kherson check plots on the basis of the yields of the adjacent Red Rustproof plots, and of those of the Red Rustproof plots on the basis of the Kherson yields increased va- riability. ACKNOWLEDGMENT The writer is indebted to Professors M. F. Miller and W. C. Etheridge for a critical reading of the manuscript, and to O. W. Letson for preparing figure 8. 78 Missouri Agr. Exp. Sta. Research Bulletin 49 REFERENCES CITED. 1. Day, James W. The relation of size, shape, and number of replications of plats to probable error in field experimentation. In Journ. Amer. Soc. Agron. 12, 3; pp. 100-105. 1920. 2. Etheridge, W. C. A classification of the varieties of cultivated oats. Cor- nell Univ. Agr. Expt. Sta. Memoir 10 ; pp. 85-172. 1916. 3. Hall, A. D. and E. J. Russell. Field trials and their interpretation. In Jour. Bd. Agr. (London) Supplement: pp. 5-14. 1911. 4. Hayes, H. K. and A. C. Arny. Experiments in field technic in rod-row tests. In Jour. Agr. Res., 11, 9: pp. 399-419. 1917. 5. Kiesselbach, T. A. Studies concerning the elimination of experimental error in comparative crop tests. Nebr. Agr. Expt. Sta. Res. Bui. 13: pp. 3-95. 1918. 6. Kiesselbach, T. A. Experimental error in field trials. In Journ. Amer. Soc. Agron. 11, 6: pp. 235-241. 1919. 7. Kiesselbach, T. A. Plat competition as a source of error in crop tests. In Journ. Amer. Soc. Agron. 11, 6 : pp. 242-247. 1919. 8. Love, H. H. The experimental error in field trials. In Journ. Amer. Soc. Agron. 11, 5: pp. 212-216. 1919. 9. Love, H. H. and W. T. Craig. Methods used and results obtained in cereal investigations at the Cornell Station. In Journ. Amer. Soc. Agron. io, 4: pp. 145-157. 1918. 10. Lyon, T. L. A comparison of the error in yield of wheat from plats and from single rows in multiple series. In Proc. Amer. Soc. Agron. 2: pp. 38, 39. 1911. 11. Lyon, T. L. Some experiments to estimate errors in field plat tests. In Proc. Amer. Soc. Agron. 3: pp. 89-114. 1912. 12. Mercer, W. B. and A. D. Hall. The experimental error in field trials. In Journ. Agr. Sci. 4, 2 : pp. 107-132. 1911. 13. Montgomery, E. G. Variation in yield and methods of arranging plats to secure comparative results. In 25th Ann. Rpt. Nebr. Agr. Expt. Sta. : pp. 164-180. 1911. 14. Montgomery, E. G. Experiments in wheat breeding. Experimental error in the nursery and variation in nitrogen and yield. U. S. Dept. Agr. Bur. Plant Indus. Bui. 269 : pp. 5-61. 1913. 15. Morgan, J. O. Some experiments to determine the uniformity of certain plats for field tests. In Proc. Amer. Soc. Agron. 1 : pp. 58-67. 1910. 16. Salmon, C. Check plats as a source of error in varietal tests. In Journ. Amer. Soc. Agron. 6, 3: pp. 128-131. 1914. 17. Surface, F. M. and Raymond Pearl. A method for correcting for soil heterogeneity in variety tests. In U. S. Dept. Agr. Journ. Agr. Res. 5, 22: pp. 1039-1049. 1916. 18. Wood, T. B. & F. J. M. Stratton. The interpretation of experimental results. In Journ. Agr. Sci. 3, 4: pp. 417-440. 1910. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 50 Certain Responses of Apple Trees to Nitrogen Applications of Different Kinds and at Different Seasons (Publication Authorized December 8, 1921) COLUMBIA, MISSOURI JANUARY, 1922 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. EANSING RAY, P. E. BURTON, H. J. BLANTON, St. Eouis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION J. C. JONES, PH. D., LL. D., PRESIDENT OF THE UNIVERSITY F. B. MUMFORD, M. S., DIRECTOR STATION STAFF January, 1922 AGRICULTURAL CHEMISTRY C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, A. M. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. SiEveking, B. S. in Agr. E. M. Cowan, A. M. AGRICULTURAL ENGINEERING J. C. Wooley, B. S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr, H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. E. F. Hopkins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale, B. S. in Agr. W. W. Swett, A. M. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B^S. in Agr. D. H. Nelson, B. S. in Agr. ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. R. McBride, B. S. in Agr. FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, A. M. O. W. Letson, A. M. B. M. King, B. S. in Agr. Alva C. Hill, B. S. in Agr. Miss Bertha Hite, A. B.* Seed Analyst Miss Pearl Drummond, A. A.* RURAL LIFE O. R. Johnson, A. M. S. D. Gromer, A. M. E. L. Morgan, A. M. B. H. Frame, B. S. in Agr. HORTICULTURE V. R. Gardner, M. S. A. H. D. Hooker, Jr., Ph. D. J. T. Rosa, Jr., M. S. F. C. Bradford, M. S. H. G. Swartwout, B. S. in Agr. POULTRY HUSBANDRY H. L- Kempster, B. S. Earl W. Henderson, B. S. SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M. W. A. Albrecht, Ph. D. F. L. Duley, A. M.t R. R. Hudelson, A. M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr. Richard Bradfield, A. B. O. B. Price, B. S. in Agr. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Secretary Sam B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsham, Librarian E. E. Brown, Business Manager *In the service of U. S. Department of Agriculture. tOn leave of absence. Certain Responses of Apple Trees to Nitrogen Applications of Different Kinds and at Different Seasons. H. D. Hooker, Jr. A careful study of the literature dealing with fertilizer appli- cations to fruit plants shows that those most commonly effective involve nitrogen. Other elements have been shown to be beneficial in many cases when applied to fruit plants, but not so consistently nor so strikingly. The relatively distinct effects that follow occa- sional applications of iron constitute an exception to the general statement, but only because the symptoms of iron deficiency are easily recognized and generally understood. For this reason, iron is seldom used as a fertilizer except where its requirement is clearly indicated by chlorosis. Were it possible to tell as readily when the other essential mineral elements become limiting factors of growth and production, the problem of fertilizer requirements would be relatively simple. In the absence of any symptoms more marked than a pale green color of the leaves or a small amount of new growth — conditions that may result from any one of a number of different causes — it is inevitable that many fertilizer applications should be without important effect, for they meet no requirement. On the whole, it is remarkable that nitrogen applications should be effective in increasing growth and crop producing power as generally as they do, a fact which indicates that nitrogen is more often the limiting factor of growth and yield in fruit trees than any of the essential mineral elements. Nitrogenous fertilizers applied to fruit trees have quite general- ly increased the set of fruit, favored vegetative growth and increas- ed yields. The experimental work by which these facts have been de- termined has been for the most part empirical. The orchard ferti- lizer problem has been attacked as a simple matter of generally in- creasing growth and productiveness. These are, to be sure, the objects of ultimate interest and of greatest importance, but the fertilizer problem is more complex. The response in terms of growth and yields is the culmination of many different activities — absorption, elaboration, utilization and storage — of correlative ef- fects on other constituents, of distinct processes of growth, fruit bud differentiation, fruit setting and development, each of which is conditioned by different factors or sets of factors. For example: 4 Missouri Agr. Exp. Sta. Research Bulletin 50 Have the increased yields following the application of nitrogenous fertilizers resulted only from the better set of fruit and the in- creased size of the tree, or have nitrogen applications had a direct or indirect influence on fruit bud differentiation? Little attention has been given the effects of fertilizer applications made at times other than early spring. Is the season of application which is best for increasing the set of fruit likewise best for increasing fruit bud differentiation? The possible advantages to be derived from ferti- lizer applications evidently have not been exhausted. The fertilizer problem should be studied as a problem in nu- trition and the effects of nitrogen applications should be measured in terms of the chemical changes produced in various parts of the plant as well as in terms of growth and yield. Only by this method can principles of more or less general significance be determined. The following questions present themselves: Is the nitrogen con- tent of a fruit tree increased by nitrogen applications? Does one form of nitrogenous fertilizer produce a greater immediate effect than another? Is the content of other constituents, particularly var- ious forms of carbohydrate, altered? Are significant changes evident at the time of fruit bud differentiation and are different effects produced by applications at different seasons? In the hope of throwing light on some of these questions, the present investiga- tion was begun in the spring of 1920. MATERIALS AND METHODS Through the kindness o f Dr. J. C. Jones, President of the University of Missouri, two apple orchards at McBaine, Mis- souri, were placed at the disposal of the Department of Horticul- ture for experimental treatment and sampling. Each orchard con- tained York trees in good condition ; those in the south orchard were 20 years old and bore in even years; those in the north or- chard were 16 years old and bore in odd years. These trees were fertilized in a manner to be described presently and samples of spurs and of bark from the scaffold limbs, four to six inches in diameter, were collected at intervals during the year. The bark samples con- sisted of strips three-quarters of an inch wide and six to eight inches long, pointed at each end and including all tissue outside of the cambium. Not more than three strips were taken from a sin- gle tree at one sampling. The wounds were covered with white lead paint to prevent infection and undue water loss. The spur samples included 1919 and 1920 growth. Responses of Apple Trees to Nitrogen Applications 5 The other trees used for experimental purposes were at the Experiment Station Fruit Farm, Turner Station, Mo. Samples of spurs including 1919 and 1920 growth were collected from ferti- lized and check trees of Jonathan and Ben Davis varieties growing in bottom land and seven years old in 1920, when the samples were collected. Another fertilizer experiment was conducted on vigor- ous four-year-old Grimes trees at the Fruit Farm. Samples of spurs of various lengths were collected from mature Ben Davis trees growing in the University orchard at Columbia. The chemical analyses were made after the manner detailed in Research Bulletin 40 of this Station. Determinations were made of dry weight, ash, potassium, phosphorus, nitrogen, reducing sug- ars, total sugars and starch. In the spring of 1921 practically all blossoms in the orchards under study were killed by late frosts. Since it was impossible to determine the effects of all the fertilizer treatments as originally planned, some of the treatments were repeated in 1921. However, a large body of data had been collected which it seems advisable to publish, incomplete and fragmentary though it be, since it shows some significant and rather unexpected facts. SPRING APPLICATIONS OF VARIOUS NITROGENOUS FERTILIZERS This first experiment was made on York trees in their bearing year. Four plots of 15 trees each were selected ; one was left as a check; another received 5 pounds of sodium nitrate per tree; an- other received 3 pounds of ammonium sulphate per tree and the other 5 pounds of a high grade of dried blood. By this treatment each fertilized tree received approximately the same amount of nitrogen. The applications were made March 19, 1920. The greater crop produced by the fertilized plots was the most striking effect produced. The yields from the plots were as fol- lows : 285 bushels from the check plot or 19.7 bushels per tree; 375 bushels from the blood plot or 25.0 bushels per tree; 381 bush- els from the nitrate plot or 25.4 bushels per tree; 376 bushels from the ammonium sulphate plot or 25.1 bushels per tree. These fig- ures show that the three types of fertilizers used produced practi- cally the same effect in increasing yield. There were minor vari- ations within the plots: one tree on the blood plot produced 40 bushels of apples ; one tree on the ammonium sulphate plot bore so large a crop that the tree split to the ground under the weight of 6 Missouri Agr. Exp. Sta. Research Bulletin 50 fruit; the yields of the trees on the nitrate plot were more nearly uniform. No definite figures are available but the apples from the nitrate plot seemed to be slightly larger and somewhat less highly colored than those from the other plots. The data in Table 1 show that neither the bearing nor the non- bearing spurs of the fertilized trees formed a larger number of leaves during the current season than did corresponding spurs on the unfertilized trees. The following spring there was no bloom on any of the plots. The chief effects of the fertilizer treatments observed were a deeper color of the foliage and an increased set of Table 1. — Average Number oe Leaves Per Spur on Fertilized and Unferti- lized York Trees. May 3 May 15 May 22 Bearing spurs Check plot _ . 7.0 9.1 9.25 Nitrate plot — 4.8 7.2 7.34 Amm. sulphate plot _ 6.8 8.4 8.6 Blood plot 5.8 7.4 7.9 Non-Bearing spurs Check plot 6.6 8.0 8.2 Nitrate plot 5.2 7.2 7.4 Amm. sulphate plot __ 6.0 7.4 7.6 Blood plot 5.7 7.1 7.4 fruit: 23.7 percent of the blossoming spurs on the check trees bore fruit and approximately 32 percent of the blossoming spurs on the fertilized trees, as determined by actual count. Samples of spurs and of bark were collected from the trees on these plots at five dates; May 22, June 19, September 6, Novem- ber 20, 1920, and April 2, 1921. The analytical determinations on these samples are given in Table 2. (See pages 8 and 9.) The greater set of fruit on the fertilized trees is probably to be associated with the greater nitrogen content of their spurs on May 22, as suggested by Harvey and Murneek (Ore. Agr. Expt. Sta. Bui. 176). At this time the nitrogen content of the spurs is on the decline, as reference to Figure 10 in Research Bulletin 40 of this Station shows. Except for this temporary increase in nitrogen content there is no very significant difference between the composition of the fertilized and unfertilized trees. The absence of such variations Responses of Apple Trees to Nitrogen Applications 7 is striking. There has been no marked starch accumulation in any of the spurs by June 19, immediately before the period of fruit bud differentiation. Moreover, on April 2 the following spring, the spurs of the fertilized trees contained very little more nitrogen than the check spurs, the difference found being well within the limits of experimental error. Similarly, though the nitrogen content of the bark from the nitrated trees is slightly greater than that of the check trees, the sulphate and blood trees have less. Consequently no consistent difference is evident. The comparison between the chemical composition of spurs and bark from the same trees as afforded by the data presented is of considerable interest. In general, the variations in the chemical composition of the bark follow rather closely those in the spurs. The low starch content of the former in May and June is particu- larly striking as it indicates that the factors which prevent carbo- hydrate accumulation in the bearing spurs likewise affect the bark of the scaffold limbs. The high total sugar content of the bark during the winter and especially in April is apparently character- istic. The bark contains, during most of the year, about half the percentage of nitrogen that the spurs contain ; its percentage phos- phorus content is also, for the most part, much less but the per- centage potassium content of these two portions of the tree is of the same order and during part of the year the bark contains an even higher percentage than the spurs. This is consistently true in the June and September analyses. The percentage ash content of bark is usually greater than that of the spurs though the difference is very nearly wiped out in the spring. In this experiment the various types of nitrogenous fertilizers have produced essentially the same effects as shown both by the chemical analyses and the crop yields. This effect has consisted principally in an increased set. There has been no effect in in- creasing the number of leaves during the current season and very little effect, if any, on the rate of growth. There has been no effect on fruit bud differentiation nor any tendency in that direction, such as might be evidenced by an accumulation of starch in the spurs during the period of fruit bud differentiation. In fact, the greater carbohydrate utilization following the increased set of fruit would tend to decrease the chances for starch accumulation in the ab- sence of an increased leaf area per spur. Nitrogenous fertilizers should, therefore, be applied to bien- nially bearing apple trees in the spring of their crop year with ex- 8 Missouri Agr. Exp. Sta. Research Bulletin 50 treme caution, for such a practice is likely to lead to overproduc- tion with its attendant evils. Table 2. — Analyses oe Spurs and Baric From Fertilized and Unfertilized York Trees in Their Bearing Year. ( Percentages of dry weight) Dry weight Reduc’g sugars Total sugars Starch Ash K P N May 22, 1920 Bearing spurs Check 40.2 0.79 1.95 0.68 9.73 0.678 0.170 1.020 Nitrate 43.2 1.00 1.65 1.15 8.74 0.626 0.162 1.047 Sulphate __ _ 43.6 0.66 1.59 0.00 11.32 0.649 0.138 1.164 Blood 41.2 0.70 2.79 0.45 7.69 0.665 0.174 1.221 Bark Check 43.4 1.48 1.74 0.11 7.97 0.621 0.060 0.578 Nitrate 43.1 1.61 1.89 0.20 12.79 0.648 0.064 0.565 Sulphate 42.7 1.69 1.74 0.00 8.12 0.585 0.072 0.605 Blood 42.3 1.53 2.10 0.14 8.32 0.513 0.075 0.526 June 19, 1920 Bearing spurs Check _ 44.4 1.26 1.65 0.00 6.44 0.554 0.140 0.916 Nitrate 45.7 1.26 1.29 0.00 6.59 0.362 0.141 0.912 Sulphate _ _ 42.5 0.77 1.08 0.36 6.02 0.639 0.148 1.024 Blood _ 43.1 0.86 1.20 0.00 7.21 0.558 0.209 1.166 Bark Check 47.9 1.32 1.62 0.00 9.91 0.570 0.145 0.578 Nitrate 42.3 1.71 2.00 0.38 8.41 0.583 0.108 0.546 Sulphate 42.6 1.62 1.89 0.70 8.11 0.647 0.086 0.575 Blood 48.1 1.73 2.04 0.54 7.77 0.522 0.096 0.551 September 6, 1920 Bearing spurs Check _ 46.7 1.21 2.25 2.88 7.97 0.408 0.155 1.030 Nitrate 45.6 0.88 1.38 1.98 7.29 0.371 0.164 0.880 Sulphate _ _ 45.3 0.72 1.14 2.41 7.29 0.420 0.211 1.110 Blood 48.8 0.72 1.20 1.98 6.58 0.414 0.176 1.010 Bark Check 44.9 1.87 2.70 2.53 10.76 0.457 0.110 0.52 Nitrate 40.6 1.72 2.60 2.27 11.45 0.411 0.107 0.44 Sulphate 43.6 1.65 2.55 2.36 11.08 0.502 0.108 0.58 Blood 44.6 1.45 1.86 2.66 12.00 0.459 0.136 0.54 November 20, 1920 Spurs Check 48.4 2.95 3.15 1.08 9.05 0.489 0.222 1.22 Nitrate 47.9 1.98 2.73 1.19 8.01 0.457 0.230 1.14 Sulphate _ 49.6 1.94 2.71 1.28 8.53 0.473 0.177 1.09 Blood 48.5 1.73 2.16 1.15 8.69 0.513 0.241 1.20 Responses of Apple Trees to Nitrogen Applications 9 Table 2. — (Continued.) Dry weight Reduc’g sugars Total sugars Starch Ash K P N Bark Check 44.1 2.68 3.78 1.58 12.35 0.498 0.098 0.69 Nitrate __ 45.3 3.07 4.32 1.72 11.48 0.393 0.102 0.61 Sulphate — 46.9 2.81 3.42 1.58 12.33 0.385 0.075 0.64 Blood 44.1 2.38 3.27 1.98 11.43 0.397 0.106 0.58 April 2 , 1921 Spurs Check 48.5 2.44 3.06 1.53 9.28 0.498 0.210 0.86 Nitrate 46.9 2.36 3.37 2.34 9.71 0.428 0.212 0.92 Sulphate 47.2 2.07 3.21 1.35 9.81 0.441 0.183 0.89 Blood 44.7 2.47 3.45 1.71 10.11 0.490 0.202 0.88 Bark Check 46.1 5.57 6.56 2.55 9.22 0.405 0.122 0.58 Nitrate 46.7 5.93 6.36 2.52 10.63 0.354 0.104 0.62 Sulphate _ _ 46.4 5.68 5.94 2.34 10.19 0.346 0.147 0.52 Blood 47.1 5.79 6.00 2.43 9.63 0.430 0.144 0.56 EFFECT OF SPRING APPLICATIONS OF NITRATE IN PROMOTING GROWTH Spring applications of nitrogenous fertilizers not only have an effect on the setting of fruit, but, as is well known, they frequently increase the amount of growth. This effect is shown by an experi- ment on Ben Davis and Jonathan trees at the University Fruit Farm. Two trees of each variety were treated with three pounds of sodium nitrate in the spring of 1919 and again March 29, 1920. The effects of this treatment have been revealed in the greater size of the fertilized trees as compared with check trees of the same variety in the same rows. These trees blossomed for the first time in 1921 but the entire bloom was killed by spring frost. Samples of spurs were collected from the fertilized trees and the checks at three dates, March 29, May 22 and June 19, 1920. The analyses of these spurs are given in Table 3. The percentage nitrogen content of the spurs from the ferti- lized trees was less on March 29 than that of the check spurs, show- ing that the fertilizer applied the year before did not increase the nitrogen content of the spurs. In May, the fertilized spurs con- tained a higher percentage of nitrogen than the checks ; but since their percentage nitrogen content continued to decline during the month following, while that of the checks increased, the fertilized 10 Missouri Agr. Exp. Sta. Research Bulletin 50 Table 3. — Analyses oe Spurs on Fertilized and Unfertilized Jonathan and Ben Davis Trees. ( Percentages of dry weight ) Dry weight Reduc’g sugars Total sugars Starch Ash K P N March 29, 1920 Jonathan Nitrated 1.64 1.92 1.35 5.315 0.489 0.193 1.23 Check 1.71 1.77 1.08 6.480 0.558 0.220 1.35 Ben Davis Check 1.98 2.13 0.47 8.020 0.414 0.190 1.30 Nitrated — 1.31 2.25 0.72 7.925 0.402 0.200 1.245 May 22, 1920 Jonathan Check 42.9 1.04 1.88 1.01 4.76 0.722 0.131 0.857 Nitrated 40.9 1.22 2.34 0.00 4.70 0.423 0.127 0.927 Ben Davis Check 44.1 1.04 1.65 0.36 4.59 0.514 0.100 0.699 Nitrated 43.9 1.01 1.77 0.90 4.33 0.480 0.103 0.822 June 19, 1920 Jonathan Check 45.1 1.08 2.67 2.16 4.64 0.457 0.157 0.897 Nitrated __ . _ 47.1 0.70 1.86 1.60 4.51 0.455 0.144 0.820 Ben Davis Check 49.3 1.08 1.38 1.42 5.94 0.325 0.224 1.128 Nitrated 47.6 0.65 1.35 1.10 4.60 0.321 0.134 0.792 spurs again contained less than the checks on June 19. This slight difference in itself may not be particularly significant at this time, immediately preceding the period of fruit bud differentiation, but when considered in its relation with other conditions with which it is associated and for which it may possibly be responsible, it may assume great importance. At this time the starch content in the spurs of the fertilized trees was distinctly less than in the check spurs. This indicates clearly that the conditions for fruit bud dif- ferentiation were not improved and in fact were made less favor- able by the spring application of nitrate of soda. The smaller ac- cumulation of starch in the spurs of fertilized trees immediately before the period of fruit bud differentiation is probably related to, if not actually caused by, the more vigorous growth of the ferti- lized trees. It is evident that the minimum nitrogen content occurred much sooner in the spurs of the check trees than in those of the Responses of Apple Trees to Nitrogen Applications 11 fertilized trees and this minimum is related to the time of growth cessation, for an accumulation of nitrogen as shown by an in- creased percentage does not usually occur in spurs until growth has ceased. The figures in Table 3 show that the potasssium and total ash content of the fertilized spurs is consistently less than in the check spurs and for the most part this is true also of the phosphorus content. These conditions may be interpreted as further conse- quences of the more vigorous growth of the fertilized trees for it is a general rule that the greater the length of the spur growth, the low- er is its ash content at the close of the growing season. This is shown by the data in Table 4 which are analyses of Ben Davis spurs collected May 21 and July 2, 1920. The samples were collected ac- cording to spur lengths as shown in this table. It will be seen that the ash, phosphorus and nitrogen content is less, the longer the spur growth of the current season. Table 4. — Analyses oe Ben Davis Spurs According to Length (1919 and 1920 wood included) ( Percentages of dry weight ) Spur length Dry Ash KPN in centimeters weight May 21, 1920 0.5- 1.0 48.8 1.1- 1.5 45.6 1.6- 3.0. 41.2 3.1- 10.0 36.9 July 2, 1920 0.5- 1.0 48.4 1.1- 1.5 47.3 1.6- 3.0 46.9 3.1- 10.0 46.8 6.15 0.597 0.179 0.891 5.28 0.568 0.177 0.876 4.91 0.536 0.137 0.705 4.49 0.572 0.133 0.700 6.44 0.513 0.142 0.754 5.53 0.531 0.134 0.754 5.04 0.527 0.138 0.726 4.18 0.475 0.118 0.684 THE EFFECT OF SPRING APPLICATIONS ON IMMA- TURE TREES On March 29, 1920, two plots of young Grimes apples were fertilized, one with 2 pounds of nitrate of soda and one with 2 pounds of dried blood to the tree. Each plot contained 10 trees and a similar block of 10 trees was left as a check. About 50 short, spur-like growths and 13 leaders on the trees of each plot were labeled. Growth measurements were made on these spur-like growths and on the four shoots arising from the terminal portion 12 Missouri Agr. Exp. Sta. Research Bulletin 50 of each leader. The effects of the fertilizer treatments on the growth of these trees is shown in Table 5. It is evident that the ni- trate of soda had a greater effect on the shoot growth than did the dried blood. Practically no difference is evident between the rates of growth from the short spur-like branches. Chemical analyses of these two types of growth are shown in Table 6. The absence of any marked differences between the va- rious plots is quite striking. Differences do appear, however, in the chemical composition of the terminal growth as compared with that of the shorter spur-like branches. In the former there is no complete disappearance of starch in June and the nitrogen content Table 5. — Average Lengths in Centimeters oe Growths From Leaders and Spur-Like Branches on Four-Year-Oed Grimes. May 5 May 14 May 21 May 31 June 14 July 9 Leader growth Check 2.24 8.5 12.2 19.5 28.0 35.8 Nitrate 2.23 9.8 14.7 22.5 31.9 42.6 Blood 2.18 8.7 12.5 19.0 27.5 34.6 Twig growth Check _ 2.4 7.2 9.4 13.0 15.3 16.1 Nitrate 2.2 7.5 10.6 13.9 15.6 16.0 Blood _ 2.7 9.3 10.4 Average number oe leaves on twig growth 13.3 15.8 16.8 Check _ _ 6.0 9.3 10.0 12.0 13.9 14.2 Nitrate 7.2 8.9 10.1 11.8 12.9 13.1 Blood 7.5 9.8 10.8 12.3 13.5 14.6 rises to a high maximum in May, much as in bearing spurs on ma- ture trees. The only apparent difference associated with the greater ter- minal shoot growth of the nitrated trees is shown in a higher per- centage of starch and total sugar and a lower percentage of ash, potassium and phosphorus in June. In September these shoots have the highest percentage of ash and potassium and the lowest of phosphorus and nitrogen. The data presented indicate a differential effect between ni- trate of soda and dried blood, which may be associated with the more quickly available character of the former. Different parts of a tree evidently may react in different ways to the same fertilizer treatment. A comparison of the responses of these four-year-old trees with those of the Ben Davis, Jonathan and York trees indi- Responses of Apple Trees to Nitrogen Applications 13 Table 6. — Analyses oe Growths From Leaders and Spur-Like Branches on Four- Year-Old Grimes. ( Percentages of dry weight ) Dry weight Reduc’g sugars Total sugars Starch Ash K P N May 21, 1920 Leader growth Check 25.8 0.97 1.17 2.09 4.49 1.435 0.234 1.625 Nitrate 25.1 0.77 1.35 2.05 4.61 0.669 0.236 1.625 Blood 25.2 1.01 1.26 2.70 4.56 1.011 0.216 1.615 Spur-like growth Check 31.0 0.86 1.44 1.96 4.63 0.599 0.204 1.299 Nitrate 33.6 0.90 1.08 1.51 4.17 0.706 0.152 1.045 Blood 33.0 1.08 1.08 1.58 4.26 0.579 0.181 1.150 June 19, 1920 Leader growth C heck 38.7 0.88 1.62 2.18 3.64 0.791 0.147 0.845 Nitrate 38.5 1.10 1.95 2.75 3.08 0.642 0.137 0.952 Blood 38.7 1.28 1.62 2.12 3.54 0.697 0.142 1.063 Spur-like growth Check 46.8 0.95 1.41 0.02 4.20 0.578 0.163 0.739 Nitrate _ 44.0 0.90 1.47 0.00 4.19 0.689 0.183 0.829 Blood _ 46.8 1.13 1.72 0.00 4.00 0.299 0.145 0.675 September 4, 1920 Leader growth Check _ _ __ 39.0 0.54 0.90 1.40 3.08 | ; 0.319 0.177 0.76 Nitrate 39.6 0.67 1.05 1.44 3.55 1 0.685 0.150 0.64 Blood 39.4 0.29 0.60 1.73 3.23 ! 0.554 0.153 0.66 Spur-like growth C heck 53.0 0.50 0.75 2.30 5.3i 0.433 0.189 0.75 Nitrate 53.0 0.81 1.23 1.89 4.98 0.411 0.197 0.73 Blood 54.3 0.68 1.23 2.05 6.01 0.406 0.196 0.81 cates that the age of the tree may be an important factor, although it is quite possible that the observed differences may have been due to the conditions under which the various trees were growing. THE EFFECT OF APPLICATIONS AT DIFFERENT SEASONS Fertilizer applications were made on 16-year-old York trees in their off year. One plot of 15 trees was fertilized March 29 with dried blood and another on June 20. Both times 5 pounds of fer- tilizer were applied to each tree. On September 20 another plot of 15 trees was given 5 pounds of sodium nitrate to the tree. Be- 14 Missouri Agr. Exp. Sta. Research Bulletin 50 cause of the severe spring frosts of 1921, the effects on these trees could not be measured except in terms of the chemical changes ob- served in the bark and spurs. These are given in Table 7. The data afford a comparison with Table 1 of the chemical composition of bark on bearing and non-bearing trees. In gener- al the seasonal changes in the chemical composition of the bark on the scaffold limbs of these alternate bearing trees follow those of the spurs. In June there is already an accumulation of carbo- hydrate in the. form of starch. In May the nitrogen content is somewhat less in the non-bearing trees. During the winter the sugar content of the bark is exceptionally high, much higher in the non-bearing than in the bearing trees. From September through March the potassium content of the non-bearing trees is distinctly higher and the phosphorus and nitrogen content like- wise, though to a lesser extent. The differential effects from applying these fertilizers at va- rious seasons is most evident in the nitrogen content of the spurs. On March 30, 1921, the nitrogen content of the spurs varies with the lateness of application the previous season. The check spurs have the least, the spring fertilized trees next, the summer ferti- lized trees more and the fall fertilized trees most of all. Moreover these differences in nitrogen content are by no means insignificant. It is unfortunate that weather conditions made it impossible to follow the later effects associated with these differences in the ni- trogen content of the spurs. It is impossible to say what advan- tage might accrue from increasing the nitrogen content of spurs in the spring of their bearing year but it is clear that late sum- mer or early fall applications of nitrogenous fertilizers are much more effective in this respect than spring applications. The effect of spring application of blood on these non-bearing trees is essentially the same as that observed on the Ben Davis and Jonathan trees. The accumulation of starch in the spurs to- ward the end of June at the critical time of fruit bud differentia- tion is reduced by the spring application of dried blood just as it was by the nitrate of soda in the Ben Davis and Jonathan spurs. Moreover, the residual effect of the spring fertilizer, as shown by the analyses of March 30, 1921, is practically nil, as in the case of the bearing York trees. The most marked effect on the nitrogen content of the bark was produced by the summer application of dried blood. In Sep- tember and December the nitrogen content of the bark on the sum- Responses of Apple Trees to Nitrogen Applications 15 Table 7. Analyses oe Spurs and Bark From Fertilized and Unfertilized York Trees in Their Oee Year. ( Percentages of dry weight ) Dry weight Reduc’g sugars Total sugars Starch Ash K P N May 27, 1920 Spurs Check _ 45.6 1.17 1.50 0.92 8.98 0.593 0.123 0.773 Blood 44.3 1.30 1.56 1.13 8.99 0.659 0.125 0.800 Bark Check 42.1 1.38 1.77 0.72 11.62 0.601 0.083 0.555 Blood 42.5 1.51 1.59 0.72 11.80 0.503 0.064 0.576 June 24, 1920 Spurs Check 49.4 0.92 1.38 2.88 6.665 0.516 0.227 0.960 Blood 48.1 0.83 1.56 1.87 5.818 0.615 0.142 0.706 Bark Check 43.5 0.94 2.30 7.845 0.558 0.084 0.550 Blood 43.8 1.10 1.89 2.00 7.640 0.540 0.144 0.721 September 20, 1920 Spurs Check 53.9 0.79 1.24 2.75 10.38 0.445 0.202 0.95 Spring blood __ 55.6 0.61 0.90 2.48 8.66 0.394 0.182 0.90 Summer blood 56.4 0.61 1.30 1.69 11.25 0.461 0.217 1.04 Bark Check 48.1 0.90 1.08 3.19 8.87 0.579 0.113 0.58 Spring blood __ 44.8 0.70 1.02 2.23 9.56 0.603 0.079 0.52 Summer blood 45.6 0.79 0.99 2.25 9.05 0.658 0.117 0.66 December 3, 1920 Spurs Check 44.7 2.78 2.79 1.51 8.85 0.539 0.233 1.09 Spring blood 46.1 2.90 3.00 2.09 10.17 0.437 0.214 1.17 Summer blood 46.0 3.05 3.30 2.23 9.96 0.461 0.240 1.19 Fall nitrate 46.8 2.32 2.70 2.18 8.82 0.450 0.253 1.33 Bark Check 53.8 4.22 5.70 1.91 10.28 0.587 0.110 0.72 Spring blood __ 53.4 5.17 5.58 1.13 10.14 0.626 0.106 0.72 Summer blood 54.6 4.57 4.98 1.93 9.84 0.659 0.123 0.91 Fall nitrate 54.4 5.02 5.22 1.22 9.82 0.595 0.159 0.77 March 30, 1921 Spurs Check 48.1 1.46 1.83 0.81 11.90 0.531 0.181 0.85 Spring blood 46.3 1.40 1.74 0.77 10.24 0.525 0.223 0.92 Summer blood 45.2 1.26 1.59 0.45 10.72 0.539 0.242 1.01 Fall nitrate 46.2 1.19 1.29 0.88 9.89 0.438 0.194 1.17 Bark Check 47.3 4.86 5.73 1.40 8.82 0.624 0.171 0.65 Spring blood __ 47.2 4.48 5.16 2.79 8.78 0.666 0.139 0.68 Summer blood 48.3 4.73 5.13 2.14 8.73 0.654 0.137 0.65 Fall nitrate 47.1 4.00 4.77 3.58 8.74 0.656 0.155 0.70 16 Missouri Agr. Exp. Sta. Research Bulletin 50 mer-fertilized trees was distinctly higher than that on the others but this difference had completely disappeared by the end of March, 1921. It is interesting that the nitrogen content of the bark on the spring-fertilized trees was apparently increased at the end of June while the nitrogen content of the spurs. was much less at that time. DISCUSSION The analyses reported in this paper confirm and extend those published in Research Bulletin 40 of this Station. Certain season- al chemical changes characteristic of bearing and of non-bearing apple spurs and certain changes in bark and spurs characteristic of apple trees in their on and off years — the years of fruit production and those of fruit bud differentiation respectively — may be consid- ered established. Particular physiological processes are apparent- ly associated with definite chemical conditions existing in the spur at critical times. Fruit bud differentiation, for example, is asso- ciated with starch accumulation in the spur late in June and, since no exception has been found to this rule, it seems safe to conclude that the conditions which bring starch accumulation about are among the factors that determine fruit bud differentiation, though the existence of other factors which are at times decisive is un- questionable. In order to account for the known facts relative to the initiation of the fruitful state it seems necessary to postulate two conditions for fruit bud differentiation: (1) that carbohydrate (or starch) accumulation be possible and (2) that no other factor such as nitrate, water or heat supply be limiting to the extent that vegetative development is stopped or seriously retarded. These points are discussed in Research Bulletin 47 of this Station, where it is also shown that the conditions determining starch accumula- tion and fruit bud differentiation are not always confined to the spurs. In a similar way other processes are found to be associated with particular features of the seasonal chemical picture. Thus recent investigators have pointed out that fruit setting seems to be related to the nitrogen content of the spur in May and vegeta- tive extension is evidently related to a number of factors. The data presented in this paper also show some of the effects of applying nitrogenous fertilizers of various kinds and at different seasons with special reference to the chemical composition of bark and spurs. These effects depend primarily on the condition of the Responses of Apple Trees to Nitrogen Applications 17 tree and might well be influenced also by climatic conditions. All the work reported here deals with apple trees in fairly good con- dition. The type of nitrogenous fertilizer is evidently important under certain circumstances; under others it is not. Various physi- ological processes are affected more or less independently and there are indications that the season when the applications are made is an important factor in determining how these processes are affected. An intelligent use of fertilizers evidently must be based on a recognition of the particular process which it is advisable to con- trol and on a knowledge of the effects that applications of various kinds and at different seasons will have on this process. In the past when nitrogenous fertilizers have appeared necessary, a suit- able amount has been determined on and has been applied in the cheapest or most readily available form in early spring. This pre- cedure is inadequate as a panacea and the facts presented show that it may produce effects directly opposite to those desired. Trees that bear light crops regularly every year and in whose an- nual yield a material increase is desired present a case for treat- ment as different from that of trees which bear heavy crops bien- nially and which it is desired to make regular producers as this in turn is different from the case of trees which bloom profusely every spring but set little or no crop because nitrogen is a lim- iting factor. In one case it may be a question of stimulating general vegetative growth and vigor; in another of affecting fruit bud differentiation ; in another of increasing the set of fruit. The same treatment will not bring about the desired result in all cases. Each is a problem for separate consideration and each involves phases which should be studied under a wide variety of conditions. There are, of course, limits to the effectiveness of nitrogenous fer- tilizers, but even where a requirement for nitrogen can be estab- lished, the best method of application in one instance may be, en- tirely different from the best for some other case. The effect o ( early spring applications of quickly available nitrogenous fertilizers in aiding the set of fruit has been established and evidence is giv- en that fruit bud differentiation and vegetative development also can be influenced in specific directions according to the time and type of application. 18 Missouri Agr. Exp. Sta. Research Bulletin 50 CONCLUSION The chief effects of spring applications of nitrogenous ferti- lizers to healthy apple trees are, on bearing trees, an increased set of fruit associated with a greater nitrogen content in the spurs during the period of fruit setting and in non-bearing trees an in- creased rate of growth. Different types of quickly available ni- trogenous fertilizers produce essentially the same effects though nitrate of soda stimulated leader growth on very young trees more than dried blood. Spring applications of nitrogenous fertilizers do not favor starch accumulation at the period of fruit bud differen- tiation and consequently they could not be expected to favor this process. No effects of spring applications are evident in the percentage nitrogen content the spring following the treatment, though larger absolute amounts would be present in the larger trees. The later in the season nitrogenous fertilizers are applied, the greater is the nitrogen content of the spurs the following spring immediately before growth begins. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 51 The Influence of the Plane of Nutrition On the Maintenance Requirement of Cattle (Publication Authorized November 21, 1921.) COLUMBIA, MISSOURI FEBRUARY, 1922 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL. THE CURATORS OF THE UNIVERSITY OF MISSOtTE EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY P. E. BURTON . j. BLANTON St. Louis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION J. C. JONES, PH. D., LL. D., PRESIDENT OF THE UNIVERSITY F. B. MUMFORD, M. S., DIRECTOR STATION STAFF February, 1922 AGRICULTURAL CHEMISTRY C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, A. M. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. SievEking, B. S. in Agr. E. M. Cowan, A.M. AGRICULTURAL ENGINEERING J. C. WoolEy, B. S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. E. F. Hopkins, Ph. D. DAIRY HUSBANDRY C. Ragsdale, B. S. in Agr. . W. Swett, A. M. m. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. R. McBride, B. S. in A. FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, A. M. O. W. Letson, A. M. B. M. King, B. S. in Agr. Alva C. Hill, B. S. in Agr. Miss Bertha C. Hite, A.B.* Seed Analyst. Miss Pearl Drummond, A. A.* RURAL LIFE O. R. Johnson, A. M. S. D. Gromer, A. M. E. L. Morgan, A.M. Ben H. Frame, B. S. in Agr. HORTICULTURE V. R. Gardner, M. S. A. H. D. Hooker, Jr., Ph. D. J. T. Rosa, Jr., M. S. F. C. Bradford, M. S. H. G. Swartwout, B. S. in Agr. POULTRY HUSBANDRY H. L- Kempster, B. S. Earl W. Henderson, B.S. SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M W. A. Albrecht, Ph. D. F. L. Duley, A.M.t R. R. HudElson, A.M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr. Richard Bradfield, A. B. O. B. Price, B. S. in Agr. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. CrislEr, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Secretary S. B. ShirkEy, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsham, Librarian. E. E. Brown, Business Manager. *In service of U. S. Department of Agriculture. tOn leave of absence. TABLE OF CONTENTS Introduction - 5 Review of Literature - 5 Experimental Procedure 7 Methods Used in Calculating Results — ~ 8 Animals Used in Experiment (Plates I-III.) . 9 Average of Results 16 Comparison of Results 19 Comparison of Maintenance Requirement During Summer and Winter Months 20 Summary and Discussion 20 Bibliography 21 Original Data and Calculations in Detail 23 Feed and Weight Record, Table 13 24 Dry Matter and Organic Matter in Feed, Table 14 30 Measurements of Experimental Steers, Table 15-16 40 Measurements of Control Steers, Table 17 44 Composition of Control Steers, Table 18 45 Energy Value of Gains by Periods, Table 19 45 Distribution of Control Steers, Table 20 46 Comparison of Control Steers and Experimental Steers. (Figures 1-5.) 46 The Influence of the Plane of Nutrition On the Maintenance Requirement of Cattle A. G. Hogan, W. D. Salmon, H. D. Fox In 1914 an investigation* was begun at the Missouri Agricul- tural Experiment Station to study effects of underfeeding. Calves of beef breeding were secured, divided into three groups, and each group was placed on a different plane of nutrition. Group I was fed to grow rapidly, but not to become fat. Group II was placed on a lower nutritional plane, and was fed to gain about one-half pound per day. Group III was placed on a still lower plane and was fed to gain about one-third of a pound per day. There were large differences in the food intake of the three groups, and after a considerable amount of data had been obtained it was decided to make a study of the maintenance requirement of these steers. REVIEW OF LITERATURE There has accumulated a considerable mass of literature con- cerning the maintenance requirement, in terms of energy, of animals as well as of man. Much of this material has no direct bearing on the problem discussed in this paper, but a short historical state- ment may be useful. Waters 1 pointed out in 1908 that if the ration of an animal were suddenly reduced to a point a little less than sufficient to maintain its weight, there would be a process of readjustment. After a time a stationary live weight would be obtained if the reduction were not too severe, and following that there might be an increase in weight. More recent data from the Missouri Experiment Station 2 show a lower maintenance requirement for animals on a low plane of •This investigation was initiated by F. B. Mumford, Dean of the College of Agriculture, and P. F. Trowbridge, formerly chairman of the department of agricultural chemistry. Since September, 1918, E. A. Trowbridge, chairman of the department of animal husbandry, has had general supervision of the project. This article was prepared by A. G. Hogan, who has been in immediate charge since September, 1920. Mr. Salmon supervised the pre- liminary calculations, and calculated the data for the summer periods. Mr. Fox made the calculations for the winter periods. A large number of workers have contributed to the success of the investigation, but it does not seem practicable* to mention them all individ- ually. A short article embodying the essential points of the investigation was published in the Journal of Agricultural Research, Vol. XXII, p. 115. 1 Refers to Bibliography, page 21. 6 Missouri Agr. Exp. Station Research Bulletin 51 nutrition. Three steers were full-fed until eleven months old, then subjected to a maintenance trial. In order of economy in maintenance requirement they ranked as follows : Steer 598 first, Steer 596 second, Steer 590 third. Following this No. 596 was fed, No. 598 was given one-half productive feed, anT fourth productive feed. The steers were agai. a maintenance trial in which they ranked as follows i,o. 590 first, No. 598 second, No. 596 third. Evidently the maintenance require- ment closely paralleled the plane of nutrition. Armsby 3a cites the observations of Zuntz and Hagemann show- ing that a surplus of feed stimulated muscular activity and rest- lessness of the horse to such an extent that a ration more than suf- ficient for maintenance of this animal when standing quietly in its stall, would not cause an increase in weight under ordinary condi- tions. Experiments with cattle 4 indicate a similar stimulating ef- fect upon their muscular movements. Armsby, therefore, con- cludes that at least a part of the lower maintenance cost may come from “voluntary restriction of motion on the part of the animals on a low nutritive plane.” An experiment by Armsby and Fries 5 showed that the main- tenance requirement of a two-year-old steer was increased 36 percent by a three-month fattening period in which the live weight was increased by about 300 pounds. The computed basal metabol- ism per 1000 pounds live weight per day showed the following var- iations in maintenance requirement : In proportion In proportion to 2-3 to weight power of live weight Unfattened 4,919 cal. 5,125 cal. Fattened 5,275 cal. 5,943 cal. “The basal katabolism increased faster than the body weight or the body surface as estimated by the Meeh formula. Apparently the accumulation of fat tended in some way to stimulate the general metabolism.” Both Kellner and Evvard have reported data, cited by Armsby 3b showing that fat steers have a higher maintenance requirement than those in medium condition. The extensive researches of F. G. Benedict and co-workers 6 in the field of human physiology are of especial significance. They have demonstrated that the basal metabolism of their subjects was markedly lower on a restricted diet than on a normal diet. In other words the maintenance requirement was lowered. In most The Influence of the Plane of Nutrition 7 cases we have mentioned it is impossible to decide to what extent decreased muscular activity accounts for the lower maintenance requirement when animals are on a low nutritive plane. EXPERIMENTAL The conditions of this investigation are unique in one respect, the animals were started on the project at weights varying from 154 to 238 pounds and thereafter were kept constantly on the same plane of nutrition. Since the animals were under observation for from four to seven years, any marked or permanent adjustment to nutritional conditions should become apparent. The ideal method of conducting an investigation of the main- tenance requirement of cattle would provide for a respiration cal- orimeter. Since that was impossible the alternative was to calcu- late the energy value of the food consumed, and correct this for the estimated value of the gains (or losses) in body weight. The net energy of the feed consumed was calculated in accordance with procedures developed by Armsby. The energy values of the changes in body weight were calculated from the composition of steers that had been analyzed by the department of agricultural chemistry at this station. Experimental Animals. — Three of the steers now under obser- vation were started on the investigation in 1914, and seven others were added in 1917. Some of the more significant early records are condensed in the following table. Table 1. — Groups, Dates oe Birth, and Breeds oe Animals. Ani- mal Group Date of birth Date put on Exp. Weight when put on Exp. Breed 528 I May 8, 1914 June 11, 1914 157 Hereford-high-grade 577 I March, 1914 Aug. 5, 1917 227 Shorthorn-grade 571 I March, 1917 Aug. 5, 1917 158 Hereford-grade 579 II May 2, 1914 May 30, 1914 154 Shorthorn-grade 573 II April, 1917 Aug. 5, 1917 203 Hereford-grade 578 II April, 1917 Aug. 5, 1917 238 Hereford-grade 585 III April 26, 1914 May 22, 1914 123 Hereford-high-grade 572 III April, 1917 Aug. 5, 1917 196 Hereford-grade 574 III April, 1917 Aug. 5, 1917 237 Hereford-grade 575 III April, 1917 Aug. 5, 1917 204 Hereford-grade 8 Missouri Agr. Exp. Station Research Bulletin 51 Quarters. — The steers had access to a shed open to the south. Adjoining this shed were dry lots sloping to the south, and having shade protection. Rations. — The concentrate consisted of the foil s ' Corn chop, 60 percent; wheat bran, 30 percent percent. The roughage fed from the beginning of me experiment until July 20, 1917, was timothy. For the next ten days a mixture of 5 parts timothy, 3 parts alfalfa and 2 parts oat straw was fed. Following this the roughage consisted of a mixture of 60 percent alfalfa and 40 percent oat straw. The animals were fed twice daily and had access to water at all times. Salt was accessible at feeding time. Weights. — The steers were weighed each morning, after feed- ing, but before watering. The weight given for the beginning of a period is the average of the ten preceding days. The weight given at the end is an average of the last ten days of the period. Periods. — The calculations are made for periods of 180 days, with the exception of the first period for each of the three older steers, which were as follows: No. 528, 130 days; No. 579, 142 days; No. 585, 150 days. In order that one period each year might be free from the disturbing effects of cold weather, the year was divided into a “summer” and “winter” period. The summer per- iods began in April or May, and ended in October. The winter per- iods began in October or November and ended in April or May. Energy Intake. — Our calculations of the energy values of the feed consumed are based on two methods described by Armsby 4 . In one case the dry matter, in the other the digestible organic nu- trients consumed, was used to calculate the net energy intake of the steers. The method of calculation based on dry matter consumed is as follows. For the concentrates the- value 83.82 therms per 100 pounds dry matter was used. This is the factor given for Armsby’s grain mixture No. 2*, which approximates the grain mixture used in this experiment. For timothy hay the value 48.63 therms per 100 pounds dry matter was used. The factor for the roughage mixture used in the latter part of the experiment was calculated from the Armsby values, for alfalfa 34.10 therms, and for oat straw *Armsby’s grain mixture No. 2 — 60 percent corn meal; 30 percent crushed oats; 10 per- cent O. P. linseed meal. Our grain mixture— 60 percent corn meal; 30 percent wheat bran; 10 percent O. P. linseed meal. The Influence of the Plane of Nutrition 9 Plate 1. — Taken at the beginning of the investigation. 5?9 10 Missouri Agr. Exp. Station Research Bulletin Plate II. — Taken after being fed three years on their respective nutritional planes. The Influence of the Plane of Nutrition 11 Plate III. — Taken after being fed six years on their respective nutritional planes. The Influence of the Plane of Nutrition 13 26.03 therms per 100 pounds dry matter. A mixture of 60 parts alfalfa and 40 parts oat straw would have a value of 30.87 therms per 100 pounds dry matter. The values used are summarized below in tabular form. Table 2. — Energy Values per 100 Pounds Dky Matter. Net energy values in therms Alfalfa hay 34.10 Oat straw 26.00 Mixture, 60 percent alfalfa, 40 percent oat straw 30.87 Timothy hay 48.63 Grain 83.82 The calculations of the energy value of the milk are also based on factors published by Armsby 3c . These are 29.01 therms per 100 pounds whole milk (4.4 percent) and 14.31 therms per 100 pounds skim milk (0.2 percent). From these values factors were com- puted for the different grades of milk used. The values used are given in Table 3. Table 3. — Net Energy Value oe Milk Used. Percent fat in milk Therms net energy per hundred pounds 4.40 (whole milk) 29.010 3.20 24.776 2.70 23.130 1.85 20.086 1.20 17.838 0.20 (skim milk) 14.310 Thef net energy intake, based on digestible organic nutrients con- sumed, was calculated by the procedure outlined below. The Armsby factor for the metabolizable energy of digestible organic matter from roughage is 1.588 therms per pound. For grains and similar feeds the factor is 1.769 therms per pound. Armsby 3d has also determined the “average energy expendi- ture” by cattle per 100 pounds of dry matter eaten. This is uiven in Table 4. 14 Missouri Agr. Exp. Station Research Bulletin 51 Table 4. — Energy Expended by Cattle per 100 lbs. Dry Matter Consumed. Roughage Energy expenditure in therms Timothy hay Alfalfa hay Oat straw Concentrate Grain mixture No. 2 35.47 53.03 46.00 51.76 The coefficients of digestibility used in these calculations were derived from digestion trials conducted under similar conditions at this Station . 7 These indicated that the digestibility of the ration varied with the relative amounts of hay and grain fed. The fac- tors used are given below. Table 5. — Digestion Factors eor Organic Matter. Ratio of 1:1 grain to hay 2:3 1:2 1:3, 4 or 5 1:6 or 7 1:8, 9 or 10 Hay only Factor .6956 .6695 .6434 .6340 .6229 .6030 .5832 Inasmuch as the thermal value of a pound of organic matter from grain differs from that of a similar weight of organic matter from roughage, the Armsby factors previously quoted in this paper could not be directly applied to the values obtained with the above digestion coefficients. Those factors would not provide for the widely varying proportions of grain and hay. The following meth- od therefore was used in computing the energy intake on the basis of digestible organic matter consumed. By use of the factors in Table 5 the w T eight in pounds of digestible organic matter in the mixed ration was determined for each period. This was multi- plied by 1.588, the Armsby factor for metabolizable energy in a pound of digestible organic matter from hay. The thermal value of digstible organic matter from grain is 1.769 however, or 0.181 therms more. Therefore each pound of digestible organic matter derived from grain was multiplied by 0.181, and the product added to the result obtained by multiplying the total digestible organic matter by 1.588. This gave the total metabolizable energy in both the hay and grain. The digestibility of the organic matter of the The Influence of the Plane of Nutrition 15 grain was estimated by difference. This ranged closely around 80 percent. The factors for energy expenditure are given in Table 4. It seemed impracticable to calculate the net energy of the milk consumed on the basis of digestible organic matter, so the calculation based on dry matter was used for milk. Since the amount was small, however, the method of calculation would have iittle effect on the final results. Changes in Body Weight. — In order to obtain data concern- ing the maintenance requirement of these steers, it is necessary to calculate the energy gained or lost through changes in body weight. Our calculations are based on analyses previously made by the department of agricultural chemistry*, University of Missouri. Control animals were selected from those on which analyses were available, on the basis of similar weights and measurements, and when possible of similar ages, daily gains and daily consumption of dry matter. In some cases suitable check animals were not avail- able, and the composition of steers for those periods was esti- mated by interpolation, using data of the preceding and succeeding periods. Using these assumed values for the composition of the steers during the different periods, the gain in protein and fat was readily calculated. The thermal equivalent of the protein and fat gained was calculated from data obtained by other investigators. Armsby 3e quotes data, computed by Kohler, giving the value 5.6776 calories per gram or 2.5753 therms per pound, for protein of mus- cular tissue of cattle. Fries 8 gives an average value of 9.4889 calories per gram of beef fat, or 4.3048 therms per pound. Since no suitable control animal was available for the last two periods of Steer 528, the gains for these two periods in terms of protein and fat were not calculated, and the energy value of a pound gain was assumed to be 3,000 therms per pound. This is the value given by Armsby 3f for animals of apparently similar condi- tion. The values we have used, also those published by Armsby are given in Table 6. Armsby’s values are consistently higher, as is to be expected. Our animals were thin, and contained less than the usual amount of fat in the gain. In calculating the maintenance requirements per 1,000 pounds live weight, Moulton’s 9 formula was used. He has shown that •These have not as yet been published. 16 Missouri Agr. Exp. Station Research Bulletin 51 Table 6. — Energy Value per Pound Gain. Approximate age Group Armsby’s > Values I II III Age Energy months therms therms therms months therms 6 .95575 .95575 .8343 1 1.170 18 1.0918 1.0583 .9445 2-3 1.374 36 1.7136 1.1608 1.0548 5-6 1.680 54 2.1993 1.4104 1.1013 11-12 2.292 66 2.50 1.5352 1.4790 18-24 3.000 78 3.00 1.660 1.6490 the surface areas of thin cattle are proportional to the five-eighths power of the live weight. The average maintenance requirement of the three groups is given in Tables 7 to 10 inclusive. The summer and winter periods are given separately, and each has been calculated by two methods. One is based on dry matter consumed, the other on digestible or- ganic nutrients consumed. In calculating the maintenance requirement on the basis of di- gestible organic matter consumed, digestion coefficients were used that had . been obtained at this station under similar conditions. This method is probably more accurate than the one based on dry matter consumed, and in this case gives a result somewhat higher. During the summer months there were four periods in all in which losses in live weight occurred. In calculating averages those periods were omitted, as the results are low. It is possible that Table 7. — Average Daily Maintenance Requirement During Summer Periods as Calculated from Dry Matter Consumed. Steer number Therms net energy per 1000 pounds based on 5-8 power of live weight Group I Group II Group III 528 — Average of 6 periods 5.870 5.280 5.073 577 — Average of 3 periods 571 — Average of 3 periods 579 — Average of 5 periods 4.920 3.830 4.409 578 — Average of 3 periods 573 — Average of 3 periods 585 — Average of 5 periods 4.221 4.041 4.302 3.256 3.921 575 — Average of 3 periods 574 — Average of 3 periods 572 — Average of 3 periods Average of each group 5.523 4.483 The Influence the Plane of Nutrition 17 Table 8. — Average Daily Maintenance Requirement During W inter Months as Calculated from Dry Matter Consumed. Steer number Therms net energy per 1000 pounds based on 5-8 power of live weight Group I Group II Group III 528 — Average of 6 periods 5.909 577 — Average of 3 periods 5.530 571 — Average of 3 periods 5.450 579 — Average of 6 periods 4.647 578 — Average of 3 periods 3.730 573 — Average of 3 periods 4.753 585 — Average of 6 periods • 4.366 575 — Average of 3 periods 4.260 574 — Average of 3 periods 4.673 572 — Average of 3 periods 3.157 Average of each group 5.770 4.444 4.164 they are correct, but the apparently diminished requirement may be due to an incorrect assumption as to the energy value of the loss in weight. One steer, No. 585, had a navel infection during the first summer period, accompanied by a very high maintenance requirement. This period also was discarded in calculating aver- ages. There is a close parallel between the intake of net energy and the maintenance requirement of the animal. The record of Steer 574 for the summer periods illustrates that tendency. For the first period the average daily intake of net energy was 3.884 therms per 1,000 pounds, based on the five-eighths power of the live weight, and the maintenance requirement was 3.818 therms. For the second period the energy intake was increased to 5.783 therms, and the Table 9. — Average Daily Maintenance Rrquirement During Summer Months as Calculated from Digestible Organic Matter Consumed. Steer number Therms net energy per 1000 pounds based on 5-8 power of live weight Group I Group II Group III 528 — Average of 6 periods 6.261 577 — Average of 3 periods 5.412 571 — Average of 3 periods 5. 174 579 — Average of 5 periods 5.260 578 — Average of 3 periods 4.192 573 — Average of 3 periods 4.893 585 — Average of 5 periods 4.725 575 — Average of 3 periods 4.454 574 — Average of 3 periods 4.591 572 — Average of 3 periods 3.649 Average of each group 5.777 4.869 4.408 18 Missouri Agr. Exp. Station Research Bulletin 51 Table 10.— Average Daily Maintenance Requirement During Winter Months as Calculated from Digestible Organic Matter Consumed Steer number Therms net energy per 1000 pounds based on 5-8 power of live weight Group I Group II Group III 528 5.965 5.713 5.553 577 571 579 5.071 4.494 5.513 578 573 585 * 4.625 4.429 5.290 3.754 575 574 572 Average of each group 5.799 5.037 4.869 maintenance requirement increased to 5.119 therms. In the third period the energy intake was 5.253 therms, and the maintenance requirement was 4.836 therms. In comparing the maintenance requirements of the three groups it should be kept in mind that Group I does not represent a high plane of nutrition. The aim was to secure maximum growth with no considerable fattening. Their maintenance requirements as computed in this paper correspond closely to the average of 22 respiration experiments by Armsby and Fries, and seven by Kellner on cattle in medium condition. A comparison of our results (com- puted on the basis of digestible organic matter consumed) and of those obtained by other investigators is given in Table 11. A few facts not shown by the data seem worthy of record. Although some of the steers were receiving a very scanty ration, they apparently did not have an unusual desire for food, and some care was necessary to prevent their “getting off feed.” This is especially true of the roughage, for it was impossible to induce them to consume a large quantity of hay. Any increase in the grain ration had to be very gradual. The dentition of these steers was apparently the same as for normal animals, as regards age. So far as could be determined by observation, the temporary teeth were lost at the normal age. Influence of Age. — The ages represented in this experiment vary from 30 days for some of the calves at the beginning of the first summer period to more than six years at the close of the seventh period. Apparently there was no relation between the age The Influence of the Plane of Nutrition 19 Table 11 . — Daily Maintenance Requirements of Cattle — Net Energy No. of Exper- iments Investigator Condition of animal Therms per 1000 lbs. live wt. Maximum Minimum Average Respiration Exp’s. 22 Armsby and Fries (3b) Medium 7.430 4.723 5.995 r. 7 Kellner (3b) Medium 6.780 4.921 5.742 Kellner (3b) Fat 8.871 7.319 7.946 Live Wt. Exp’s. 10 Armsby (3b) Thin 7.044 6.136 6.505 3 Armsby (3b) Thin 6.039 4.713 5.423 6 Haecker (3b) Medium 5.676 4.662 5.021 3 Eward (3b) Medium 7.079 5.841 6.173 7 Eckles (3b) Medium 7.079 5.841 6.173 1 Shirky (7) Medium a 7.732 2 Shirky (7) Thin b 5.0959 4.953 5.0245 3 Our results, summer periods Group I 7.380 4.915 5.777 3 Our results, summer periods Group II 5.724 3.809 4.869 4 Our results, summer periods Group III 5.217 3.276 4.408 3 Our results, winter periods . Group I 7.431 4.314 5.799 3 Our results, winter periods . Group II 7.598 3.246 5.037 4 Our results, winter periods . Group III 5.574 3.475 4.869 a Corresponds to Group I animals this experiment. b Corresponds to Group II of this experiment. and the maintenance requirement of these animals. Some of the steers showed a gradual decrease in the maintenance cost from the beginning to the end of the experiment. In such cases it was found that the energy intake per 1,000 pounds had also decreased. On the other hand, steers with an increasing energy intake showed an increased maintenance requirement. Maintenance trials on young animals usually give higher results than have been obtained with mature animals, but if age does influence the maintenance requirement the effect is too slight to be shown in a live weight ex- periment of this kind. Influence of Season. — The maintenance requirement of the steers in Groups I and II is slightly higher during the winter, as compared to the summer months. The animals in Group III how- ever required considerably more energy for maintenance during the winter periods than they did in the summer periods. Presumably the energy expenditure incident to the greater consumption of feed by the steers of Groups I and II is sufficiently great to make un- necessary the oxidation of a large quantity of additional nutrients during the winter months' in order to maintain the body tempera- ture. This is not the case with the steers on a lower nutritional plane, and so during periods of prevailingly low temperatures they 20 Missouri Agricultural Experiment Station Bulletin 51 must oxidize a larger amount of material in order to counteract the more rapid loss of heat from the body surface. The contrast be- tween the two seasons is shown in Table 12. Table 12— Daily Maintenance Requirement in Therms oe Cattle During Summer and Winter Months. During Summer and Winter Months Calculated on basis of digestible organic matter consumed Group I Group II Group III Summer 5.777 4.869 4.408 Winter 5.779 5.037 Calculated on basis of dry matter consumed 4.869 Summer 5.523 4.483 3.921 Winter 5.770 4.444 Average of results obtained by the two methods 4.164 Summer 5.650 4.676 4.165 Winter 5.775 4.741 4.517 SUMMARY AND DISCUSSION There is a close relation between the amount of net energy consumed and the maintenance requirement. Periods of high en- ergy intake were periods of high maintenance cost, while periods of low energy intake were accompanied by a lowered maintenance requirement. The averages of all periods show the following daily mainten- ance requirements per 1,000 pounds live weight, in terms of net energy. Summer months : Group I, (high plane) 5.650 therms ; Group II, (medium plane) 4.676 therms; Group III, (low plane) 4.165 therms. Winter months: Group I, 5.775 therms; Group II, 4.741 therms; Group III, 4.517 therms. The maintenance requirement of Group I is about 20 percent higher than that of Group II, and about 30 percent higher than that of Group III. If there is a definite relation between the age of animals and their maintenance requirements, it was obscured in this investi- gation by variations in the food intake. The maintenance requirement of these animals is higher in the winter than in the summer. The Influence of the Plane of Nutrition 21 BIBLIOGRAPHY 1. Waters, H. J. 1908 — Capacity of Animals to Grow Under Adverse Conditions. Proc. Soc. Promotion Agr. Science 29th Annual Meeting, p. 71. 2. Trowbridge, P. F., Moulton, C. R., Haigh, L. D. 1915 — The Maintenance Requirement of Cattle, Missouri Agricultural Experiment Station, Research Bulletin 18. 3. Armsby, H. P. 1917 — The Nutrition of Farm Animals. New York, The MacMillan Company. (a) Page 306 (c) Page 719 (e) Page 54 (b) Page 291 (d) Page 652 (f) Page 400 4. Armsby, H. P., Fries, J. A. 1915 — Net Energy of deeding Stuffs for Cattle. Jour. Agr. Research Vol. 3, p. 435. 5. Armsby, H. P., Fries, J. A. 1917 — Influence of Degree of Fatness of Cattle Upon Their Utiliza- tion of Feed. Jour. Agr. Research, Vol. 11, p. 451. 6. Benedict, F. G., Miles, W. R., Roth, Paul, Smith, H. M. Human Vitality and Efficiency under Prolonged Restricted Diet. Carnegie Institution of Washington, Publication No. 280. 7. Shirky, S. B. 1919 — The Extent to Which Growth Retarded During the Early Life of the Animal Can Be Regained. University of Missouri, Thesis for the degree, Master of Arts. 8. Fries, J. A. 1907 — Investigations in the Use of the Bomb Calorimeter. U. S. Dept, of Agriculture, Bur. Animal Industry, Bulletin 94, p. 13. 9. Moulton, C. R. 1916 — Units of Reference for Basal Metabolism and Their Interre- lations. Jour. Biol. Chem., Vol. 24, p. 299. ORIGINAL DATA AND CALCULATIONS IN DETAIL 24 Missouri Agricultural Experiment Station Bulletin 51 Table 13. — Weight in Pounds of Animals, and of Feed Consumed by Thirty Day Periods Date beginning of period Period No. Live 0 weight Pounds Grain Pounds Hay Pounds Milk Pounds Date beginning of period Period No. Live 0 weight Pounds Grain Pounds Hay Pounds Steer 585 585 (Cont. ) 5-22-14 1 143.1 8.75 321 8- 5-17 40 495 15.0 220.5 6-21-14 2 155 5.00 15.5 317.5 9- 4-17 41 498 37.0 225.0 7-21-14 3 173 59.5 300 10- 4-17 42 512 70.0 225.0 8-20-14 4 188 90.0 300 11- 3-17 43 529 67.0 232.0 9-19-14 5 194 104.0 299.8 12- 3-17 44 532 72.5 240.0 10-18-14 6 200 120.0 290.0 1- 2-18 45 554 76.0 240.0 11-18-14 7 192 140.0 100.0 2- 1-18 46 576 36.5 240.0 12-18-14 8 198 150 0 3- 3-]8 47 586 50.0 236.0 1-17-15 9 193 150 0 4- 2-18 48 576 225.0 2-16-15 10 194 150 0 5- 2-18 49 596 224.5 3-18-15 11 218 150 0 6- 1-18 50 578 240.0 4-17-15 12 223 ' 150.0 7- 1-18 51 568 240.0 5-17-15 13 229 150.0 7-31-18 52 554 240.0 6-16-15 14 233 155.0 8-30-18 53 542 253.0 7-17-15 15 235 11 75 153 5 9_29-] 8 54 532 255.0 8-16-15 16 251 36.7 163.5 10-29-18 55 506 263.5 9-15-15 17 256 47.0 150.5 1 1-28-18 56 532 278.0 10-14-15 18 272 60.0 154 0 12-28-18 57 558 328.5 11-14-15 19 290 60.0 180.0 1-27-19 58 562 317.0 12-14-15 20 303 60.0 180.0 2-26-19 59 591 330.0 1-13-16 21 312 60.0 180.0 3-28-19 60 603 330.0 2-12-16 22 337 56.5 180.0 4-27-1 9 61 604 317.5 3-13-16 23 345 34.0 180.0 5-27-19 62 614 326.5 4-12-16 24 356 30.0 180.0 6-26-19 63 622 335.0 5-12-16 25 360 30.0 180.0 7-26-19 64 643 360.0 6-11-16 26 361 31 .5 180.0 8-25-19 65 630 360.0 7-11-16 27 370 45.0 180.0 9-24-19 66 645 31.4 316.5 8-10-16 28 387 45.0 180.0 10-24-19 67 653 60.0 333.0 9- 9-16 29 386 42.5 170.0 11-23-19 68 688 58.0 257.5 10- 9-16 30 403 45.0 187.5 12-23-19 69 682 60.0 334.0 11- 8-16 31 413 45.0 201.0 1-22-20 70 716 60.0 360.0 12- 8-16 32 419 51.0 211.5 2-21-20 71 741 60.0 360.0 1- 7-17 33 437 75.0 225.0 3-22-20 72 763 60.0 360.0 2- 6-17 34 465 66.0 218.5 4-21-20 73 785 60.0 360.0 3- 8-17 35 484 55.0 210.0 5-21-20 74 825 60.0 359.0 4- 7-17 36 482 38.0 210.0 6-20-20 75 850 60.0 360.0 5- 7-17 37 496 30.0 210.0 7-20-20 76 858 60.0 359.0 6- 6-17 38 499 25.0 210.5 8-19-20 77 878 60.0 360.0 7- 6-17 39 504 15.0 208.5 9-18-20 78 885 60.0 360.0 Steer 579 579 (Cont. ) 5-30-14 1 154.0 2.55 226 8- 5-17 40 709 75.0 225.0 6-21-14 2 183.9 7.3 14.5 374 9- 4-17 41 707 86.0 225.0 7-21-14 3 204.6 24.0 47.0 420 10- 4-17 42 725 141.5 231.0 8-20-14 4 240.0 30.0 60.5 208 11- 3-17 43 744 136.5 247.5 9-19-14 5 270 37.0 74.0 420 12- 3-17 44 758 147.0 251.5 10-18-14 6 304 45.0 90.0 420 1- 2-18 45 789 164.5 253.0 11-18-14 7 305 45.0 103.0 140 2- 1-18 46 820 120.0 255.0 12-18-14 8 316 45.0 120.0 3- 3-18 47 822 67.0 255.0 1-17-15 9 321 45.0 120.0 4- 2-18 48 831 73.5 255.5 2-16-15 10 322 45.0 122.0 5- 2-18 49 851 22.5 292.5 3-18-15 11 335 45.0 120.0 6- 1-18 50 834.5 309.0 “Average weight of last ten days of period. The Influence of the Plane of Nutrition 25 Table 13 (Continued). — Weight in Pounds of Animals and of Feed Consumed by Thirty Day Periods Date beginning of period Period No. Live 0 weight Pounds Grain Pounds Hay Pounds Milk Pounds Date beginning of period Period No. Live 0 weight Pounds Grain Pounds Hay Pounds Steer 579 (Cont.) Steer 579 (Cont.) 4_17_15 12 334 120.0 7- 1-18 51 821 315.0 5-17-15 13 344 45.0 120.0 7-31-18 52 786 315.0 6-16-15 14 344 50.0 120.0 8-30-18 53 788 26.0 330.0 7 17-15 15 351 71 5 143 0 9-29-18 54 781 2.0 339.0 8-16-15 16 379 76 25 178 0 10-29-18 55 705 253.0 9-15-15 17 399 70.0 180 0 11-28-18 56 730 304.5 10-14-15 18 409 60.0 180 0 12-28-18 57 750 330.0 11-14-15 19 419 64.5 184 5 1-27-19 58 750 351.5 12-14-15 20 430 76.5 194.0 2-26-19 59 780 360.0 1-13-16 21 434 75.0 195 0 3-28-1 9 60 788 360.0 2-12-16 22 453 75 0 195 0 4-27-19 61 808 357.0 3-13-16 23 469 75.0 195.0 5-27-19 62 729 287.0 4-12-16 24 489 75.0 195.0 6-26-19 63 776 326.0 5-12-16 25 495 74.0 194.0 7-26-19 64 799 359.0 6-11-16 26 505 68.0 180.0 8-25-19 65 776 360.0 7-11-16 27 521 75.0 180.0 9-24-19 66 766 8.5 360.0 8-10-16 28 536 75.0 180.0 10-24-19 67 771 59.0 360.0 9- 9-16 29 552 75.0 180.0 11-25-19 68 782 60.0 360.0 10- 9-16 30 557 75.0 187.5 12-23-19 69 797 60.0 329.5 11- 8-16 31 565 81.0 189.0 1-22-20 70 823 60.0 360.0 12- 8-16 32 560 90.0 200.0 2-21-20 71 843 60.0 360.0 1- 7-17 33 582 90.0 216.0 3-22-20 72 855 60.0 360.0 2- 6-17 34 601 93.0 211.0 4-21-20 73 857 60.0 360.0 3- 8-17 35 632 105.0 225.0 5-21-20 74 866 59.0 328.0 4- 7-17 36 653 95.5 224.5 6-20-20 75 893 60.0 360.0 5- 7-17 37 674 94.0 229.5 7-20-20 76 910 60.0 358.0 6- 6-17 38 701 91.5 224.5 8-19-20 77 907 60.0 360.0 7- 6-17 39 708 75.0 225.0 9-19-20 78 918 60.0 360.0 Steer 528 528 (Cont. ) 6-11-14 1 179.0 2.5 124.9 8- 5-17 40 1008 188.0 227.0 6-21-14 2 202.8 14.0 14.0 489 9- 4-17 41 1029 196.0 238.0 7-21-14 3 252.0 44.0 53.5 582 10- 4-17 42 1043 209.0 231.0 8-20-14 4 303.8 69.0 75.0 600 11- 3-17 43 1070 218.0 240.0 9-19-14 5 363.8 110.5 101.5 600 12- 3-17 44 1076 225.0 240.0 10-18-14 6 425.1 120.0 120.0 600 1- 2-18 45 1080 225.0 240.0 11-18-14 7 426.8 105.2 120.0 176 2- 1-18 46 1178 225.0 240.0 12-18-14 8 439.4 120.0 120.0 3- 3-18 47 1147 225.0 240.0 1-17-15 9 457.2 120.0 120.0 4- 2-18 48 1163 224.5 240.5 2-16-15 10 460.6 120.0 119.0 5- 2-18 49 1184 150.0 293.5 3-18-15 11 482 120.0 120.0 6- 1-18 50 1177 97.0 318.0 4-17-15 12 497 120.0 120.0 7- 1-18 51 1159 28.0 330.0 5-17-15 13 507 120.0 120.0 7-31-18 52 1139 30.0 330.0 6-16-15 14 517 120.0 130.0 8-30-18 53 1122 30.0 330.0 7-17-15 15 527 120.0 150.0 9-29-18 54 1107 30.0 336.0 8-16-15 16 559 145.7 152.0 10-29-18 55 1103 87.0 333.0 9-15-15 17 586 150.0 180.0 11-28-18 56 1122 120 353.5 10-14-15 18 620 150.0 180.0 12-28-18 57 1155 120.0 357.5 11-14-15 19 629 155.0 184.5 1-27-19 58 1158 120.0 360.0 12-14-15 20 651 165.0 195.0 2-26-19 59 1172 150.5 360.0 1-13-16 21 659 165.0 195.0 3-28-19 60 121 1 150.0 360.0 2-12-16 22 678 165.0 195.0 4-27-19 61 1231 150.0 360.0 “Average weight of last ten days of period. 26 Missouri Agr. Exp. Station Research Bulletin 51 Table 13 (Continued). — Weight in Pounds of Animals and of Feed Consumed by Thirty Day Periods Date beginning of period Period No. Live 0 weight Pounds Grain Pounds Hay Pounds Milk Pounds Date beginning of period Period No. Live 0 weight Pounds Grain Pounds Hay Pounds Steer 528 (Cont.) Steer 628 (Cont.) 3-13-16 23 699 165.0 195.0 5-27-19 62 1258 150.0 360.0 4-12-16 24 727 165.0 195.0 6-26-19 63 1244 149.0 357.0 5-12-16 25 745 165.0 195.0 7-26-19 64 1267 150.0 360.0 6-11-16 26 768 165.0 194.5 8-25-19 65 1278 150.0 360.0 7-11-16 27 785 165.0 180.0 9-24-19 66 1278 150.0 360.0 8-10-16 28 805 165.0 180.5 10-24-19 67 1274 150.0 360.0 9- 9-16 29 837 179.5 189.0 11-23-19 68 1285 148.5 360.0 10- 9-16 30 869 180.0 202.5 12-23-19 69 1304 150.0 360.0 11- 8-16 31 881 180.0 211.0 1-22-20 70 1314 150.0 360.0 12- 8-16 32 878 180.0 214.5 2-21-20 71 1333 150.0 360.0 1- 7-17 33 887 180.0 226.0 3-22-20 72 1342 150.0 360.0 2- 6-17 34 915 180.0 225.0 4-21-20 73 1341 150.0 360.0 3- 8-17 35 933 180.0 225.5 5-21-20 74 1355 150.0 360.0 4- 7-17 36 942 180.0 225.0 6-20-20 75 1367 150.0 360.0 5- 7-17 37 964 190.0 228.5 7-20-20 76 1369 150.0 360.0 6- 6-17 38 986 180.0 225.0 8-19-20 77 1381 147.0 348.0 7- 6-17 39 990 180.0 225.0 9-19-20 78 1401 150.0 360.0 Date beginning of period Period No. Live 0 weight Pounds Grain Pounds Hay Pounds Milk Pounds Period No. Live 0 weight Pounds Grain Hay Pounds Pounds Milk Pounds Steer 578 8- 5-17 1 9- 4-17 2 10- 4-17 3 11- 8-17 4 12- 3-17 5 1- 2-18 6 2- 1-18 7 3- 3-18 8 4- 2-18 9 5- 2-18 10 6- 1-18 11 7- 1-18 12 7-31-18 13 8-30-18 14 9-29-18 15 10-29-18 16 11-28-18 17 12-28-18 18 1-27-19 19 2-26-19 20 3-28-19 21 4-27-19 22 5-27-19 23 6-26-19 24 7-26-19 25 8-25-19 26 9-24-19 27 252 268 279 314 317 331 365 383 378 411 418 413 402 393 383 372 375 395 409 432 438 455 458 457 466 490 496 25.0 99.5 152.5 44.3 111.0 216.4 72.2 130.8 54.5 90.0 140.4 76.1 194.0 81.0 164.5 81.6 165.0 29.0 164.0 21.5 165.5 10.0 192.0 196.0 195.0 195.0 208.0 210.0 2.0 210.0 233.5 226.5 240.0 240.0 240.0 240.0 241.5 245.0 17.0 269.0 30.0 270.0 30.5 270.0 Steer 577 1 249 2 272 3 288 4 317 5 339 6 361 7 404 8 441 9 472 10 504 11 524 12 538 13 555 14 560 15 578 16 599 17 617 18 640 19 671 20 690 21 730 22 750 23 764 24 778 25 813 26 820 27 810 23 .5 79, .4 33 .0 104 .1 71 .1 118. .2 94 .9 140. .4 105. .0 194. 4 105 .5 164. .5 107. .5 165. .0 120. .0 165. .0 119 .5 165. .5 91 .5 195. .0 90. .0 198. .5 90 .0 210. ,0 90 .0 210 .0 90. .0 223. .0 90, .0 225. .0 113. .0 211 .5 120. .0 210. 0 120. 0 210. 0 120 .0 231. 0 120 .0 240. 0 120. .0 240. 0 119 .0 228. ,5 120 .0 240. .0 120 .0 240. 0 119 .0 240. .0 120 .0 239. .0 107 .5 226. .5 ’Average weight of last ten days of period. The Influence of the Plane of Nutrition 27 Table 13 (Continued). — Weight in Pounds of Animals and of Feed Consumed by Thirty Day Periods Date beginning of period Period No. Live" weight Pounds Grain Pounds Hay Pounds Milk Pounds Period No. Live" weight Pounds Grain Pounds Hay Pounds Milk Pounds Steer 578 (Cont.) Steer 577 (Co nt.) 10-24-19 28 512 30.0 270.0 28 833 120.0 258.0 1 1-23-19 29 519 30 0 270.0 29 855 120.0 270.0 12 23 IQ 30 541 30 0 270 0 30 875 120.0 270.0 1-22-20 31 555 30 0 270 0 31 905 120.0 270.0 2-21-20 32 580 30.0 270.0 32 915 120.0 270.0 3-22-20 33 595 30 0 270 0 33 941 120.0 270.0 4-21-20 34 580 30.0 270.0 34 957 120.0 270.0 5-21-20 35 591 30 0 270 0 35 966 120.0 270.0 6-20-20 36 696 30.0 270.0 36 976 120.0 270 0 7-20-20 37 613 30.0 270.0 37 995 120.0 270.0 8-19-20 38 603 30 0 270.0 38 990 120.0 270.0 9-18-20 39 619 30.0 270.0 39 1000 120.0 270.0 Steer 575 Steer 574 8- 5-17 1 217 13.4 75.8 2.35 1 242 18.0 78.6 240.0 9- 4-17 2 227 .2 75.0 236 5 2 244 9.8 90.0 226.9 10- 4-17 3 228 109.0 67.0 3 252 23.4 126.2 62.5 11- 3-17 4 237 17.1 135.0 4 264 45.0 141.0 12- 3-17 5 246 30 0 143.9 5 268 46.5 150.0 1- 2-18 6 257 30.0 164.5 6 284 51.0 164.5 2- 1-18 7 275 18.3 165.0 7 309 51.2 165.0 3- 3-18 8 283 155.0 8 315 11.5 165.0 4- 2-18 9 290 150.0 9 315 5.5 165.0 5- 2-18 10 308 150.0 10 337 174.0 6- 1-18 11 304 150.0 11 337 174.0 7- 1-18 12 307 150.0 12 332 180.0 7-31-18 13 294 150.0 13 322 180.0 8-30-18 14 298 161.5 14 324 193.0 9-29-18 15 299 165.0 15 321 195.0 10-29-18 16 294 27.5 152.5 16 316 210.5 1 1-28-18 17 295 53.5 151.0 17 309 32.5 192.5 12-28-18 18 308 59.0 148.0 18 321 60.0 175.5 1-27-19 19 328 60.0 150.0 19 333 60.0 180.0 2-26-19 20 343 60.0 150.0 20 348 60.0 180.0 3-28-19 21 358 60.0 150.0 21 368 60.0 180.0 4-27-19 22 380 60.0 150.0 22 382 60.0 180.0 5-27-19 23 394 60.0 154.0 23 412 60.0 180.0 6-26-19 24 409 60.0 179.0 24 417 60.0 180.0 7-26-19 25 426 60.0 180.0 25 445 60.0 180.0 8-25-19 26 425 60.0 180.0 26 445 60.0 180.0 9-24-19 27 436 60.0 180.0 27 432 60.0 180.0 10-24-19 28 441 60.0 180.0 28 439 60.0 180.0 11-23-19 29 447 60.0 180.0 29 435 60.0 180.0 12-23-19 30 458 60.0 180.5 30 448 60.0 180.5 1-22-20 31 472 60.0 180.0 31 453 60.0 180.0 2-21-20 32 480 60.0 180.0 32 464 60.0 180.0 3-22-20 33 496 60.0 180.0 33 474 60.0 180.0 4-21-20 34 504 60.0 180.0 34 481 60.0 180.0 5-21-20 35 515 60.0 180.0 35 499 60.0 180.0 6-20-20 36 528 60.0 180.0 36 511 60.0 180.0 7-20-20 37 538 60.0 180.0 37 518 60.0 180.0 8-19-20 38 530 60.0 180.0 38 506 60.0 180.0 9-18-20 39 537 60.0 180.0 39 518 60.0 180.0 "Average weight of last ten days of period. 28 Missouri Agricultural Experiment Station Bulletin 51 Table 13 (Continued). — Weight in Pounds of Animals and of Feed Consumed by Thirty Day Periods Date beginning of period Period No. Live weight Pounds Grain Pounds Hay Pounds Milk Pounds Period No. Live weight Pounds Grain Pounds Hay Pounds Milk Pounds Steer 573 Steer 572 8- 5-17 1 197 14.1 64.9 214.0 1 210 16.4 72.6 215.4 9- 4-17 2 220 34.1 102.3 253.0 2 216 22.1 92.1 42.3 10- 4-17 3 237 75.7 108.4 50.5 3 231 28.8 124.9 11- 3-17 4 269 105.8 128.1 4 243 30.0 134.5 12- 3-17 5 290 87.5 145.4 5 252 30.0 146.5 1- 2-18 6 312 72.0 164.3 6 263 30.0 150.0 2- 1-18 7 335 62.5 165 0 7 280 16.3 154.5 3- 3-18 8 345 30.0 165 0 8 289 155.0 4- 2-18 9 354 29.5 165 6 9 295 0.5 150.0 5- 2-18 10 366 15.0 192 0 10 305 150.0 6- 1-18 11 364 6.0 202.5 11 307 150.0 7- 1-18 12 369 210.0 12 311 150.0 7-31-18 13 355 210 0 13 306 150.0 8-30-18 14 367 223 0 14 304 165.0 9-29-18 15 364 225 0 15 309 165.0 10-29-18 16 361 213 5 16 309 179.0 11-28-18 17 339 9.5 167 5 17 304 182.0 12-28-18 18 354 30 0 180 5 18 309 180.0 1-27-19 19 383 30.0 221.5 19 317 187.0 2-26-19 20 399 30.0 221.5 20 328 210.0 3-28-19 21 406 43.0 213.5 21 341 210.0 4-27-19 22 431 60.0 210.0 22 358 210.0 5-27-19 23 438 60.0 212.5 23 382 206.0 6-26-19 24 454 60.0 217.5 24 370 210.5 7-26-19 25 477 60.0 209.0 25 375 210.0 8-25-19 26 492 60.0 210.0 26 380 210.0 9-24-19 27 485 60.0 206.0 27 378 210.0 10-24-19 28 499 60.0 200.5 28 375 210.0 11-23-19 29 503 60.0 210.5 29 372 224.5 12-23-19 30 522 60.0 209.5 30 392 241.5 1-22-20 31 532 60.0 210.0 31 414 254.0 2-21-20 32 537 60.0 210.0 32 434 270.0 3-22-20 33 552 60.0 210.0 33 448 270.0 4-21-20 34 557 60.0 210.0 34 433 238.5 5-21-20 35 569 60.0 210.0 35 460 270.0 6-20-20 36 581 60.0 210.0 36 469 270.0 7-20-20 37 595 60.0 210.0 37 479 270.0 8-19-20 38 588 60.0 210.0 38 485 270.0 9-18-20 39 594 60.0 209.5 39 488 270.0 Steer 571 8- 5-17 1 173 24.5 79.3 82.8 9- 4-17 2 181 32.5 103.0 10- 4-17 3 200 81.1 96.9 11- 3-17 4 237 109.6 111.0 12- 3-17 5 264 105.9 123.3 1- 2-18 6 288 102.0 134.8 2- 1-18 7 321 102.0 136.5 3- 3-18 8 355 102.0 135.0 4- 2-18 9 383 103.0 142.0 5- 2-18 10 412 76.0 177.5 6- 1-18 11 433 70.0 189.0 The Influence of the Plane of Nutrition 29 Table 13 (Continued). — Weight in Pounds of Animals and of Feed Consumed by Thirty Day Periods Date beginning of period Period No. Live weight Pounds Grain Pounds Hay Pounds Milk Pounds Period No. Live weight Pounds Grain Pounds Hay Pounds Milk Pounds Steer 571 7- 1-18 (Cont.) 12 453 75.0 195.0 7-31-18 13 469 75.0 195.0 8-30-18 14 478 75 0 208.0 9-29-18 15 483 75.0 210.0 10-29-18 16 494 88.5 210.5 11-28-18 17 499 90.0 163.0 12-28-18 18 510 90.0 180.5 1-27-19 19 536 90.0 207.0 2-26-19 20 559 90.0 210.0 3-28-19 21 502 90.5 210.0 4-27-19 22 496 90.0 210.0 5-27-19 23 612 90.0 210.0 6-26-19 24 616 88.5 206.5 7-26-19 25 635 90.0 210.0 8-25-19 26 637 89.9 210.0 9-24-19 27 617 90.0 213.0 * 10-24-19 28 627 90.0 230.0 11-23-19 29 649 90.0 242.0 12-23-19 30 667 90.0 240.0 1-22-20 31 681 88.5 228.5 2-21-20 32 699 90.0 240.0 3-22-20 33 716 90.0 240.0 4-21-20 34 727 90.0 240.0 5-21-20 35 747 90.0 240.0 6-20-20 36 755 90.0 240.0 7-20-20 37 754 90.0 240.0 8-19-20 38 752 90.0 240.0 9-18-20 39 757 90.0 240.0 30 Missouri Agr. Exp. Station Research Bulletin 51 Table 14. — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer 585 1 8.020 7.655 7.655 4.464 2 4.714 14.211 4.548 18.107 13.559 11.480 3 53.811 48.683 48.683 28.392 4 84.158 67.090 76.090 44 . 376 5 97.228 87.906 87.906 51.267 6 26.974 111.024 100.449 100.449 58 . 582 7 127.800 115.769 115.769 67.516 8 136.098 123.498 123.498 72 . 024 9 136.098 123.498 123 . 498 72 . 024 10 136.098 123.498 123.498 72 . 024 11 136.098 123.498 123.498 72.024 12 136.098 123.498 123.498 72.024 13 136.098 123.498 123.498 72 . 024 14 144.047 131.017 131.017 76 . 409 15 10 616 142.540 10.231 129.640 139.871 84 . 342 16 33.196 151.424 31.983 138.174 170.157 102 . 605 17 61.760 139.866 60 . 208 138.656 188.864 113.885 18 54 . 639 143.105 52.691 129.375 182.066 115.430 19 54.330 167.263 52 . 394 51.213 103.617 65.687 20 53.710 168.654 51.810 152.515 204 . 324 129 . 542 21 53.724 178.837 51.847 162.701 214.548 136.023 22 50 . 568 164.763 48.801 148.733 197.534 125.236 23 30.431 164.763 29.311 148.733 178.044 110.904 24 27.072 165.125 26.013 149.135 175.148 109.100 25 27.072 165.125 26.013 149.135 175.148 109.100 26 28.422 164.641 27.311 157.199 184.510 114.931 27 40.609 166.170 39.021 152.310 191.331 121.304 28 40 . 609 166.170 39.021 152.310 191.331 121.304 29 38.357 156.923 36.857 143.843 180.700 114.564 30 40.228 172.617 38.630 158.217 196.837 124.795 31 39 . 846 185.540 38.217 170.070 208.287 132.054 32 45.156 193.915 43.311 176.576 219.887 139.408 33 66.404 207 . 059 63 . 690 187.969 251.659 159.552 34 58 . 432 201.112 56.044 182.562 238.606 151.276 35 48 . 702 193.255 46.712 175.435 222.147 140.841 36 33 . 684 193.255 32.324 175.435 207.759 129.413 37 26.769 193.255 25.721 175.435 201.156 125.300 38 22 . 343 193.255 21.477 175.435 196.912 118.738 39 13.406 191.933 12.886 174.233 187.119 112.833 40 13.406 202.915 12.886 184.205 197.091 118.846 41 33 . 063 207 . 059 31.781 187.969 219.750 136.882 42 62.557 210.216 60.132 191.418 251.550 159.483 43 59 . 770 218.126 57.487 198.876 256.363 162.534 44 64 . 609 221.997 62.164 201 . 753 263.917 167.323 45 67.736 219.940 65.172 199.900 265.072 168.056 46 32.538 219.940 31.307 199.900 231.207 144.019 47 44 . 566 215.383 42 . 880 195.753 238.633 151.283 48 207 . 593 187.163 187.163 109.143 49 206 . 844 187.950 187.950 109.612 50 219.508 199.066 199.066 116.095 51 222.130 197.050 197.050 114.920 52 222.130 197.050 197.050 114.920 53 234 . 484 208.361 208.361 121.516 The Influence of the Plane of Nutrition 31 Table 14 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer~685 (Cont.) 54 236.436 210.176 210.176 122.575 55 243.117 215.919 215.919 125.924 5fi 244.898 215.688 215.688 125.789 57 289.470 254.970 254.970 148.698 58 . . 279 . 340 246.020 246.020 143.479 5Q 290.748 256.060 256.060 143.502 fiO 290.740 256.060 256.060 149.354 fil 281.412 249.172 249.172 145.317 «2 291.199 260.959 260.959 152.191 88 297.227 . .267. 117 267.117 155.783 84 319.520 287.150 287.150 167.466 319 520 287.150 287.150 167.466 66 27.663 280.876 26.661 252.426 279.087 168.298 67 52.877 304.072 50.961 275.429 326.390 196.813 68 51.105 275.950 49.254 254.840 304 . 094 192.796 69 53.460 297.126 51.281 269.746 321.027 199.968 70 53.460 320.353 51.281 290.853 342.134 213.115 71 53.460 320.353 51.281 290.853 342.134 213.115 72 53.460 320.353 51.281 290.853 342.134 213.115 73 54.787 320.353 51.513 290.853 342.366 283.260 74 55.552 319.243 53.227 289.813 343.040 213.680 75 55.552 320.353 53.227 290.853 344.080 214.327 76 55.552 324.764 53.227 294 . 509 347.836 216.667 77 55.552 338 . 393 53.227 305.928 358.155 223.095 78 53.233 338.393 50.292 305.928 355.221 221.267 Steer 579 1 2.337 2.230 2.230 1.301 2 6.835 13.297 6.595 12.687 19.289 12.406 3 21.317 41.92 20.434 37.937 58.371 37.556 4 26.663 56.577 25.563 51.153 76.716 49.359 5 33.813 69.177 32.228 62.544 94 . 772 60.976 6 39.738 82.904 38.192 75.032 113.224 72.848 7 39.738 66.513 38.172 57.667 95.839 61.662 8 39.738 108.874 38.192 98 . 804 136.996 88.143 9 40.674 108.874 39.156 98.804 137.960 88.763 10 40.614 110.697 39.152 100.447 139.599 89.817 11 40.614 108.874 39.152 98.804 137.956 88.761 12 40.614 108.874 39.152 98.804 137.956 88.761 13 40.614 108.874 39.152 98.804 137.956 88.761 14 45.162 114.739 43 . 540 104.549 148.189 95.345 15 64.604 132.805 62.276 120.795 183.071 117.788 16 68.975 168.623 66.455 154.178 220.633 141.955 17 61.512 167.275 59.199 153.855 213.054 137.079 18 54 . 639 167.253 52.691 151.213 203 . 904 131.191 19 58.361 171.440 56.282 155.000 211.282 133.953 20 68.488 180.275 66.066 162.906 228.972 147.321 21 67.142 179.823 64 . 796 162.344 227.140 146.142 22 67.142 178.459 61.796 161.099 225.895 145.341 23 67.129 178.459 64.642 161.099 225.741 145.242 24 07.677 179.865 65.031 161.545 226.576 145.779 25 66.766 177.97 64.155 160.730 224.885 144.691 32 Missouri Agricultural Experiment Station Bulletin 51 Table 14 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer 579 (Cont.) 26 61.361 165.772 58.961 151.134 210.095 135.175 27 67.677 166.17 65.031 15^.310 217.341 139.837 28 67.677 166.17 65.031 152.31 217.341 139.837 29 67.677 166.17 65.031 152.31 217.341 139 . 837 30 73.553 173.084 70.868 158.654 229.523 147.675 31 71.727 174.565 68.796 159.945 228.741 147.171 32 79.687 184.228 76.431 167.765 244.196 157.116 33 79.687 198.794 76.431 180.464 256.895 165.286 34 82.341 194.189 78.976 176.279 255.255 164.231 35 92.967 207.059 89.168 187.969 277.137 178.309 36 84.667 206.591 81.239 187.541 268.770 172.927 37 83 . 882 211.187 80.602 191.717 272.319 175. 2’0 38 81.771 206.591 78.600 187.541 266.141 171.235 39 67.068 207.059 64.469 187.969 252.438 160.046 40 67.068 207.059 64.469 187.969 252.438 160.046 41 76.865 207.059 73.885 187.261 261.854 166.015 42 126.453 215.797 121.550 1»6.499 318.049 204.633 43 121.745 232.210 117.109 211.670 328.779 211.536 44 131.020 232.557 126.062 211.336 337.398 217.082 45 146.614 231.830 141.067 210.700 351.767 226.327 46 106.958 233.635 102.910 212.335 315.245 202.829 47 59.722 233.635 57.462 212.335 269.797 171.051 48 65.942 235.717 63.459 212.527 275.986 174.975 49 20.256 269.474 19.495 244.911 264.406 159.437 50 282.572 256.351 256.351 149 . 504 51 291.554 258.634 258.634 150.835 52 291.554 258.634 258.634 150.835 53 23.476 308.734 22.572 274.357 296.929 179.048 54 1.805 314.352 1.736 279.442 281.178 163.983 *»5 233.442 207.339 207.339 120.920 56 268.290 236.290 236.290 137.805 57 290.760 256 . 080 256 . 080 149 346 58 309 . 700 272.760 272.760 159.074 59 278.747 240.907 240.907 140.497 60 278.747 240.907 240.907 140.497 61 316.923 280.723 280.723 163.718 62 256.276 229.544 229 . 544 133.870 63 289.234 259.924 259.924 151.588 64 318.530 286.250 285.250 166.941 65 319.620 287.150 287.150 167.466 66 7.489 19.520 7.218 287.150 294.368 171.675 67 51.991 320.059 50.107 289.896 340.003 198.290 68 53.442 317.243 51.279 287.753 339.032 211.183 69 53.460 293.057 51.281 266.047 317.328 197.664 70 53.460 293.057 51.281 266.047 317.328 197.664 71 53.460 293.057 51.281 266.047 317.328 197.664 72 53.460 293.057 51.281 266.047 317.328 197.664 73 54.556 317.243 52.576 287.753 340.329 205.218 74 54.629 291.776 52.340 264.906 317.246 191.299 75 55.555 325.603 53.227 295.275 348.502 210.147 76 55.555 338.393 53.227 305.928 359.155 216.570 77 55.555 338.393 53.227 305.928 359.155 216.570 78 53.233 338.393 50.293 305.928 356.221 214.801 The Influence of the Plane of Nutrition 33 Table 14 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer 528 1 2.292 2.187 2.187 1.275 2 13.199 12.836 12.735 12.247 24.982 17.377 3 24.449 48.210 18.829 43.620 62.449 43.440 4 61.320 70.133 58.791 63.410 122.201 85.003 5 97.749 94.913 93.861 85.814 179.675 124.982 6 105.963 110.548 101.839 100.052 201.891 140.435 7 92.871 109.557 89.255 99.249 188.504 131.123 8 106.422 108.884 102.297 98.804 201.101 139.886 9 108.485 108.485 104.436 104.436 208.872 145.291 10 108.311 107.972 104.411 97.977 202.388 140.781 11 108.311 108.884 104.411 98.804 203.215 141.356 12 108.311 108.884 104.411 98.804 203.215 141.356 13 108.311 108.884 104.411 98.804 203.215 141.356 14 108.322 120.711 104.430 109.791 214.221 149.012 15 108.374 138.046 104.467 125.436 229.903 159.921 16 131.778 141.910 126.964 130.344 257.308 178.983 17 135.684 167.303 130.728 153.883 284.611 197.975 18 136.606 167.253 131.736 151.213 282.949 196.819 19 - 140.308 171.440 135.309 155.000 290.309 210.939 20 147.715 181.653 142.492 164.154 306.646 213.303 21 147.658 179.823 142.496 162.244 304 . 740 211.977 22 147.658 178.459 142.496 161.099 303.595 211.181 23 147.668 178.459 142.194 161.099 303 . 293 210.971 24 148.916 178.865 143.094 161.545 304 . 639 211.907 25 148.916 178.865 143.094 161.545 304.639 211.907 26 148.916 179.161 143.094 163.418 306.512 213.210 27 148.916 166.170 143.094 152.310 295.404 205.483 28 148.916 166.638 143.094 152.738 295.832 205.781 29 162.008 174.495 155.674 159.945 315.619 219.545 30 160.901 186.971 154.469 171.381 325.840 226.654 31 159.286 294 . 778 152.773 178.528 331.301 230.459 32 159.286 197.634 152.773 179.975 332.748 231.459 33 159.286 207.974 152.773 188.794 341.567 237.594 34 159.286 207.059 152.773 187.969 340.742 237.020 35 159.286 207.547 152.773 188.407 340.180 236.629 36 159.615 207.059 153.136 187.969 341.105 237.273 37 169.556 210.292 162.924 190.892 353.816 246.114 38 160.889 207 . 059 154.652 187.969 342.621 238 . 327 39 100.889 207.059 154.652 187.969 342.621 238.327 40 168.125 208.878 161.610 189.218 351.228 244.314 41 175.158 219.033 168.366 198.833 367.199 255.424 42 186.821 215.766 179.570 196.467 376.037 261.571 43 200.193 225.637 192.791 205.717 398.508 277.202 44 200.504 221.956 192.914 201.706 394 . 620 274.498 45 200.504 219.940 192.914 199.900 392.814 273.241 46 200.504 219.940 192.914 199.900 392.814 273 . 24 1 47 200 . 504 219.940 192.914 199.900 392.814 273.241 48 201.409 221.858 193.821 200.028 393.849 273.961 49 132.998 270.41 6 129.924 145.763 375.687 241.717 50 87.287 290.807 84 . 007 263.791 347.798 223 . 773 51 25.168 305 .412 24.211 270.932 295. 143 177.971 52 27.026 305.412 26 . 539 270.932 297.471 179.375 34 Missouri Agr. Exp. Station Research Bulletin 51 Table 14 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer 528 (Cont.) 53 27.085 306.048 26.042 271.986 298.028 179.711 182.640 54 27.085 311.443 26 . 042 276.843 302.885 55 78.548 307.178 75 . 522 272.820 348.342 220.848 56 107.548 311.360 103.381 274.210 377.591 239.393 57 105.910 314.960 101.791 277.380 378.171 240.394 58 105.576 317.150 101.468 279.310 380.778 241 .413 59 132.253 317.150 127.092 279.310 406.392 261.472 60 131.819 317.150 126 . 665 297.310 405.975 261.204 61 131 .006 319.497 126.089 283.017 409.106 263.219 266.241 62 130.710 321.207 125.894 287.909 413.803 63 131.769 316.860 126.888 284.760 411.648 264 . 854 64 132.111 319.520 127.268 287.150 414.418 266.637 65 132.181 319.520 127.392 287.150 414.542 266.716 66 132.181 319.520 127.392 287.150 414.542 266.716 67 132.181 320.062 127.392 289.899 417.291 268.485 68 132.249 320.273 126.891 290.783 417.670 268.731 69 133.678 320.273 128.232 290.783 419.015 269.594 70 133.678 320.273 128.232 290.783 419.015 269 . 594 71 133.678 320.273 128.232 290.783 419.015 269.594 72 122 . 678 320.273 128.232 290.783 419.015 269 . 594 73 136.974 320.027 131.291 290.783 422.074 271.562 74 136.573 320.027 130.850 290.783 421.633 271.279 75 138.888 320.027 130.850 290 . 783 421.633 271.279 76 138.888 320.061 130.850 290.246 421.096 270.933 77 136.110 327.113 130.407 295.730 426.137 274.177 78 133.085 338.093 125.734 305.928 431.662 277.731 Steer 578 1 22.361 104.971 21.945 96.528 118.023 74 . 827 2 39 . 593 102.156 38 . 058 92.737 130.795 82 . 924 3 64.522 124.266 62.020 111.347 173.367 111.544 4 80.281 131.978 77.222 120.328 197.550 127.104 5 67.831 180.588 65.270 164.178 229.448 147.627 6 72.191 150.751 69 . 459 137.011 206.470 132.843 7 112.607 151.207 109.855 137.427 247.232 159.101 8 25.852 151.207 24 . 874 137.427 162.301 102.899 9 19.304 152.693 18.577 137.668 156.245 94.216 10 8.998 176.927 8.660 160.696 169.356 98.768 11 180.748 161.434 161.434 94.148 12 180.485 160.100 160.100 93.370 13 180.485 160.100 160.100 93.370 14 192.763 171.302 171.302 99 . 903 15 194.679 173.056 173.056 100.926 16 1.805 159.232 1.736 144.700 146.436 85.400 17 204 . 744 180.211 180.211 105.099 18 199.535 175.740 175.740 102.492 19 211.430 186.214 186.214 108 . 600 20 211.430 186.214 186.214 108.600 21 211.430 186.214 186.214 108 . 600 22 212.996 188.682 188.682 110.039 23 215.343 193.013 193.013 112.565 24 217.377 195.347 195.347 113.926 The Influence of the Plane of Nutrition 35 TableT 4 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer]578 (Cont.) 25 14.981 238.663 14.438 214.483 228.921 133.507 26 26.438 239.560 25.480 215.280 240.760 145. 178 27 26.879 239.560 25.905 215.280 241.185 145.435 28 26.438 216.989 25.480 194.367 219.847 132.568 29 26.718 240.187 25.633 218.066 243.699 146.950 30 26.727 240.187 25.638 218.066 243.704 146.954 31 26.727 240.187 25.638 218.066 243 . 704 146.954 32 26.727 240.187 25.638 218.066 243 . 704 146.954 33 26.727 240.187 25.638 218.066 243 . 704 146.954 34 27.065 240.187 26.652 218.006 244.718 147.565 35 27.778 240.187 26.614 218.066 244.680 147.542 36 27.778 240.187 26.614 218.066 244 . 680 147.542 37 27.778 244 . 278 26.614 221.518 248.132 149.624 38 27.778 253.795 26.614 229.446 256.060 154.404 39 26.616 253.795 25.147 229.446 254.593 153.520 Steer 577 1 18.999 73.079 18.185 66.338 84.523 53.588 2 29.488 94.806 28.344 85.969 114.313 72.474 3 63.460 110.490 61.000 100.619 161.619 108.204 4 84.664 121.482 81.436 120.317 201.750 135.072 5 93.583 180.626 90.041 163.214 253 . 255 169.554 6 93 . 583 150.736 90.041 136.996 237.037 158.696 7 95.834 151.206 92.208 137.426 229 . 634 153.740 8 106.958 151.206 102.910 137.426 240.336 160.905 9 107.214 152.228 103.176 137.248 240.424 160.964 10 82 . 340 179.679 79.245 163.279 242.524 156.040 11 80.991 181.545 77.947 164.658 242.605 156.092 12 80.889 194.372 77.812 172.419 250.231 160.999 81.809 194.372 78.276 172.419 250.695 161.297 14 81.260 206.674 78.130 183.643 261.773 168.425 15 81.260 208.596 78.130 185.426 263.556 169.572 16 102.026 224.104 98.096 202 . 282 300.378 193.263 17 107.569 185.009 103.402 162.939 266.341 171.364 18 105.926 185.009 101.801 162.939 264 . 740 170.334 19 105.602 203.512 101.493 179.233 280.726 180.619 20 105.460 211.451 101.337 186.221 287.558 185.015 21 105.460 211.451 101.337 186.221 287.558 185.015 22 103.973 202.745 100.071 179.586 279.657 179.931 23 104.567 211.007 100.714 191.817 292.531 188.214 24 106.204 212.964 102.268 191.384 293 . 652 188.936 25 101 . 308 212.964 97.469 191.384 288 . 853 185.848 26 105.765 212.174 101.933 190.584 292.517 188.205 27 94.751 200.989 91.317 180.629 271.946 174.970 28 105.765 229.396 101.933 207.780 309.713 199.269 29 106.849 240.173 102.528 218.043 320.571 206 . 255 30 106.917 240.173 102.560 218.043 320.603 206 . 276 31 106.917 240. 173 102.560 218.043 320.603 206 . 276 32 106.917 240.173 102.560 218.043 320.603 206.276 33 106.917 240.173 102.660 218.043 320.603 206 . 276 34 110.112 240.173 105.556 218.043 323 . 599 208.204 35 111.110 240.173 106.454 218.043 324.497 208.781 36 Missouri Agr. Exp. Station Research Bulletin 51 Table 14 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer 577 (Cont.) 36 111.110 240.173 108.454 218.043 324.497 208.781 37 111.110 244.278 106.454 221.518 327.972 211.017 38 111.110 253.795 106.454 229.445 335.900 216.118 39 106.465 253.795 100.597 229.464 330.051 212.355 Steer 575 1 12.158 69.762 11.512 63.329 74.841 46.618 2 .182 69.026 .175 62.661 62.836 36.646 3 115.261 106.173 106.173 61.920 4 15.236 151.560 14.659 140.360 155.019 90.407 5 26.742 142.210 26.730 129.225 154.955 90.370 6 26.742 150.379 26.730 136.669 162.399 94.711 7 16.314 151.206 15.696 137.426 153.122 89 . 008 R 142.028 129.078 129.078 75.278 g 147.389 133.779 133.779 78 . 020 10 139.213 125.591 125.591 73 . 245 11 137.162 124.453 124.453 72.581 12 138.828 123.158 123 158 71.826 13 138.828 123.158 123.158 71.826 14 149.691 133.024 133 . 024 77.580 15 152.834 136.004 136.044 79.318 16 24.831 140.477 23 . 875 124.735 148.610 94.219 17 47.748 178.010 45.892 162.140 208.032 131.892 18 52.087 130.398 50.059 114.848 164.907 104.551 19 52.798 136.154 50.744 120.404 171.148 108.508 20 52 . 728 136.154 50.668 120.404 171.072 108.460 21 52 . 728 136.154 50.668 120.404 171.072 108.460 22 52.419 133.109 50.452 117.913 168.365 106.743 23 52.284 137.308 50.357 123.075 173.432 109.956 24 53 . 047 158.877 51.084 142.777 193.861 122.908 25 52.848 159.736 50.911 143.556 194.467 123.292 26 52.877 159.736 50.961 143.556 194.517 123.324 27 52.877 159.736 50.961 143.556 194.517 123.324 28 52.887 149.840 50.961 144.933 195.894 124.197 29 21.274 160.112 19.105 145.362 164.467 104.272 30 53.460 160.552 51.281 145.762 197.043 124.925 31 53.460 160.552 51.281 145.762 197.043 124.925 32 53.460 160.552 51.281 145.762 197.043 124.925 33 53.460 160.552 51.281 145.762 197.043 124.925 34 54.556 160.552 52.578 145.762 198.340 125 748 35 55.555 160.552 53.227 145.762 198.989 126.159 36 55.555 160.552 53.227 145.762 198.989 126.159 37 55.555 162.844 53.227 147.764 200.991 127.428 38 55.555 169.196 53.227 152.964 206.191 130.725 39 53.233 169.196 50.293 152.964 203.257 128.845 Steer 574 1 16.086 72.337 15.462 65.667 81.129 51.436 2 8.758 82.829 8.419 75.192 83.611 50.417 3 20.911 118.144 20.100 107.764 127.864 81.066 4 54.842 132.642 53.328 120.860 174.188 110.435 5 41.451 1 139.717 39 . 882 127.064 166.946 105.844 The Influence of the Plane of Nutrition 37 Table 14 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer 574 (Cont.) 6 45.109 150.755 43 . 389 137.013 180.402 114.375 7 45.686 151.215 43.959 137.431 181.390 115.001 8 10.198 151 .215 9.810 137.431 147.241 88.786 9 4.942 152.268 4.762 137.288 142.050 82.844 10 160.321 145.659 145.659 84 . 948 11 159.116 144.338 144.338 84.178 12 166 605 147.795 147.795 86.194 13 166 605 147.795 147.795 86.194 14 169 588 150.701 150.701 87.889 15 180.783 160.703 160.703 93 . 722 16 262.436 240.717 240.717 140.386 17 29.015 169.589 27.887 149.359 177.246 110.407 18 52.969 154.553 50.906 136.166 187.072 118.604 19 52.799 158.582 50.736 139.662 190.398 120.712 20 52.730 158.582 50.668 139.662 190.330 120.669 21 52.730 158.582 50.668 139.662 190.330 120.669 22 52.823 162.733 50.856 144.498 195.354 123.954 23 52.284 160.518 50.357 143.869 194.226 123.139 24 53.047 159.736 51.081 143.556 194.637 123.400 25 52.850 159.736 50.913 143.556 194.469 123.293 26 52.877 159.736 50.961 143.556 194.517 123.324 27 52.877 159.736 50.961 143.556 194.517 123.324 28 52.877 185.616 50.961 170.530 221.491 140.425 29 53.402 160.127 51 .250 145.377 196.627 124.662 30 53 . 460 160.571 51.281 145.781 197.062 124.937 31 53.460 160.571 51.281 145.781 197.062 124.937 32 53.460 160.571 51.281 145.781 197.062 124.937 33 53.460 160.571 51.281 145.781 197.062 124.937 34 54.556 160.571 52.578 145.781 198.359 125.760 35 55.556 160.571 52.578 145.781 199.008 126.171 36 55.555 160.571 53.227 145.781 199.008 126.171 37 55.555 162.844 53.227 147.764 200.991 127.428 38 55.555 169.196 53.227 152.964 206.191 130.725 39 53.233 169.196 50.293 152.964 203.257 128.865 Steer 573 1 ' 12.605 59.714 12.117 54 . 205 66.322 42.048 2 34.491 94.115 29 . 300 85.441 114.741 72.746 3 67.653 101.288 65.030 92.233 157.263 105.288 4 94.395 120.418 90.791 109.788 200.579 139.523 5 77.992 134.305 75.042 122.030 197.072 131.940 6 65.162 138.195 62.733 130.822 199.555 128.394 7 55.708 151.216 53 . 600 137.436 191.036 123.913 8 26.740 151.216 25.728 137.436 163.164 103.446 9 26.458 152.228 25.462 137.248 162.710 103.158 10 13.499 193.531 12.992 177.521 190.513 114.879 11 5.399 185.064 5.196 168.046 173.242 101.035 12 194.340 172.390 172.390 100.538 13 194.340 172.390 172.390 100.538 14 206 712 183 . 652 183 . 652 107.106 15 208.516 185.426 185.426 108.140 16 196.976 174.949 174.949 102.030 38 Missouri Agr. Exp. Station Research Bulletin 51 Table 14 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounds Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer 573 (Cont.) 17 8.437 147.573 8.108 129.973 137.081 82.660 18 26.485 159.023 25.453 140.053 165.506 103.094 19 26.401 195.150 25.374 171.870 197.244 122.863 20 26.365 195.150 25.334 171.870 197.204 122.839 21 36.957 187.201 35.479 164.871 200.350 127.022 22 52.070 186.347 50.103 165.079 215.182 136.425 23 52.285 189.491 50.358 169.844 220.202 139.608 24 53 . 046 193.005 51.081 173.465 224.536 142.356 25 52.849 185.425 50.912 166.635 217.547 137.925 26 52.877 186.335 50.961 167.455 218.416 138.476 27 52.877 182.819 50.961 164.299 215.260 136.475 28 52 . 877 178.251 50.961 161.515 212.476 134.710 29 53.447 186.960 51.277 169.782 221.019 140.126 30 53 . 460 170.905 51.281 169.188 220.469 139.777 31 53.460 171.309 51.281 169.588 220.869 140.031 32 53.460 171.309 51.281 169.588 220.869 140.031 33 53.460 171.309 51.281 169.588 220.869 140.031 34 54.556 171.309 52.578 169.588 222.166 140.853 35 55.555 171.309 53.227 169.588 222.815 141.265 36 55.555 171.309 53.227 169.588 222.815 141.265 37 55.555 189.994 53.227 172.293 225.520 142.980 38 55.555 197.396 53.227 178.458 231.785 146.952 39 53.233 197.396 50.293 178.458 228.751 145.028 Steer 572 1 14.652 66.795 14 . 084 60.635 74.719 47.372 2 19.763 48.768 18.997 76.952 95.949 60.832 3 25.738 116.754 24.740 106.329 131.069 83.098 4 26.760 116.404 25.741 115.288 141.029 89.412 5 26.731 135.427 25.729 123.064 148.793 94.335 6 26.741 164.213 25.729 % 151.683 177.412 112.479 7 14.535 141.583 13.985 ' 128.630 142.665 86.027 8 103.436 90.486 90.486 52.771 9 .445 138.389 .428 124.779 125.207 73.020 10 138.416 125.794 125.794 73.363 11 137.202 124.236 124.236 72 . 454 12 138.838 123.158 123.158 71.826 13 138.838 123.158 123.158 71.826 14 152.948 135.909 135.909 79.262 15 152.973 135.983 135.983 79.305 16 165.518 147.204 147.204 85 . 849 17 165.933 146.688 146.688 85.546 18 163.897 145.077 145.077 84 . 609 19 170.337 150.687 150.687 87.880 20 191 .319 169.249 169.249 98.706 21 191.319 169.249 169.249 98.706 22 188.313 167.389 167.389 97.621 23 189.891 164.650 164.650 96.024 24 186.759 167.839 167.839 97.884 25 186.314 167.455 167.455 97.660 26 186.333 167.455 167.455 97.660 27 186.333 167.455 167.455 97.660 The Influence of the Plane of Nutrition 39 Table 14 (Continued). — Dry Matter and Organic Matter in Feed by 30-Day Periods Period Dry matter in grain Pounsd Dry matter in hay Pounds Organic matter in grain Pounds Organic matter in hay Pounds Total organic matter Pounds Digestible organic matter Pounds Steer 572 (Cont.) 28 177.007 161.031 161.031 93.913 29 199.681 181.281 181.281 105.723 30 214.805 195.015 195.015 113.733 31 225.909 205.099 205 . 099 119.614 32 240 613 218.433 218.433 127.390 33 240 613 218.433 218.433 127.399 34 212.135 192.595 192.595 112.321 35 240 613 218.433 218.433 127.390 36 240 613 218.433 218.433 127.390 37. . . 244 276 221.518 221.518 129.199 38. . . 253 795 229.446 229.446 133.813 39 253.795 229.446 229.446 133.813 Steer 571 1 21.901 72.984 21.025 56.247 87.272 55 . 292 2 29.045 94.772 27.918 86 . 032 113.950 72.244 3 72.497 96.069 69 . 688 87.970 157.658 109.666 4 97.728 104.381 94 . 004 95.171 139.175 96.801 5 94 . 529 127.521 90.957 117.834 208.791 145.235 6 90.913 123.562 87.472 112.292 199.764 138.955 7 90.913 125.082 87.472 113.682 201.154 139.922 8 90.913 127.725 87.472 112.445 199.917 139.062 9 127.496 95.030 122.696 85.680 108.376 144.946 10 68.385 163.855 65.815 148.935 214.750 136.152 11 62.991 172.918 60.624 156.704 217.328 137.786 12 67.402 180.741 64 . 838 160.371 225.209 142.783 13 86.448 180.741 83.134 160.371 243.505 154.382 14 67.705 193.738 65.097 172.265 237.362 150.488 15 67.705 194.670 65.097 173.040 238.157 150.979 16 79 . 906 193.852 76 . 829 172.117 248.946 160.171 17 80.680 143.604 77.554 126.474 204 . 028 131.271 18 79 . 688 159.023 76.581 140.053 216.634 139.382 19 79 . 204 182.380 76.122 160.620 236.742 152.319 20 79.095 185.009 76.003 162.939 238.942 153.735 21 78.346 185.009 75.236 162.939 238.175 153.241 22 78.263 183.664 67.282 165.090 232.372 149.508 23 78.425 187.272 75.535 167.851 243.386 156.595 24 78.236 183.263 75.394 164.693 240.087 154.472 79.228 186.335 76.287 167.455 243.742 156.824 26 79.235 186.335 76.364 167.455 243.819 156.873 27 79.316 189.013 76.442 169.863 246.305 158.473 28 79.316 204 . 484 76.442 1S5.209 261.651 168.346 29 80. 160 215.246 76.905 195.416 272.321 172.651 30 80.188 213.455 76.920 193.795 270.615 171.633 31 78.852 208.103 75.639 184.533 260.172 164.949 32 80.188 208.103 76.920 184.533 261.453 165.761 33 80.188 208.103 76.920 184.544 261.464 165.768 34 82.282 208. 103 78.865 184.544 263 . 409 167.001 35 83.333 208. 103 79.841 184.544 264.385 167.620 36 83.333 208.103 79.841 184.544 264.385 167.620 37 83.333 217.133 79.841 196.902 276.743 175.455 38 83.333 225.595 79.841 203 . 952 283 . 793 179.925 39 79 . 850 225.595 77.434 203.952 281.386 177.399 40 Missouri Agr. Exp. Station Research Bulletin 51 Table 15. — Measurements in Centimeters of Steers at Beginning of Summer Periods and End of Winter Periods STEER 585 5-22-14 4-17-15 4-12-16 4-7-17 5-2-18 4-27-19 4-21-20 Height at withers 79.5 91.0 103.5 114.5 121.5 124.0 128.0 Height at hips 83.5 93.0 107.0 117.0 126.5 128.0 130.0 Girth of throat 47.0 52.0 64.0 72.0 79.0 78.0 85.0 Depth of chest 32.5 41.0 49.5 55.0 60.0 61.0 62.5 Width of chest 18.0 21.0 23.0 25.0 26.0 29.0 30.0 Width of paunch 20.4 31.0 38.0 42.0 45.5 52.5 54.5 Foreleg elbow to ground 49.0 53.0 63.0 65.0 72.0 72.0 74.0 Point of shoulder to top hip point . . . 60.0 68.0 87.0 94.0 96.0 101.5 109.0 Point of shoulder to ground 60.0 63.0 73.0 77.0 81.5 85.0 92.0 Poll to point of muzzle 26.0 31.0 39.0 44.0 49.0 49.0 49.0 Heart girth 85.0 102.0 121.0 140.0 149.0 153.0 163.0 Paunch girth 85.0 124.0 139.0 157.0 162.0 183.0 192.0 Width of hips 20.0 27.0 34.0 39.0 42.5 43.0 46.0 Width of loin 14.0 18.5 21.5 24.5 25.0 32.5 31.5 STEER 528 Date 6-11-14 4-17-15 4-12-16 4-7-17 5-2-18 4-27-19 4-21-20 Height at withers 81.8 109.0 122.5 130.0 138.5 140.0 143.0 Height at hips 86.3 110.5 126.5 136.0 141.0 143.0 144.5 Girth of throat 54.0 75.0 83.0 94.0 100.0 96.0 100.0 Depth of chest 36.5 53.0 61.5 66.0 73.0 76.0 78.0 Width of chest 20.3 30.0 33.5 40.0 41.0 41.0 43.5 Width of paunch 25.0 41.0 46.0 59.0 61.0 61.0 64.0 Foreleg elbow to ground 50.5 62.0 70.0 75.0 77.0 79.5 80.0 Point of shoulder to top hip point . . . 64.0 87.0 103.0 112.0 114.0 122.0 125.0 Point of shoulder to ground 60.5 72.0 82.0 86.0 91.0 92.0 96.0 Poll to point of muzzle 27.5 38.0 45.0 54.0 56.0 57.0 58.0 Heart girth 95.0 136.0 157.0 175.0 191.0 194.5 201.0 Paunch girth 98.0 144.0 166.0 193.0 206.0 209.0 217.0 Width of hips 22.7 35.0 42.5 49.5 54.5 56.0 59.0 Width of loin 15.3 24.0 27.0 31.5 35.0 38.5 36.0 STEER 579 Date 5-30-14 4-17-15 4-12-16 4-7-17 5-2-18 4-27-19 4-21-20 Height at withers 81.5 104.0 114.5 126.0 135.5 137.5 139.0 Height at hips 75.4 105.0 117.0 128.5 136.5 137.5 140.0 Girth of throat 49.0 64.0 69.0 73.0 83.0 79.0 85.0 Depth of chest 34.5 47.0 54.5 59.5 66.0 67.5 69.0 Width of chest 18.5 23.0 25.5 29.0 32.0 31.5 36.0 Width of paunch 23.0 32.0 40.0 46.0 53.0 49.5 52.0 Foreleg elbow to ground 51.0 63.0 68.0 75.0 79.0 83.0 84.0 Point of shoulder to top hip point . . . 62.0 84.0 99.0 105.0 114.0 121.0 121.0 Point of shoulder to ground 57.0 68.0 78.0 85.0 88.0 92.0 95.0 Poll to point of muzzle 27.5 36.0 43.0 49.0 53.0 52.5 53.0 Heart girth 90.0 120.0 137.0 150.0 168.0 169.0 171 .0 Paunch girth 92.0 129.0 142.0 162.0 181.0 181.0 187.0 Width of hips 21.5 30.0 36.0 41 .0 47.0 47.0 49.0 Width of loin 14.9 21.0 20.0 23.5 27.0 31.5 31.5 The Influence of the Plane of Nutrition 41 Table 15 (Continued). — Measurements in Centimeters of Steers at Beginning of Summer Periods and End of Winter Periods STEER 578 STEER 577 STEER 575 5-2-18 4 - 27-19 4-21-20 5-2-18 4-27-19 4-21-20 5-2-18 Height at withers 107.0 113.0 118.5 113.5 127.0 134.0 100.0 Height at hips 110.0 114.0 120.5 116.75 130.0 136.5 101.5 Girth of throat 64.0 63.0 75.0 73.0 79.0 90.0 60.0 Depth of chest 51.0 53.5 57.5 56.0 64.5 71.5 46.5 "Width of chest 24.0 25.0 27.5 28.0 32.5 37.0 21.0 Width of paunch 39.5 46.0 55.0 40.0 52.0 55.0 36.5 Foreleg elbow to ground 64.5 67.0 69.0 70.0 76.5 79.0 60.0 Point of shoulder to top hip point . . . 86.0 91.5 97.0 89.0 106.0 109.0 78.0 Point of shoulder to ground 72.0 76.5 82.0 78.0 88.0 95.0 69.0 Poll to point of muzzle 40.0 42.0 44.0 43.5 50.0 53.0 38.5 Heart girth 128.0 132.0 145.0 141.0 165.0 180.0 119.0 Paunch girth 146.0 162.0 183.0 149.0 179.0 197.0 131.0 Width of hips 34.0 35.5 41.0 34.5 41.5 46.5 27.5 Width of loin 19.0 24.0 26.0 23.5 31.0 35.0 18.0 STEER 575 (Cont.) STEER 574 STEER 573 Date 4-27-19 4-21-20 5-2-18 4-27-19 4-21-20 5-2-18 4-27-19 Height at withers 106.5 114.0 99.0 104.0 111.0 102.0 109.0 Height at hips 109.0 116.5 103.0 107.0 114.5 102.5 109.5 ■Girth of throat 63.0 03.0 63.0 65.5 72.0 66.0 68.0 Depth of chest 52.0 55.0 49.0 52.5 57.5 52.5 56.5 Width of chest 25.0 27.5 23.5 26.0 25.5 26.0 27.0 Width of paunch 43.0 44.5 37.0 42.0 42.5 39.0 44.5 Foreleg elbow to ground 64.0 70.0 59.0 64.0 68.0 58.0 63.0 Point of shoulder to top hip point . . . 87.0 94.0 80.0 87.0 96.0 77.0 87.0 Point of shoulder to ground 72.0 79.0 66.5 71.0 76.0 68.0 71.0 Poll to point of muzzle 42.0 47.0 39.0 42.5 46.0 40.0 43.0 Heart girth 130.0 142.0 127.0 132.0 145.0 131.0 137.0 Paunch girth 148.0 159.0 135.0 144.0 154.0 141.0 153.0 Width of hips 31.0 36.0 30.0 33.0 36.0 31.0 33.5 Width of loin 22.0 23.0 21.0 22.0 23.0 20.0 25.0 STEER 573 (Cont.) STEER 572 STEER 571 Date 4-21-20 5-2-18 4-27-19 4 21-20 5-2-18 4-27-19 1-21-20 Height at withers 117.0 98.5 105.0 107.5 100.25 114.5 118.5 Height at hips 115.5 99.25 105.4 110.5 104.75 117.5 123.0 Girth of throat 74.0 60.0 62.0 67.0 65.0 75.5 80.0 Depth of chest 61.0 46.0 48.5 51.0 51.0 57.5 62.5 Width of chest 28.5 23.0 24.0 26.0 26.5 31.5 33.0 Width of paunch 47.0 33.5 40.0 47.0 40.0 47.5 50.0 Foreleg elbow to ground 70.0 61.0 62.0 69.0 60.5 69.0 73.0 Point of shoulder to top hip point. . . 90.0 76.0 83.5 89.0 81.0 98.0 101.0 Point of shoulder to ground 77.0 69.0 71.5 76.5 67.5 77.5 81.0 Poll to point of muzzle 46.0 39.0 41.5 44.0 39.0 47.0 49.0 Heart girth 155.0 116.0 124.0 133.0 131.0 149.0 160.0 Paunch girth 166.0 124.0 136.0 158.0 143.0 162.5 175.0 Width of hips 37.0 29.25 31.0 34.0 31.5 38.0 41.0 Width of loin 27.5 18.25 19.5 27.0 19.75 23.0 28.0 42 Missouri Agr. Exp. Station Research Bulletin 51 Table 16. — Measurements in Centimeters of Steers at End of Summer Periods and Beginning of Winter Periods STEER 585 Date 10-19-14 10-13-15 10-8-16 10-3-17 10-28-18 10-23-19 10-17-20 Height at withers 87.5 94.7 109.0 118.5 122.0 125.0 129.5 Height at hips 92.0 100.0 114.0 122.0 126.0 127.5 130.5 Girth of throat 52.0 53.0 64.0 70.0 73.0 79.0 81.0 Depth of chest 39.5 42.0 50.0 55.0 59.0 61.0 65.5 Width of chest 19.0 19.0 23.0 25.0 25.0 29.5 36.0 Width of paunch 29.5 33.5 40.0 43.5 44.0 52.0 61.0 Foreleg, elbow to ground. . . . 52.0 59.0 65.0 69.0 73.0 74.0 74.0 Point of shoulder to top hip point 68.5 73.0 91.0 91.0 99.0 102.0 114.0 Point of shoulder to ground. . 64.0 64.0 75.0 81 .0 82.0 85.5 89.0 Poll to point of muzzle 32.0 33.0 43.0 46.5 48.0 48.5 51.0 Heart girth 99.0 107.0 123.0 139.0 145.0 154.0 169.0 Paunch girth 116.0 129.0 145.0 160.0 166.0 185.0 204.0 Width of hips 26.5 29.7 36.0 38.0 41.0 43.5 47.5 Width of loin 17.0 18.0 20.0 24.0 26.0 30.0 36.0 STEER 528 Date 10-19-14 10-13-15 10-8-16 10-3-17 10-28-18 10-29-19 10-17-20 Height at withers 101.5 115.5 127.0 133.5 139.0 144.0 144.0 Height at hips 103.5 120.5 131.5 138.5 143.0 144.5 143.0 Girth of throat 68.0 73.0 83.0 93. 95.0 99.0 102.0 Depth of chest 46.5 55.5 64.5 71.0 73.5 78.0 79.0 Width of chest 26.0 28.0 35.5 38.5 38.0 41.0 46.0 Width of paunch 39.5 46.7 52.5 55.5 55.0 62.5 65.0 Foreleg, elbow to ground .... 60.0 70.0 72.0 76.0 81.0 81.5 77.0 Point of shoulder to top hip point 80.0 95.0 104.0 113.0 122.0 124.5 120.0 . Point of shoulder to ground. . 67.0 77.0 87.0 88.0 89.0 94.0 96.0 Poll to point of muzzle 35.0 40.0 50.0 55.5 55.0 57.0 58.0 Heart girth 125.0 144.0 165.0 180.0 189.0 198.5 205.0 Paunch girth 144.0 164.0 183.0 192.0 198.0 214.0 222.0 Width of hips 32.0 39.2 46.5 51.0 54.5 59.0 60.0 Width of loin 20.5 23.0 30.0 31.0 34.5 36.5 39.5 STEER 579 Date 10-19-14 10-13-15 10-8-16 10-3-17 10-28-18 10-29-19 10-17-20 Height at withers 96.0 110.0 121.0 129.5 135.5 138.5 139.5 Height at hips 101.5 114.0 125.0 133.5 137.0 138.5 141.0 Girth of throat 59.0 62.0 69.0 80.0 77.0 78.0 81.5 Depth of chest 42.0 49.5 55.0 61.5 64.0 65.5 68.5 Width of chest 21.0 24.0 29.0 30.5 29.5 31.5 39.0 Width of paunch 31.5 39.0 43.5 45.5 48.0 47.5 52.0 Foreleg, elbow to ground .... Point of shoulder to top hip 60.0 67.5 69.0 74.0 82.5 83.0 78.0 point 79.0 91.0 100.0 112.0 120.0 119.0 118.0 Point of shoulder to ground. . 65.0 75.0 85.0 85.0 90.0 90.5 94.5 Poll to point of muzzle 34.0 38.0 46.0 51.5 53.0 52.0 54.0 Heart girth 110.5 124.0 143.0 155.0 163.0 164.0 176.5 Paunch girth 124.0 143.0 155.0 167.0 172.0 176.0 191.0 Width of hips 28.0 32.7 39.0 43.5 47.0 48.0 50.0 Width of loin 20.0 20.5 22.0 25.0 26.5 30.5 31.5 The Influence of the Plane of Nutrition 43 Table 16 (Continued). — Measurements in Centimeters of Steers at End of Summer Periods and Beginning of Winter Periods STEER 678 STEER 577 Date 11-3-17 10-28-18 10-29-19 10-17-20 11-3-17 10-28-18 10-29-19 Height at withers 97.0 110.5 113.5 121.0 97.0 122.0 132.0 Height at hips 100.0 111.5 117.0 123.0 102.0 124.5 135.0 Girth of throat 56.0 60.0 66.0 69.0 60.0 73.0 79.0 Depth of chest 43.0 50.5 53.5 59.0 45.0 59.0 67.0 Width of chest* 21.5 24.5 24.0 27.5 27.0 30.0 35.5 Width of paunch 40.5 43.0 49.5 51.0 39.0 46.0 52.5 Foreleg, elbow to ground .... 57.0 64.0 69.0 69.0 61.0 71.0 80.0 Point of shoulder to top hip, point 76.0 90.0 94.0 105.0 74.0 100.0 109.0 Point of shoulder to ground . . 67.0 75.0 81.0 82.0 68.0 81.0 89.5 Poll to point of muzzle 35.5 41.0 43.0 48.0 35.5 46.0 50.5 Heart girth 111.0 126.0 137.0 151.0 115.0 150.0 171.0 Paunch girth 138.0 154.0 171.0 177.0 137.0 166.0 186.0 Width of hips 29.0 34.0 37.5 41.0 27.5 38.0 44.0 Width of loin 17.5 20.5 24.0 26.5 18.5 25.0 33.0 STEER 577— (C ont .) STEER 575 STEER 574 Date 10-17-20 11 3-17 10-28-18 10-29-19 10-17-20 11-3-17 10-28-18 Height at withers 136.5 91.5 102.5 111.0 118.5 91.0 102.0 HeiTht at hips 138.0 95.0 104.5 115.5 119.0 95.5 105.0 Girth of throat 89.0 55.0 57.0 70.0 72.0 58.5 57.0 Depth of chest 73.0 42.0 47.5 54.0 57.5 44.5 49.5 Width of chest 40.0 21.0 22.0 26.0 29.5 23.0 24.0 Width of paunch 56.0 36.0 42.5 47.5 49.5 39.0 40.0 Foreleg, elbow to ground .... 81.5 57.0 62.0 69.0 70.0 56.0 60.0 Point of shoulder to top hip point 122.0 72.0 85.0 91.5 100.0 72.0 83.0 Point of shoulder to ground. . 93.0 65.0 69.0 78.5 79.0 63.0 67.0 Poll to point of muzzle 56.0 34.0 40.0 45.0 50.0 36.0 41.0 Heart girth 181.0 107.0 119.0 137.0 146.0 113.0 128.0 Paunch girth 201.0 129.0 148.0 165.0 169.0 132.0 148.0 Width of hips 48.0 24.0 28.0 33.0 37.5 27.0 30.5 Width of loin 35.5 16.5 18.5 23.0 26.5 17.0 22.0 STEER STEER 574— (Cont.) STEER 573 572 Date 10-29-19 10-17-20 11-3-17 10-28-18 10-29-19 10-17-20 11-3-17 Height at withers 108.5 114.0 92.0 106.5 114.5 119.0 89.5 Height at hips 112.0 116.5 94.0 107.0 115.0 118.5 93.0 Girth of throat 69.0 70.0 58.0 64.0 74.0 75.0 55.0 Depth of chest 56.0 60.5 44.5 51.5 58.0 68.0 40.5 Width of chest 29.0 32.5 24.5 25.0 30.0 30.5 22.5 Width of paunch 46.5 47.0 39.0 43.0 48.5 50.5 36.0 Foreleg, elbow to ground .... 66.0 65.0 55.0 62.0 68.0 66.5 56.0 Point of shoulder to top hip point 92.5 100.0 71.0 83.0 94.0 101.0 70.0 Point of shoulder to ground. . 75.0 78.0 63.0 72.0 75.5 79.0 62.0 Poll to point of muzzle 45.5 49.0 35.0 41.0 44.5 49.0 34.0 Heart girth 145.0 150.0 116.0 133.0 149.0 159.0 116.0 Paunch girth 160.0 165.0 138.0 150.0 167.0 175.0 126.0 Width of hips 35.0 38.0 28.0 31.5 36.0 39.0 25.0 Width of loin 25.0 27.5 19.0 21.0 28.0 27.5 16.5 44 Missouri Agr. Exp. Station Research Bulletin 51 Table 16 (Continued). — Measurements in Centimeters of Steers at End of Summer Periods and Beginning Winter Periods STEER 572— (Cont.) STEER 671 10-28-18 102.25 10-29-19 107.5 10-17-20 11-3-17 10 28-18 107.0 10-29-19 116.0 10-17-20 120.0 Height at withers 115.5 83.5 Height at hips 102.25 109.0 113.0 88.5 111.0 120.5 123.5 Girth of throat 58.0 65.0 65.0 54.0 68.0 72.0 78.0 Depth of chest 46.5 50.0 54.5 39.0 53.0 59.5 63.5 Width of chest 23.5 24.0 29.0 20.0 29.0 33.5 34.5 Width of paunch 39.5 41.0 49.0 35.0 48.0 50.0 52.5 Foreleg, elbow to ground. . . . 63.0 67.0 67.0 53.0 66.0 71.5 72.0 Point of shoulder to top hip point 82.0 85.5 98.0 65.0 88.0 101.0 110.0 Point of shoulder to ground . . 67.0 76.0 77.0 57.0 70.0 81.5 80.0 Poll to point of muzzle 39.0 42.0 46.0 32.0 42.0 49.0 51.0 Heart girth 120.0 129.0 137.0 100.0 140.0 156.0 165.0 Paunch girth. 138.0 145.0 163.0 125.0 163.0 167.0 182.0 Width of hips 30.0 32.0 35.0 24.0 35.0 39.5 43.0 Width of loin 18.5 23.0 26.5 16.0 23.0 29.0 28.5 Table 17. — Measurements* in Centimeters of Control Animals at Time of Slaughter Steer No 554 552 523 526 512 531 525 509 Height at withers . 90.5 96.5 128.8 130.0 153.0 114.75 124.8 140.0 Height at hips .... 96.8 98.0 130.0 140.5 150.5 115.5 126.8 139.0 Girth of throat. . . . 55.0 60.0 87.0 94.0 100.0 72.0 80.0 90.5 Depth of chest .... 38.5 44.0 66.0 71.5 80.0 54.5 62.0 67.0 Width of ohest .... 22.3 25.0 36.0 43.0 44.5 26.5 30.5 40.5 Width of Paunch. . Foreleg, elbow to 25.6 33.3 62.0 57.0 62.0 42.0 54.0 53.0 ground Point of shoulder to 60.0 61.2 76.5 85.0 88.0 72.5 73.8 84.5 top hip point . . . Point of shoulder to 70.5 75.0 112.0 124.0 127.0 94.0 105.0 116.0 ground Poll to point of 67.0 70.0 88.0 94.0 101.0 88.5 85.0 98.5 muzzle 31.5 35.0 52.5 54.0 55.0 43.0 51.0 52.0 Heart girth 101.0 114.0 175.0 197.0 210.0 146.0 160.0 186.0 Paunch girth 103.0 125.0 210.0 212.0 220.0 160.0 189.0 204.0 Width of hips 23.1 27.0 46.0 53.0 56.0 35.5 40.5 48.5 Width of loin 17.0 21.7 37.0 41.0 43.5 27.5 31.5 41.0 *From unpublished data furnished' by C. R. Moulton, department agricultural ohemistry, Missouri Agricultural Experiment Station. The Influence on the Plane of Nutrition 45 Table 18. — Composition of Control Animals Steer Age days Weight pounds Composition of body Dry matter per cent Protein per oent Fat per cent 554 90 196. 0 32 . 836 20.038 7.395 552 160 256.2 35.928 19.469 10.555 523 798 864.2 39.479 19.219 15.134 526 1217 1088.2 44.552 18.813 20.435 512 1454 1250.4 48.001 18.094 24.299 531 588 479.6 37.227 20.263 10.355 525 800 694.6 37.338 20.031 11.787 509 1363 1004.2 42.079 20.181 16.232 From unpublished data furnished by C. R. Moulton, department agricultural chemistry Missouri Agricultural Experiment Station. Table 19. — Energy Value of Gains, Calculated for Summer Ferioes Periodt Steer 1 Therms 2 Therms 3 Therms 4 Therms 5 Therms 6 Therms 7 Therms Group 1 528 577 .95575 1.0918 1.0918 1.2279 1.2279 1.7136 1.7136 2.1993 2.500 3.000 571 1.0918 1.2279 1.7136 Group II 579 578 .95575 1.0583 1.0583 1.0583 1 . 0583 1 . 1608 1.1608 1.4104 1.5352 1.66 573 1 . 0583 1.0583 1 . 1608 Group III 585 575 .8343 .9445 .9445 1 . 0548 .9445 1.1013 1.0548 1.1013 1.1479 1.649 574 .9445 1.0548 1.1013 572 .9445 1.0548 1.1013 Table 18 shows the ages, weights, and percentage composition of live weight of the control animals used in this study. tThese same values apply also to the winter periods of the seven younger steers. In the case of the three older animals, Nos. 528, 579, and 585, the first winter period corresponds to the second summer period and so on, thus making the sixth winter period correspond to the seventh summer period. From these data the composition of the gains was estimated for all periods except period seven of 528. No control animal to fit this period could be found. The value of three therms per pound gain was used in this case. This value was assumed on the basis of Armsby’s calculation (3f) of the energy value of gains from Lawes and Gilbert’s analyses on four-year-old fattening cattle. Table 19 shows the estimated energy value of the gains by periods for all the steers. Table 20 shows distribution of the control animals. Check animals were first fitted to the old steer in eaoh group; then the young animals were oompared with the old animal in their re- spective groups rather than with check animals direct. 46 Missouri Agr. Exp. Station Research Bulletin 51 Table 20. — Distribution oe Control Animals. Period 1 2 3 4 5 6 7 Steer 528 552 Inter- polate 523 Inter- polate 526 512 577 571 Period 2 of 528 523 Period 4 of 528 579 552 I nter- polate Inter- polate 525 Inter- polate Inter- polate 509 578 573 Period 2 of 579 Period 3 of 579 525 585 554 Inter- polate Inter- polate 531 Inter- polate 525 523 575 574 572 Period 3 of. 585 531 Period 5 of 585 Fig. 1. — Weights — Comparison of control animals and experimental animals. The Influence on the Plane of Nutrition 47 Fig. 2. — Heart Girth — Comparison of control animals and experimental animals. Fig- 3. — Depth of Chest — Comparison of control animals and experimental animals. 48 Missouri Agr. Exp. Station Research Bulletin 51 Fig. 5. — Height • at Withers — Comparison of control animals and experimental animals. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 52 SCARRED ENDOSPERM AND SIZE INHERITANCE ERRATA transpose table headings of the two (ties opposite page 6. Page 7, table 3: Strike out “Average Ker- Weight in mgs.” in box head of last col- !n. 5 OF MAIZE rized June 1, 1922.) COLUMBIA, MISSOURI JULY, 1922 48 Missouri Agr. Exp. Station Research Bulletin 51 O — f\> to. -C* cn O' CD <£> o — rj to K 0> 0' 3 CO Fig. 5. — Height at Withers — Comparison of control animals and experimental animals. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 52 SCARRED ENDOSPERM AND SIZE INHERITANCE IN KERNELS OF MAIZE (Publication authorized June 1, 1922.) COLUMBIA, MISSOURI JULY, 1922 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL, THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY P. E. BURTON H. J. BLANTON St. Louis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION J. C. JONES, PH. D., LL. D., PRESIDENT OF THE UNIVERSITY F. B. MUMFORD, M. S., DIRECTOR STATION STAFF JULY, 1922 AGRICULTURAL CHEMISTRY C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, Ph. D. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. SiEveking, B. S. in Agr. A. M. Cowan, A. M. AGRICULTURAL ENGINEERING J. C. WoolEy, B. S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumeord, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY VV. J. Robbins, Ph. D. E. F. Hopkins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale, B. S. in Agr. W. W. Swett, A. M. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. C. McBride, B. S. in Agr. FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, A. M. O. W. Letson, B. S. in Agr. Alva C. Hill, B. S. in Agr. Miss Pearl Drummond, A. A.* RURAL LIFE O. R. Johnson, A. M. S. D. Gromer, A. M. E. L. Morgan, A.M. Ben H. Frame, B. S. in Agr. Owen Howells HORTICULTURE V. R. Gardner, M. S. A. H. D. Hooker, Jr., Ph. D. J. T. Rosa, Jr., M. S. F. C. Bradford, M. S. H. G. Swartwout, B. S. in Agr. poultry husbandry H. L. Kempster, B. S. Earl W. Henderson, B.S. SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M I W. A. Albrecht, Ph. D. F. L. Duley, A.M. R. R. Hudelson, A.M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr Richard Bradfield, A. B. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Secretary S. B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsham, Librarian. E. E. Brown, Business Manager. In service of U. S. Department of Agriculture. Scarred Endosperm and Size Inheri- tance in Kernels of Maize William H. Eyster* In the summer of 1920 the writer found in a field of corn in cen- tral Pennsylvania a number of plants with striking chlorophyl patterns which are unlike any that have yet been described. These plants were numbered and marked in the field so that they could be identified at harvest time. The matured ears were sent to the writer at the Univer- sity of Missouri by Mr. Webster Snyder in whose field they were dis- covered. Plantings were made from each ear in the greenhouse the following winter and the seedlings were found to be entirely green. One plant from each ear was grown to maturity in the greenhouse and self pollinated. In the summer of 1921 field plantings were made from the original ears and also from the self pollinated greenhouse ears. The F 2 progenies segregated plants with the chlorophyl patterns of the original plants together with a number of other characters, including a pistillate plant similar in appearance to tassel ear (Emerson, 1920) and the endosperm character described in this paper, which has been desig- nated scarred endosperm. The field plantings of 1921 were made at the Missouri Agricul- tural Experiment Station as part of a project in the genetics of maize carried on in the Department of Field Crops. DESCRIPTION OF SCARRED ENDOSPERM Maize kernels with scarred endosperm can usually be recognized on the ear, even though the kernels are so closely arranged that only the crowns are visible. The scarred kernels are not so large as the normal kernels on the same ear and are commonly pinched ofif so that they are somewhat similar in appearance to kernels with “rough Men- tation”. The scarred character can more easily and certainly be recognized upon examination of the abgerminal surface of the kernel. Its external appearance is that of a scar left after the healing up of a deep wound. •Assistant Professor of Botany, Department of Botany, University of Missouri, as- sociated with the Department of Field Crops, Missouri Agricultural Experiment Station, in the genetic studies of corn, carried on by this Department. 4 Missouri Agr. Exp. Sta. Research Bulletin 52 In Fig. 1 are shown camera lucida drawings of a number of kernels with scarred endosperm. In Fig. 2 are similar drawings of two scarred kernels from which the pericarp overlying the abgerminal surface was removed. The nature of this endosperm character can best be seen from the drawings. It is an irregular cavity in the endosperm on the abgerminal side of the kernel. The cavity consists in a crater-like ex- cavation near the crown with divergent and often branched furrows extending towards the base of the kernel. The pericarp over the crater-like excavation near the crown nearly always collapses and causes the kernels to have a rough indentation. Occasionally a kernel is found with the pericarp over the crater of the cavity in the form of a blister. Scarred Endosperm and Size of Kernel. — Scarred kernels are uniformly smaller than normal kernels. In Fig. 3 is shown a crown view of a series of representative normal kernels (upper row) and scarred kernels (lower row) which were taken from the same ear. In Fig 4 is shown the same series of kernels, as in Fig. 3, but from the side. These figures show in a general way the relative differences in size between normal and scarred kernels of maize. Scarred Endosperm and Thickness of Kernel. — The most con- spicuous size difference between normal and scarred kernels is in the thickness of the kernels. Thickness here refers to the distance between the germinal and abgerminal surfaces of the kernel. The thickness of each kernel of individual ears segregating normal and scarred ker- nels was measured by using a sliding caliper rule and tabulated as shown in Table 1. Readings were made to the nearest one-half millimeter. The measurements were made by clamping the caliper over the end of the kernel at a uniform distance from the crown. The distributions given in Table 1 show that for each ear the nor- mal kernels are thicker than those with scarred endosperm. The mean thickness of normal kernels from individual ears varies from 3.856 to 5 .722 millimeters. The mean thickness of the scarred kernels from the same ears varies from 3.100 to 5.360 millimeters. The mean dif- ference in thickness of the normal and scarred kernels from the ears studied varied from 0.295 to 0.823 millimeter. The mean thickness of the normal kernels of the eight distributions listed in Table 1 con- sidered collectively is 4.500 — 0.144 millimeters. The mean thickness of the scarred kernels of these distributions is 3.926^0.258 millimeters. The normal kernels are 0.574 ± 0.295 millimeter thicker than the scarred Scarred Endosperm and Size Inheritance in Maize 5 kernels. In Fig. 5 are given curves which show graphically the vari- ation in thickness of the normal and scarred kernels. These curves represent the total frequencies of the distributions listed in Table 1. Scarred Endosperm and Weight of Kernel. — The kernels of each ear studied were weighed individually to the nearest milligram and tabulated as shown in Table 2. In every case the mean weight of the normal kernels is higher than the mean weight of the scarred kernels from the same ear. The mean weights of the normal kernels from the ears studied varied from 251.92 to 341.10 milligrams. The mean weights of the scarred kernels from the same ears varied from 232.70 to 329.60 milligrams. The differences in the means of the in- dividual ears varied from 1.24 milligrams for ear 1243-2 to 19.13 milli- grams for ear 1238-16. In order to obtain a general expression of the mean weights of the normal and scarred kernels the distributions in Table 2 are considered collectively. The mean weight of the normal kernels from the eight ears is 274 milligrams and the mean weight of Fig. 5. — Variation in thickness of normal and scarred kernels. 6 Missouri Agr. Exp. Sta. Research Bulletin 52 the scarred kernels is 259.57 milligrams. This is a difference of 14.43 — 1.29 milligrams. In Fig. 6 are curves of variation in weight of nor- mal and scarred kernels when the eight distributions of Table 2 are considered collectively. In many respects these curves are similar to those for thickness of kernel given in Fig. 5. The normal and scarred kernels respectively of each ear were weighed en masse with the results given in Table 3. From these total weights average kernel weights were obtained that do not involve the errors due to the separate weighing of the individual kernels. The av- erage kernel weights are in fairly close agreement with the mean weights as given in Table 2. The normal and scarred kernels from ear 1243-2 differ only slightly in average thickness and have approximately the same kernel weight. The mean weight of the normal kernels is given in Table 2 as 1.24 milligrams greater than that of the scarred kernels. The average weight, however, of the normals was found to be 1.36 milligrams less than the average weight of the scarred kernels. For the other ears the average weight of the normal kernels varies from 2.17 to 20.63 milli- grams heavier than the scarred kernels taken from the same ear. The Fig. 6. — Variation in weight of normal and scarred kernels. Numbei Descripi 90 1 450 Total 1 Mean Difference 1238-16 Normal 1 384 288.07 Scarred — — 139 268.94 19.13 1243-2 Normal 424 233.94 Scarred — — 145 232.70 1.24 1243-8 Normal 1 340 312.12 Scarred — — 121 300.17 11.95 1243-13 Normal 301 260.86 Scarred — — 128 249.84 11.02 1243-7 Normal __ 181 341.10 Scarred — — 24 329.60 11.50 1245-6 Normal 422 251.92 Scarred — — 128 23703 14.89 1245-7 Normal 58 287.93 Scarred — — 25 284.82 3.11 1245-8 Normal 85 255.65 Scarred — — 36 247.22 8.43 Total Normal 1 1 2195 274.00 Scarred -- — 746 259.57 14.43 Table 2. — F 2 Kernels of the Cross Normal X Scarred Thickness of Kernels in Millimeters Pedigree Number Description 1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5 5.0 5.5 6.0 6.5 7.0 7.5 Total Mean Difference 1238-16 Normal __ 1 4 10 43 93 77 73 32 27 17 10 387 4.637 .572 Scarred - 1 5 5 11 28 32 25 17 8 5 1 - - 138 4.065 1243-2 Normal 1 28 95 167 54 37 23 11 3 419 4.123 .332 Scarred - 1 2 5 28 41 39 12 7 6 5 146 3.791 1243-6 Normal 3 12 26 30 15 17 29 23 2 157 5.070 .815 Scarred - - - 1 3 9 13 7 10 4 1 1 - 49 4.255 1243-8 Normal 5 23 69 62 69 62 41 12 3 346 4.860 .484 Scarred - - 4 5 5 10 26 26 19 13 9 117 4.376 1243-13 Normal 1 13 43 86 72 49 16 14 9 3 __ 306 4.433 .823 Scarred - - 11 13 25 25 15 14 13 6 1 123 3.610 1245-6 Normal 3 104 118 102 33 22 18 9 6 3 418 3.856 .756 Scarred "I 6 15 29 32 27 5 4 3 2 1 125 3.100 1245-7 Normal 3 4 2 15 15 19 8 2 3 71 5.655 .295 Scarred _ 1 1 4 6 7 0 4 2 25 5.360 1245-8 Normal 1 1 7 14 19 23 18 5 __ 88 5.722 .379 Scarred 1 3 12 11 6 2 - - 35 5.343 Total Normal 1 9 163 338 548 337 294 202 173 96 28 3 2192 4.500 Scarred _ — "I ~8 37 58 104 141 132 95 87 57 28 8 2 — 758 3.926 .574 Scarred Endosperm and Size Inheritance in Maize 7 data in Table 3 show a greater difference in weight of normal and scarred kernels for some ears than the data in Table 2. Table 3. — F 2 Kernels From the Cross Normal X Scarred Noi -mal Kerr els Sea rred Kerr lels Difference Total Average Total Average Average Pedigree Num- weight kernel Num- weight kernel kernel Number ber in mgs. weight in mgs. ber in mgs. weight in mgs. weight in mgs. 1238-16 387 108880 281.34 138 37700 273.19 8.15 1243-2 442 102325 231.50 124 28875 232.86 —1.36 1243-7 185 52200 337.57 25 8200 328.00 9.57 1243-8 345 107775 312.10 119 36200 304.20 7.90 1243-13 299 80850 270.40 126 32240 255.87 14.53 1245-6 420 89750 213.69 126 24325 193.06 20.63 1245-7 60 17650 294.17 25 7300 292.00 2.17 1245-8 87 21750 250.00 35 8650 247.14 2.86 Total 2225 581180 261.20 718 183490 255.56 5.64 INHERITANCE OF SCARRED ENDOSPERM The factor pair for scarred endosperm is designated by the sym- bols S c s c . F x Generation. — F x kernels from the cross scarred x normal , or its reciprocal, have normal endosperm. F 2 Generation. — When F x plants are self pollinated, ears are produced which have normal and scarred kernels in ratios approximat- ing 3 : 1. In Table 4 are recorded the numbers of normal and scarred Table 4. — F 2 Kernels oe the Cross Normal X Scarred Pedigree Normal Scarred Total Ratio per 4 Numbers kernels kernels 1238-16 390 138 528 2.955 : 1.045 1243-2 442 124 566 3.124 : 0.876 1243-7 185 25 210 3.524 : 0.476 1243-8 345 119 464 2.974 : 1.026 1243-13 299 126 425 2.814 : 1.186 1245-6 424 128 552 3.072 : 0.928 1245-7 60 25 85 2.828 : 1.172 1245-8 87 35 122 2.853 : 1.147 Total observed 2232 720 2952 3.026 : 0.974 Total expected 2214 738 2952 3.000 : 1.000 Deviation 18 ± 15.88 8 Missouri Agr. Exp. Sta. Research Bulletin 52 kernels taken from eight ears of F x plants that had been self pollinated. The ratios of the individual ears vary from 2.814 : 1.186 to 3.524 : 0.476. The average ratio for all the kernels from the eight ears is 3.026 : 0.974. The total numbers observed were 2232 normal and 720 scarred kernels. This is a deviation from the expected distribution of 18 — 15.88 kernels. F 3 Generation. — A field planting under family number 1238 was made from a self pollinated ear that segregated kernels with scar- red endosperm. Twenty-one F 2 plants were grown to maturity. Three of these were wholly pistillate plants. The remainder were self pol- linated and produced ears with kernels as indicated below : Kernels all Normal and All scarred Normal Scarred kernels Kernels Observed 6 9 3 Expected 4.5 9 4.5 Deviation 1.5 0 — 1.5 These numbers are small but are in close agreement with expecta- tion. SUMMARY AND DISCUSSION Scarred is a new endosperm character in maize which consists in an irregular cavity in the endosperm on the abgerminal side of the kernel. Kernels with scarred endosperm usually have a rough indenta- tion. Scarred kernels have been compared in thickness and weight with normal kernels and it is evident both from the general appearance of the kernels and from the data given in this paper that scarred kernels are smaller than the kernels with normal endosperm. Scarred endo- sperm is inherited as a simple Mendelian recessive character. Cor- related with scarred endosperm is a difference in size of kernel that is apparently due to the same factor. Emerson and East (1913) found in their study of quantitative characters in maize size differences, such as height of plant, length of ear, and size of kernel, to be due to multiple factors. Such quantitative characters, however, are not always due to multiple fac- tors. A difference in size, or in any quantitative character, between certain individuals may be due to multiple factors, but a similar size or quantitative difference between certain other individuals may be due to a single factor. Thus differences in height of plants are com- monly due to a number of factors, but a large number of height dif- Scarred Endosperm and Size Inheritance in Maize 9 ferences in maize plants have already been found that are due to single factors. As examples may be mentioned dwarf and anther ear (Emer- son, R. A., and Emerson, S. H., 1921), brachytic (Kempton, 1920) and others from the cultures of the writer and other workers in corn. By inter crossing these different types, progenies can be produced which segregate a number of factors for height of plant, and size inheritance becomes quantitative. The same principle may be applied to other quantitative differences. Scarred endosperm represents a difference in size of kernel which is quantitative, but due apparently to a single factor. In this re- spect it is similar to size differences in seeds of beans observed by Johannsen (1913). In one of his pure lines Johannsen found a mu- tant with a longer seed than the parent stock. Seed length of this mu- tant bean was found by Leitch (1921) to be inherited as a single Mendelian character. Johannsen (1913) also found in his cultures a broad bean which, when crossed with the type, gives an F 2 progeny of 1 type : 2 intermediate : 1 broad. It is reasonable to expect that maize plants will be found with other quantitative differences than height of plant and size of kernel that are inherited as simple Mendelian characters. 10 Missouri Agr. Exp. Sta. Research Bulletin 52 ACKNOWLEDGMENTS The writer is indebted to George T. Kline for the drawings in Figs. 1 and 2, and to James F. Barham for the photographs in Figs. 3 and 4. LITERATURE CITED Emerson, R. A., 1920. Heritable Characters of Maize. II. Pistillate Flowered Maize Plants. Jour. Heredity 11 : 65-76. Emerson, R. A. and East, E. M., 1913. The inheritance of quantita- tive characters in maize. Neb. Agr. Exp. Sta. Research Bui. 2. Emerson, R. A. and Emerson S. H., 1921. Genetic interrelations of two andromonoecious types of maize : dwarf and anther ear (In manuscript). Johannsen, W., 1913. Elemente der exakten Erblichkeitslehre. Ver- lag von Gustav Fischer, Jena. Zweite Auflage: 652-654. Kempton, J. H., 1920. Heritable characters of maize. III. Brachytic Culms. Jour. Heredity 11 : 111-115. Leitch, I., 1921. A study of the segregation of a quantitative char- acter in a cross between a pure line of beans and a mu- tant from it. Jour. Genetics, 11 : 183-204. *#*• Fig. 1. — Maize kernels with scarred endosperm. Fig. 2. — Maize kernels with scarred endosperm. The pericarp has been removed from these kernels to show the nature of the scarred endosperm. 4.^6^ inaii ; fe i 8 4 I Fig. 3. — Kernels with normal endosperm (upper row) and scarred endosperm (lower row) from the same ear. Fig. 4. — Kernels with normal endosperm (upper row) and scarred endosperm (lower row) from the same ear. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 53 THE RELATION OF TEMPERATURE TO BLOSSOMING IN THE APPLE AND THE PEACH (Publication Authorized August 18 , 1922 ) COLUMBIA, MISSOURI AUGUST, 1922 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY P. E. BURTON H. J. BLANTON St. Louis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION J. C. JONES, PH. D., LL. D., PRESIDENT OF THE UNIVERSITY F. B. MUMFORD, M. S., DIRECTOR STATION STAFF AUGUST, 1922 AGRICULTURAL CHEMISTRY RURAL LIFE O. R. Johnson, A. M. S. D. Gromer, A. M. E. L. Morgan, A.M. C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, Ph. D. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. Sieveking, B. S. in Agr. A. M. Cowan, A. M. AGRICULTURAL ENGINEERING J. C. Wooley, B. S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. E. F. Hopkins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale, B. S. in Agr. W. W. SwETT, A. M. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. ENTOMOLOGY Leonard Haseman. Ph. D. K. C. Sullivan, A. M. O. C. McBride, B. S. in Agr. FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm. A. M. L. J. Stadler, Ph. D. O. W. Letson, B. S. in Agr. Miss Regina Schulte* Ben H. Frame, B. S. in Agr. HORTICULTURE V. R. Gardner, M. S. A. H. D. Hooker, Tr., Ph. D. J. T. Rosa, Jr., Ph. D. F. C. Bradford, M. S. H. G. Swartwout, B. S. in Agr. POULTRY HUSBANDRY H. L. Kempster, B. S. Earl W. Henderson, B.S. SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M W. A. Albrecht. Ph. D. F. L. Duley, A.M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr Richard Bradfield, Ph. D. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S.. Treasurer Leslie Cowan, B. S.. Secretary S. B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B.. Agricultural Editor J. F. Barham. Photographer Miss Jane Frodsham, Librarian. E. E. Brown, Business Manager. In service of U. S. Department of Agriculture. Relation of Temperature to Blossoming in the Apple and the Peach F. C. Bradford Observation of the responsiveness of plants to certain tem- peratures and of the poleward progress of vegetative activity more or less concurrently with the advance of warm weather led to the formulation many years ago of the doctrine of thermal constants. According to this theory a given stage in the development of any plant is reached when that plant has received a certain amount of heat, regardless of the time required or of the temperatures in- volved. For each plant and for each successive stage there was assumed to be a definite heat requirement, which generally re- ceived a mathematical expression in the form of so-called “heat units.” The unit was a degree on one of the several thermometer scales. However taken, temperature observations were generally reduced to terms of average or mean daily temperatures. The readings for all the days involved in the period in question were combined and the sum called the “thermal constant,” since it was assumed to be constant for the plant wherever and whenever grown. Units based on this system may be designated “day-de- grees” to distinguish them from the “hour-degrees” obtained by computation from hourly temperatures. Enunciated first, probably, by Reaumer 29 in 1735, the original conception has been modified by later workers. Adanson 2 pointed out that temperatures below freezing do not reverse plant activity and discarded them from his summations. Others used as bases of calculation some still higher temperature, at which vegetative activity supposedly began. The number of heat units for one day was obtained by subtracting the base temperature from the actual — mean or maximum as the case might be — thermometer reading for that day; consequently this has been called the “remainder system.” Later graduated values were assigned to various tem- peratures in recognition of accelerated growth at certain tempera- tures. The Livingstons, 22 particularly, have worked out a scale of weighted temperature values based on the principle of Van’t Hoff and Arrhenius, pointing out, however, that the purely physical proces- ses involved in growth are not governed by this principle. This system they called the “exponential.” More recently Livingston 21 4 Missouri Agr. Exp. Sta. Research Bulletin 53 evolved a third system called “physiological”, based on Lehen- bauer’s observations of root growth in maize at various tempera- tures. This system differs from the others in that it recognizes an optimum temperature above which the values assigned decrease. It is put forward, evidently, only as tentative, since Livingston states several qualifications of its applicability. Progress in plant physiology and particularly the recogni- tion that numerous factors influence plant growth have modified the original conception of thermal constants. Numerous objec- tions to the original conception have been stated aptly by Schimper, 31 and its rigid application is not often attempted, ex- cept in the use of growing season summations to characterize var- ious regions, as exemplified by Merriam’s 24 work on life zones, and Swingle’s 33 on the date palm. Ihne, 18 though inclined to con- sider phenological observations a measure of the weather, express- ly repudiates the assignment of definite thermal constants to any plant. Unfortunately there has survived a supposed connotation be- tween the old thermal constant conception and phenology which has retarded the study of phenological observations in ways that might otherwise have been attempted. One purpose of the present paper is to point out how the thermal constant conception, though full recognition be accorded the many objections to it, still may furnish, in connection with phenological observations, a valuable tool in the study of the response by plants to some of the factors composing climate. The comparative meagerness of the available data and the limited number of localities they represent preclude the possibility of formulating much that is conclusive, and this paper can be regarded only as suggestive of what might be at- tempted with abundant data for the same plant under many con- ditions. PHENOLOGY OF FRUIT TREES IN NORTH AMERICA Systematic observations on the blossoming of fruit trees at various points in North America began, perhaps, in 1817 when Bigelow 7 compiled a list of the dates of blossoming of the peach at various points from Fort Claiborne, Alabama, to Montreal. Early reports of the Smithsonian Institution and of the Army Signal Ser- vice, the forerunner of the present Weather Bureau, contain many scattered observations on blossoming dates. Following the recog- Relation of Temperature to Blossoming — Apple and Peach 5 nition of the importance of cross pollination in many fruits, blos- soming data have been published by a number of agricultural ex- periment stations. Since these were intended merely to show the overlapping in blossoming seasons of horticultural varieties, complementary temperature data are not ordinarily available and study of them shows little beyond: (1) a general similarity in se- quence of species and of varieties, (2) differences between places in the average lengths of the blossoming seasons and in the in- tervals between the blossoming of the several fruits, and (3) a general, though not uniform, recession of the blossoming dates with increased latitude and altitude. The Initial Date. — Those who have attempted to fit thermal constants to phenological observations on perennial plants have found much perplexity in fixing an initial date for temperature summations. Some have computed from leaf fall in the previous autumn, some from the coldest period of the winter (which is, in many cases, early in February) and some from the date when the average daily temperature rises above the freezing point. Others, as Fritsch, 15 have considered the precise date of little im- portance and have used January 1, as a matter of convenience. This is, perhaps, the most commonly used starting point. It is interesting that in many plants heat summations from this date to blossoming are not the same everywhere. Waugh 34 found in 1898 a general tendency for blossoming at lower summations in the north with the “American wild plum”, than in the south. More pronounced differences are evident in the summations for the Late Crawford peach at Pomona, California, and at Wauseon, Ohio, shown in Table 1. These are compiled from reports of the California Agricultural Experiment Station 9 ’ 10> 11 and from the Mikesell records. 25 Though the California figures are calculated from monthly means of daily maximum temperatures and the Ohio figures from daily maximum readings, errors arising from this cause must be slight in proportion; and the great differences shown in heat summations to blossoming actually exist. The highest summation from January to blossoming for any of the 27 years of the Ohio records is 925, considerably below the lowest shown here for Pomona and the minimum for Wauseon is barely more than one-fourth the maximum for Pomona. 6 Missouri Agr. Exp. Sta. Research Bulletin 53 Table 1. — Heat Summations in Day-Degrees (Maximum Above 43) eor the Tate Crawford Peach From January 1 to Blossoming in Ohio and in California. Year Wauseon, Ohio Pomona, California Date of first blossoming Day- degrees Date of first blossoming Day- degrees 1894 Apr. 17 745 Mar. 15 1222 1895 May 2 860 Mar. 2 1266 1896 Apr. 23 650 Mar. 20 2217 1902 Apr. 29 804 Apr. 1 2329 1903 Mar. 25 1895 Average (27 yrs.) 732 (5 yrs.) 1786 There may be, then, a considerable and consistent inequality in heat summations from January 1 to blossoming for the same fruit grown at points differing considerably in climate. This is in accord with observations of Palladin, 27 who records similar dif- ferences in many plants at Brussels and at Petrograd ; these dif- ferences were much more pronounced in the early blossoming than in the late blossoming plants. According to one view, ad- vanced by Linsser 20 the total heat requirements for any stage of plant development are not identical at all places, but their proportion to the total heat summation of the year is everywhere constant. In other words, there is supposed to be an acclimatiza- tion so that the same function may be performed with less heat at one point than at another, but require the same proportion to the total for the year at all points. This hypothesis appears un- tenable, in some cases at least, since this numerical ratio between the accumulation to ripening in the peach and the total for the year varies widely, from 48.8 per cent in Alabama to 83.1 in Massachusetts. 16 Furthermore, it does not take into account sea- sonal variations at the same point. Seeley 32 found great fluctua- tions from year to year in heat accumulations for various epochs in the Late Crawford peach at Wauseon, Ohio. In some years the minimum accumulation was 70 per cent of the maximum for the same period, in another 50 and in one, only 38. Evidently, then, Linsser’s constant or “aliquot” will not explain such differ- ences as those in summations in California and in Ohio from Janu- ary 1 to blossoming. Finally, since heat accumulations at one point may vary considerably from year to year, if carried to ex- Relation of Temperature to Blossoming — Apple and Peach 7 tremes this hypothesis implies a rather remarkable prescience in the plant. The differences between localities in heat accumulations to blossoming may be due in part to different normal temperature distributions. Price 28 demonstrated an acceleration in blossom- ing of peach and plum with high temperatures; yet his data show that the twigs held at the lower temperatures, though they re- quired a longer time for blossoming, actually received in some cases less total heat (in day-degrees). In some localities it is possible that even before blossoming there occur temperatures high enough to exercise an inhibitory effect, or, perhaps, the winters are not cold enough to make subsequent high temperatures fully effective. Twigs of ash and linden cut before the end of the rest period were kept by Weber 35 in a dormant state in a warm green- house for 15 months; at the end of this time most of the buds opened normally. VARIABILITY IN SUMMATIONS TO BLOSSOMING Since fruit bud differentiation in several fruits is first evident about July 1, it has been suggested that summations should be computed from this time to blossoming in the following spring. If this date is used for beginning computations on the apple and on the plum in Wisconsin, there is apparently a closer agreement from year to year than when summations are made from January 1 ; this has been interpreted to indicate July 1 as the proper starting point. 30 Much of this apparent agreement, however, is due to the tendency of meteorological elements to average alike over long periods. Summations calculated from July 1 to the following May 1, the approximate date of fruit bud opening, fit very nearly as closely as those figured to the dates of actual blossoming. The ratio be- tween the smallest summation and the largest is, in the Doney plum, 86.8 per cent; in the calendar summations, the check, it is 82.9 per cent. The very fact that the Doney plum came into blossom in 1904 with 4,494 day-degrees from July 1 while in 1901 the accumulation was 5,174, or 680 more, suggests that in the latter year some heat was received when it was ineffective in forwarding blossoms or was received in surplus quantities or that at some periods in the cycle heat is not a controlling factor. Indeed, something of the sort may be deduced from the data published by Sandsten. If the summations from successive dates 8 Missouri Agr. Exp. Sta. Research Bulletin 53 be averaged and their respective mean deviations determined, it becomes evident that the ratio betwen the average of the sum- mations and the mean deviation (in other words, the variability of the summations) changes and that it does not diminish in strict accordance with the tendency of meteorological values toward greater uniformity with increasing time. This is shown in Table 2 , arranged from Sandsten’s data for the Forest Garden plum, where the ratio just mentioned is designated the coefficient of variability. The “coefficient of variability” used in this paper is calculated from the mean, rather than from the standard, devia- tion, to lessen the effects of extreme variations. 37 Table 2. — Heat Summations in the Forest Garden Plum at Madison. Wis- consin, 1900-1905 Inclusive. (Compiled from data by Sandsten 30 ) To blossoming from Mean of summations Average deviation Coefficient of variability July 1. 4836 181 3.74 Aug. 1 3608 157 4.35 Sept. 1 2416 71 2.93 Oct. 1 1540 97 6.29 Nov. 1 893 80 8.96 Dec. 1 678 61 9.99 Jan. 1 667 70 10.49 Feb. 1 662 71 10.72 Mar. 1 653 63 9.64 Apr. 1 523 58 11.08 Sept. 1 (omitting Nov., Dec., Jan., and Feb.) 2159 50 2.31 The significance of these coefficients is more apparent if they are studied beginning with the coefficient for January 1. On either side of this (December and March) are lower values, signi- fying greater agreement in summations beginning at other times and suggesting that temperature accumulations from this date are not altogether effective in advancing blossoming. In summations beginning March 1 there is closer agreement; and the high varia- bility from April 1 may be interpreted to mean that advancement toward blossoming has begun, in some years at least, by that time. The difference in coefficients between November 1 and October 1 is striking and suggests that, beginning possibly about November 1, the temperatures received are not ordinarily effective. The greatest agreement in summations in the whole series is in those Relation of Temperature to Blossoming — Apple and Peach 9 dating from September 1 ; the low coefficient at this point is re- markable. If, however, the November, December, January and February temperatures are omitted the coefficient of variability in summations from this date is diminished even further. From August 1 and July 1, though these months in them- selves usually show relatively slight variability in their tempera- ture summations, the coefficients of variability are higher. Their low value as compared with that of March is due to the longer period covered, and their significance is probably slight. Here, then, though caution must be observed against infer- ring too much, there seems to be reason to consider tentatively for this fruit at Madison: (1) that temperature deficiency during July and August is not a limiting factor in any ordinary season, (2) that it becomes in some measure a limiting factor during Sep- tember and October, (3) that temperature is ineffective during November, December, January and February, possibly because there is not enough heat received to have any appreciable effect and (4) that about March 1 it again becomes for a time a determ- ining factor. Under other conditions, of course, very high temper- atures may become limiting. Even though these indications be true for the Forest Garden plum, caution should be exercised in applying them to another plant, for example a Japanese plum, in Wisconsin, or to the same plum in another locality. In other words, significant dates for phenological data may conceivably differ with the plant and with the locality. Angot 3 carried this idea of flexibility to the extreme, stating that the significant date varied not alone with the plant and the locality but also from year to year. Evidence is intro- duced in this paper indicating the variation of the significant date with plant and with locality; as to the yearly variation in the same plant and in the same locality the evidence is less clear. If, however, the chemical composition of the plant be considered to have an influence, as seems quite plausible, the effective date may vary as well. Furthermore, the stage of blossom development at- tained in the fall has been shown by Magness 23 to vary from year to year in the same variety. Consequently some variation in opening in the spring might be expected even in seasons that pre- sent practically the same temperatures. THE APPLE AND THE PEACH IN OHIO A study covering a number of seasons at one point has cer- 10 Missouri Agr. Exp. Sta. Research Bulletin 53 tain advantages over studies of a few seasons at many points. If, for example, the date when temperatures become effective be conceived to vary from place to place there is no satisfactory way of ascertaining this date from scattered observations unless the minimum accumulation to blossoming observed at any point be subtracted from the accumulations at other points and the dates computed from the day-degree remainders. This is, in effect, shap- ing the problem to fit the answer. In observations at one point over a series of years a certain degree of variation in other limit- ing factors is presumably reduced and if there is any validity in the thermal constant conception it should appear in observations of this sort. The publication by the United States Weather Bureau of the Mikesell records, 25 comprising phenological observations on num- erous plants at Wauseon, Ohio, over a period of 30 years, to- gether with daily meteorological records, makes possible a rather critical comparison of heat accumulations and phenological obser- vations. Since these records cover a longer period than any other avail- able data they are analyzed here and used in the study of the records of the Missouri Agricultural Experiment Station, which cover a much shorter time. Methods Used. — Heat accumulations may be measured in var- ious ways. In the work reported here the simple summations of temperature to blossoming, both maximum and mean, above sev- eral thermometric points, were computed. In addition one series was computed on the exponential system. For each series the yearly summations were averaged, the mean deviations from the averages determined and variability coefficients derived by divid- ing the mean deviations by the averages of the total accumula- tions. Occasional trials showed no material relative changes in coefficients resulting from the use of standard or mean deviations. The coefficients derived by the several methods are shown in Table 3. — Variability Coefficients of Heat Summations From January 1 to Blossoming at Wauseon, Ohio, as Calculated on Different Bases. Base System Apple Peach 32°F. Max. Remainder 7.69 8.26 43 °F. Max. Remainder 8.79 9.80 50°F. Max. Remainder 10.48 12.73 43 °F. Mean Remainder 12.20 16.06 40 °F. Max. Exponential 10.38 9.97 Relation of Temperature to Blossoming — Apple and Peach 11 The magnitude of the variability seems to vary inversely with the number of units involved ; for this reason the lower variabil- ity resulting from the use of 32° as the base point is not neces- sarily significant. Since this comparison did not show any base- point or system to be markedly superior to any other, the series based on maximum temperature above 43° was chosen for most of the further computations. This system appeared to give inter- mediate values and its results would be comparable with other work which has been based on the same temperature. Temperature observations taken according to conventional meteorological methods are not true records of plant tempera- tures and since the disparity between the two varies no correc- tions can be applied. For present purposes, however, since in sunshine twigs are generally warmer than the air, maximum air temperatures probably approximate those of the plant more closely than mean air temperatures. For other seasons or for other tem- perature ranges or in other climates mean temperatures might be preferable. However, even on summer stages for the peach at Wauseon, Seeley 32 found less variation in computations involving maximum than in those involving mean temperatures, though it is true this lower figure may be due to the larger number of units involved. Calculations. — Though it seems unlikely that heat deficiency is a limiting factor with apples or peaches in Ohio during the sum- mer months, computations were made from July 1 to blossoming the following year. From these figures the summations from other dates to blossoming were readily secured and the respec- tive variability coefficients determined. As a check on these, sum- mations to April 28 and to May 7, the average blossoming dates of the peach and of the apple, respectively, were similarly com- puted. These may be considered as measures of the independent variability of the weather and are valuable for comparison with the variability to the actual dates of blossoming. If heat accumulations are plotted vertically and a horizontal scale be adopted for time such that the spread of the projections of blossoming dates on the abscissa is equal in length to the spread of the projections of the accumulations on the ordinate, mathe- matical expression of the trend of the line connecting blossoming dates is possible. This is, in effect, done when the coefficient of variability in accumulations to blossoming is divided by the coef- ficient of variability to the average date of blossoming. With per- 12 Missouri Agr. Exp. Sta. Research Bulletin 53 feet uniformity in total day-degrees to blossoming the line would be horizontal ; with perfect uniformity in totals to a given date the line would be vertical. With an equal degree of uniformity in both it would be at a slope of 45°. This would be the case were coefficients of variability to blossoming and to average date of blossoming equal. In short, the numerical ratio obtained by dividing the coef- ficient of variability in summations to blossoming by the coef- ficient of variability in summations to average date is the tan- gent of the angle with the horizontal made by a smooth line con- necting the dates of blossoming. When this ratio is above one, the angle is greater than 45° and nearer vertical. In other words, the agreement is closer with the average date than in the total accumulations. Accordingly the figures in the columns headed “Tangent” in Table 4 are in reality tangents of slopes of lines connecting the graphical positions of the blossoming dates. The value 0.94 for the apple indicates a slope of approximately 43° for this line- practical neutrality. The value 0.51 indicates a slope of ap- proximately 27.° In the peach the 1.47 value indicates a slope of 56°, nearer vertical than horizontal. However, even without ex- pression in degrees, the tangents serve for comparison. Table 4. — Variability Coefficients of Heat Accumulations From Various Dates to Blossoming in the Apple and in the Peach at Wauseon, Ohio. Beginning date Apple Peach To actual blossom- ing To average date of blossoming Tangent To actual blossom- ing To average date of blossoming Tangent July 1 4.55 5.36 0.85 5.27 5.00 1.05 Aug. 1 5.63 6.45 0.87 6.22 6.34 0.98 Sept. 1 7.37 7.86 0.94 8.04 7.66 1.05 Oct. 1 9.33 11.11 0.84 9.71 10.59 0.92 Nov. 1 11.08 13.80 0.80 7.77 14.33 0.54 Dec. 1 13.65 15.47 0.88 9.55 16.94 0.56 Jan. 1 8.79 15.64 0.56 9.80 16.37 0.60 Feb. 1 8.48 15.78 0.54 11.34 16.73 0.68 Mar. 1 9.79 16.76 0.58 13.40 17.51 0.76 Mar. 15 7.93 15.51 0.51 14.91 15.60 0.95 Apr. 1 14.70 15.79 0.93 25.22 17.09 1.47 Relation of Temperature to Blossoming — Apple and Peach 13 Indications. — The lower the tangent the more significant, pre- sumably, is the coefficient for the corresponding period. Conse- quently the low variability coefficients for the summer and au- tumn months lose their weight and those of some of the later dates become more significant. An interesting difference between the apple and the peach is revealed by inspection of the tangents. The lowest value in the peach is in the summations figured from November 1 ; in the apple the lowest value is in the figures dating from March 15. This dif- ference seems to indicate that, under the conditions obtaining at Wauseon, high temperatures during winter are effective in promot- ing growth in the peach, but not in the apple. In the apple the period of effective temperatures seems more definitely fixed than in the peach. From January 1 to March 15 in the apple the tangents change but little, with the smallest fig- ure on March 15. It should be considered, however, that heat accumulations are small during this time and can affect the total variability but little. Other things equal, such changes as do occur as the date of summations moves backward should, through augmenting somewhat the total of day-degrees involved, reduce the variability. Therefore even the slight difference in tangents shown may be significant in the apple. The occurrence of the lowest figure on March 14 does not signify that the rest period ends then. It is, in a sense, an average date and means that, broadly speaking, advancement starts in half the years at that time. Consequently the end of the rest period must be earlier. In the peach, the succession of low values is in the reverse order and so far as this array affords evidence, the decrease from January 1 or December 1 may be due merely to the longer period and the consequently greater total of units involved. In either case, however, it seems clear that the peach becomes responsive to high temperature earlier than the apple and that its earlier blossoming is not necessarily due to a lower total requirement of heat. The low temperature of the ordinary winter at Wauseon would keep the trees dormant and microscopic study of buds for several years might show no development during this time, unless the period of investigation happened to include a mild winter. Dormancy of the peach in the north and in the south may be quite different; in the one case imposed by low temperature and in the other by the rest period. Johnston 19 found that the mois- ture content of peach buds in Maryland increases after January 1 14 Missouri Agr. Exp. Sta. Research Bulletin 53 in a definite relationship to the “sum of the effective daily mean temperature above 43°. ” Seasonal Differences. — Illustration of the difference between the two types of fruit is found in the graphs of heat accumula- tions from January 1 in 1890 and in 1912, shown in figure 1. These years are selected because they represent respectively the maxi- mum and the minimum accumulations of heat from January 1 to March 1. In 1912, with little accumulation of heat prior to April 1, the apple came into blossom very close to the peach both in time and in heat accumulations. This year, in fact, marked the Fig. 1. — Blossoming of peach and apple in years of maximum and of minimum accumulation on March 1, at Wauseon, O. lowest summation to blossoming for the apple. In 1890, with con- siderable heat accumulation throughout the winter months, the peach came into blossom earlier, but with substantially the same heat accumulation as in 1912. The apple, however, though its date of blossoming was nearly the same as in 1912, had re- ceived 300 day-degrees more of heat. This difference is about the same as the margin by which the accumulation to April 1 in 1890 exceeded that of 1912. This may be interpreted to signify that in 1912 practically all the heat received came when it was Relation of Temperature to Blossoming — Apple and Peach 15 effective, while in 1890 much of it was ineffective for the apple. Apparently the peach started from dormancy earlier than the apple in 1890, while in 1912 there was little difference, because the low temperatures held both dormant. Since these two winters were so widely different, it seems logical to infer that localities with average winters differing (as do these extreme types) would show for regions with mild winters a considerable spread in blossoming season between the peach and the apple — well known to be the case — while those with cold winters would show little or no difference. In extreme cases the peach and the apple may bloom simultaneously. This condition occurred at Wauseon in 1895, following a cold January and Feb- ruary and was closely approached in other years, invariably fol- lowing winters of small heat accumulations. Indeed, in 1912 which, according to Hedrick 17 was not an unusual blossoming sea- son, at Geneva, New York, blossoming in the apples began a day ahead of the peaches. The same phenomenon occurred in 1905 at Columbia, Missouri. The variations sometimes reported in the sequence of blossoming in other fruits may be due to simi- lar causes. If a certain validity be assumed for the thermal constant con- ception, the rather wide difference from year to year in the sum- mations from any given date to blossoming suggest that the higher figures may be due to accumulations occurring during per- iods when they are ineffective or in greater quantities than can be fully effective — and of course other factors than temperature may intervene. To facilitate comparison, data for the years of maxi- mum and of minimum accumulations from January 1 to blossom- ing in the peach and in the apple are arranged in Table 5. It is interesting and significant that these years are not identical for the two fruits, only three duplications occurring. In both fruits the minimum accumulations from January 1 to blossoming average practically the same in relation to the maximum, but here the sim- ilarity stops. In the peach the years of lowest summation from January 1 to blossoming generally succeed periods of consider- able accumulation in November and December, the accumulations preceding the years of minimum accumulation being in fact 141 per cent of those preceding the maximum. In the apple it is ap- parently a matter of indifference, since the November and De- cember accumulations preceding the minimum and the maximum years average practically the same. It should be stated that the 16 Missouri Agr. Exp. Sta. Research Bulletin 53 Table 5. — Analysis oe Accumulations in Years oe Greatest and oe Lowest Summations From January 1 to Blossoming in the Peach and in the Apple at Wauseon, Ohio. (In day-degrees) Year Jan. 1 to blossoming Previous Nov. 1 to Jan. 1 Jan. 1 to Mar. 1 Jan. 1 to Mar. 15 Mar. 15 to blossoming: Peach Years of Minimum Summations 1910 632 363 14 112 520 1908 588 180 18 104 484 1900 627 305 86 100 527 1896 650 286 50 57 593 1891 594 296 125 142 452 Years of Maximum Summations 1902 804 184 52 144 660 1898 834 266 72 198 636 1895 860 222 31 45 815 1893 853 150 13 104 749 1887 870 195 80 213 657 Apple Years of Minimum Summations 1912 752 195 6 6 746 1908 791 302 18 104 687 1905 787 290 13 15 772 1896 779 286 12 57 722 1886 803 217 83 196 607 Years of Maximum Summations 1901 1005 259 42 66 939 1894 1217 323 124 324 893 1890 1052 296 294 327 725 1889 1056 308 70 148 908 1887 1079 195 80 213 866 Averages Peach Min. 618 286 59 103 515 Max. 844 203 49 141 703 Apple Min. 781 258 26 56 707 Max. 1081 276 122 215 866 Minimum in per cent of Maximum. Peach 73.2 141 120 73 73 Apple 72.2 93 21.0 26 82 Relation of Temperature to Blossoming — Apple and Peach 17 two years of greatest heat accumulation in November and Decem- ber were followed by crop failures in the peach, constituting two out of the three in the 30 years of record. These facts, together with the low variability coefficient in the peach summations from November 1 (Table 4) indicate that rather marked accumulations of heat in November and December have some influence in the forwarding of Late Crawford peach blossoms toward opening, but are not important in the King apple. Johnston 19 found that the relation between temperature ac- cumulations from January 1 and moisture content of peach fruit buds, though constant in any one year, varies from year to year; and that “certain conditioning influences that are operative dur- ing or preceding dormancy apparently ‘predetermine’ the exact re- lationship between air temperature and the moisture content of the buds for the period following dormancy.” The averages in Table 5 show an excess of heat during Jan- uary and February of the years of minimum accumulation for the peach, but inspection of the detailed figures shows that this is of doubtful significance, since it is due to a high value in one year only. The low ratio of the minimum to the maximum years (21 per cent) in the apple, however, apparently signifies that the apple is unresponsive at this time and that heat accumulations during this period merely swell the total without having any marked effect in advancing the blossoms. The same negative re- lationship appears in the figures to March 15 for the apple (16 per cent) while the figures for the peach change markedly and assume the same relationship as the total accumulations. The similarity in the relationship in the peach of the years of maximum to those of minimum accumulations from January 1 to blossom- ing, from January 1 to March 15 and from March 15 to blossom- ing suggests that the same influences are operative during all three periods; in other words, that development is progressing. In the apple the change at this time is abrupt — from 28 to 86 per cent — the accumulations from March 15 to blossoming being more nearly alike as between maximum and minimum years than those from January 1 and closer than in the peach — 86 as compared to 73 per cent. Assuming, for the reasons given above, November 1 to mark the commencement of possible effective temperatures in advanc- ing the peach toward blossoming and March 15 for the apple, data for the five years of maximum summations for these respec- 18 Missouri Agr. Exp. Sta. Research Bulletin 53 tive periods in each fruit are assembled in Table 6 to show their relation to the temperatures of October, September and August preceding. Tabus 6. — Temperature Summations From Date oe PossibeE Eeeectiveness in Relation to Temperature oe Previous Months, at Wauseon, Ohio. Peach Apple Year Nov. 1 to blos- soming Oct. Sept. Aug. Year Mar. 15 to blos- soming Oct. Sept. Aug. Years of Minimum Accumulation 1911-12 879 554 979 1203 1910-11 669 647 979 1245 1910-11 782 647 979 1245 1908-09 694 699 1202 1249 1907-08 768 380 932 1190 1907-08 687 380 932 1190 1906-07 878 326 1154 1326 1901-02 706 740 1063 1350 1890-91 890 502 839 1213 1885-86 707 430 952 1075 Years of Maximum Accumulation. 1897-98 1100 882 1270 1211 1900-01 939 940 1174 1448 1894-95 1082 620 1109 1361 1893-94 893 597 1048 1317 1893-94 1068 597 1048 1317 1888-89 908 476 958 1289 1891-92 1071 587 1201 1254 1887-88 903 460 990 1284 1883-84 1104 378 866 1137 1883-84 892 378 866 1137 Averages Min. yrs. 839 481 977 1235 693 579 1026 1222 Max. yrs. 1085 613 1099 1256 907 550 1072 1295 Minimum in per cent of Maximum. Peach 77 78 89 98 76 105 96 94 These figures suggest, though not very strongly, a tendency toward an association between lower temperatures in October and a low summation from November 1 to blossoming in the peach. In the apple there is little or no appearance of any relationship. This difference may possibly be associated with some effect of the high October temperatures in prolonging or of the low temperatures in breaking the rest period in the peach, while in the apple at this time they have, ordinarily, no apparent effect. However, since rainfall in September is likely to be important in connection with September and October temperatures, no clear evidence is afforded by the data in Table 6 as to the effects of October temperature, though the essential similarity in September and August summa- tions indicates that temperature variations in these months have little effect on these fruits in this locality. Relation of Temperature to Blossoming — Apple and Peach 19 MISSOURI RECORDS Rather complete phenological records of numerous varieties of apples and peaches were kept at the Missouri Agricultural Ex- periment Station from 1905 to 1918 inclusive, with the exception of the blossoming records for 1910. This was an early season and the records show most varieties in full bloom on March 28 but the dates of first blossoming are not recorded ; consequently this year is omitted from calculations reported here. Through the kindness of Mr. George Reeder, of the United States Weather Bureau, temperature records for the period cov- ered by the phenological data have been made available. These observations were made at the Weather Bureau office, about one- fourth mile from the University Orchard in which the phenologi- cal observations were taken. An interesting commentary on the hazards of peach growing in this section is the appearance of blossoming dates for peaches for only 8 of the 13 years of the record. Since the observations were made with considerable care it is safe to presume that no blossoms appeared in other seasons during this period. Com- pared with the 27 crops in 30 years at Wauseon, Ohio, and with the uninterrupted, though brief, sequence reported from Pomona, California, they suggest that this particular section may be termed a no-man’s land for the common varieties of peach, being sub- jected to the hazards of both northern and southern types of winter injury, (extreme cold and untimely warm weather respec- tively) while regions north and south are subject ordinarily to only one form. Because of the scarcity of data no attempt is made here to study extensively the climatic relations of the peach in central Missouri. The comparative brevity of the period for which data are available at Columbia increases the difficulty of formulating any hypothesis as to the periods of effective temperatures. Similarly, though data are available for a considerable number of varieties, the brevity of the record for each makes varietal comparisons rather uncertain. However, some generalizations seem safe. The warmer winter months at Columbia make the average heat summations up to the date of blossoming greater than those at Wauseon, though the difference is not so marked as that between California and Ohio for the peach. Since the comparison in Table 7 between summations at blossoming at Columbia and at Wauseon is be-, tween the King apple at Wauseon and the Fameuse at Colum- 20 Missouri Agr. Exp. Sta. Research Bulletin 53 Table 7. — Average Temperature Accumulations (Max. Above 43°) From January 1 to Blossoming in the Apple at Wauseon, Ohio, and at Columbia, Mo. Wauseon, Ohio Columbia, Missouri To King Apple Fameuse Apple February 1 36 122 March 1 72 261 March 15 127 389 April 1 254 646 Blossoming 912 950 bia, the actual difference in any one variety would be somewhat greater. It is interesting that Fameuse blossomed in 1895, ap- parently a normal season, on April 1 at Paso Robles, California, with a day-degree accumulation of 1421 from the first of January 9 ; in 1902 the blossoming at Pomona, California, was on April 5 with an accumulation of over 2300 day-degrees 10 and in 1903 on April 15 with an accumulation of about 2273 day-degrees. 11 Varietal Differences. — Of the varieties for which data are available for all the years of record, Minnesota, Fameuse and Pri- mate are the earliest blossoming; Rome, Ralls and Ingram the latest. Data are presented in Table 8 showing the coefficients of variability in summations to blossoming in these varieties from dif- Table 8. — Variability in Day-Degree Summations From Various Dates to Blossoming in the Apple at Columbia, Mo., Computed From Maximum Temperatures Above 43°F. Oct. 1 Nov. 1 Dec. 1 Jan. 1 Feb. 1 Feb. 15 Mar. 1 Mar. 15 Apr. 1 Early blos- soming Minnesota 7.82 8.95 8.96 9.41 8.98 8.21 8.68 16.78 Fameuse 7.91 8.21 8.32 9.15 9.15 7.99 7.69 13.45 Primate 7.48 8.81 9.28 10.90 10.24 9.51 10.41 15.56 Av. 7.77 8.66 8.85 9.75 9.46 8.57 8.93 15.26 Weather Late blos- 8.33 11.87 16.58 18.11 20.89 20.09 22.46 18.41 soming Rome 7.61 9.27 9.27 10.32 10.49 10.18 9.82 8.41 29.65 Ralls 7.84 9.01 10.42 10.14 11.21 11.12 11.23 8.69 23.79 Ingram 7.58 9.01 7.68 8.15 8.63 9.60 8.76 9.62 27.74 Av. 7.68 9.10 9.12 9.54 10.11 10.29 9.94 8.91 27.06 Weather 6.65 8.50 9.46 9.37 10.62 9.89 9.88 7.01 15.15 Relation of Temperature to Blossoming — Apple and Peach 21 ferent dates on the 43° maximum basis. The variability in the weather to average data of blossoming as compared with the Ohio figures is generally greater in the early blossoming varie- ties and lower in the late blossoming. Much of this difference may be attributed to the smaller number of years considered, since 1912 was marked by great deficiency in temperature until near the av- erage date of blossoming for the early varieties, but was more nearly normal by the average date for the late blossoming var- ieties. Omission of this year from the record would reduce the variability in the summations to the average date of the early blossoming varieties very materially. The lower variability in the Columbia figures to the average date for the late blossoming varieties may be due to the greater number of day-degrees involved or it may be accidental. The probable error of the mean from January 1, is, for Columbia —40, as compared with —21 for Wau- seon. As they stand, the figures in Table 8 show, though not at all clearly, the same general tendencies in the early blossoming varieties as those appearing in the Wauseon data, with the ap- parently significant date earlier. Those for the late blossoming varieties, however, show no agreement greater than that in the weather to their average date of blossoming. The drop to 8.91 on March 15 might be significant were it not for the even lower figure (7.01) for the weather check. Though the low value of the latter is obviously accidental, it precludes the attachment of any significance to the former. Another way of comparing these two groups of apple varieties is through coefficients of correlation between accumulations and the date of blossoming, somewhat after the manner used by Aoki and Tazika 4 in the sweet cherry. In this case any relationship would be shown by a negative correlation. As shown in Table 9 the correlation, wherever there is one, is stronger in the early Table 9. — Coefficients of Correlation Between Heat Accumulations (Above 43°, Max.) and Date of First Blossoms in Apple at Columbia, Mo. Period of Early blossoming Late blossoming accumulation varieties varieties January 0.174±0.13 0.168±0.18 February — 0.369±0.16 — 0.142±0.11 March — 0.856±0.05 — 0.510±0.14 February 15— March 15 — 0.473±0.15 —0.065 22 Missouri Agr. Exp. Sta. Research Bulletin 53 blossoming varieties. Since the time interval between the per- iods of accumulation considered and the blossoming is shorter in the early blossoming than in late blossoming varieties, there is less opportunity for disturbing variations in the unmeasured in- terval and the correlation would be expected to be greater in the former. However, even with this allowance, there seems some indication that the date of effective temperatures is earlier in the early blossoming than in the late blossoming varieties. Different Temperature Basis. — Since it seems possible that the late blossoming of some varieties may be due to lack of re- sponse to certain temperatures which are effective with the early blossoming varieties, variability coefficients based on a higher minimum, 50°, are presented in Table 10. Here, curiously enough in view of the Wauseon results, the variability for the early blos- soming varieties is generally decreased, though the variability of the weather is increased. The full significance of this is not clear though the study of the records for single years which follows may explain it in part. In the late blossoming varieties the varia- bility in summations to blossoming generally decreases somewhat, while that of the summations to the average date increases. The changes are too slight, however, to be indicative. One possibly significant change is in the figures for March 15 where the vari- ability increases enough to give the tangent a value of 0.7034. Of itself this is not sufficiently low to have much weight, but in con- Table 10. — Variability in Day-Degree Summations From Various Dates to Blossoming in the Apple at Columbia, Mo., Computed From Maximum Temperatures Above 50° F. Nov. 1 Dec. 1 Jan. 1 Feb. 1 Feb. 15 Mar. 1 Mar. 15 Apr. 1 Early blossoming Minnesota 10.55 8.45 8.31 8.59 9.81 10.59 15.96 Fameuse 10.04 8.78 9.47 8.61 7.45 6.92 13.37 Primate 9.42 9.38 8.95 7.81 8.41 8.30 13.12 Av. 10.00 8.87 8.91 8.34 8.56 8.60 14.15 Weather Late blossoming 14.44 20.69 21.45 24.22 24.09 25.86 23.17 Rome 11.51 9.34 8.96 9.50 9.34 8.94 8.42 34.14 Ralls 10.10 10.59 9.94 10.65 9.93 9.33 8.10 27.76 Ingram 10.61 8.69 7.72 8.46 9.36 8.89 10.65 31.73 Av. 10.74 9.54 8.87 9.54 9.54 9.05 9.06 31.21 Weather 8.73 9.80 8.28 10.10 10.81 10.66 12.88 21.07 Relation of Temperature to Blossoming — Apple and Peach 23 nection with the condition shown for this date in Table 8 it may have some meaning. Seasonal Differences. — Some interesting weather variations with related responses are shown by the graphs of yearly accum- ulations shown in figures 2, 3 and 4. These are grouped more or less at random, the chief aim being to present the years of peach blossoming in two diagrams. The first four years of the record are shown in figure 2. Two of the four, 1906 and 1907, were rather high in accumulations to March 15 and diverged widely from that time ; the 1905 curve Fig. 2. — Accumulations (in day-degrees) to blossoming at Columbia, Mo. shows a markedly low winter accumulation followed by rapid ad- vance; 1908 is noteworthy for steadiness of the accumulation from March 1. Another interesting relationship is the identity of ac- cumulations about March 15 of the two pairs of curves. The agreement in these pairs in the accumulations to blossoming for Rome and the agreement in summations from March 15 are re- markably close. The tangent in this case is 0.244. The agreement in Primate is even closer, the tangent being in fact 0.222, but the agreement is not within the same pairs as in Rome. In both cases 24 Missouri Agr. Exp. Sta. Research Bulletin 53 of blossoming at the lower summation the accelerating influence of high temperature is apparent in the steepness of the gradient. The 1906 and 1908 curves are rather close to parallel for some time and blossoming of Primate occurs at the same level on them. The Elberta peach blossoms at a lower level on the 1908 curve. These curves crossed about March 10 and accumulations then were identical, but the more rapid rise from that point in 1908 evidently had more effect on Elberta than on Primate. Compari- son of the 1905 and 1908 curves indicates the effect of the sharp rise after March 15 in hastening the development of the Primate blossoms. In 1905 the Primate apple blossomed ahead of the Elberta peach. The arrangement in the figure shows that this was due to Elberta being late in blossoming rather than Primate being early. This was the year of very little accumulation until after March 1. Apparently the cold weather held the peach dormant until high temperatures could become effective on the apple, as in Ohio in 1912, shown in figure 1. It should be stated, however, that a considerable amount of winter-killing of buds occurred during the winter of 1904-1905 and that the blossoming of Elberta as re- corded is doubtless later than it would have been with a full crop, since generally the more advanced buds are more readily killed. Furthermore, Chandler 13 mentions a mild form of winter injury which retards, but does not prevent blossoming. Even with this allowance, however, the closeness of Elberta and Primate is in- dicative of the influences mentioned. It is interesting that Morgan 26 working at Ithaca, New York, reported the apple to start development earlier in the spring than the peach but that the peach rapidly overtook it. This might well be the case if the investigation were carried on in such years as 1905 or where the common season resembles the 1905 season. On the other hand Drinkard 14 in Virginia reported more ad- vancement during the winter in the peach than in the apple. As- suming the peach to require higher temperatures than the apple it might start later than the apple in seasons that are cool, but with warm temperatures it starts before the apple. In figure 3 are presented curves for the remaining years for which peach blossoming dates are available, with that for 1912 added for comparison. The successive spring freezes of 1921 de- stroyed apple blossoms so extensively that blossoming records were not taken, consequently the curve for that year is not car- Relation of Temperature to Blossoming — Apple and Peach 25 ried beyond the blossoming of Elberta, which occurred before any damage had been inflicted and is therefore reliable. At first glance the high accumulations for Elberta in 1909 and 1911 are outstand- ing and apparently inconsistent. These years, however, were char- acterized by a considerable amount of winter killing of buds, the damage in 1909 in Elberta at Columbia amounting, according to Chandler, 12 to 97.3 per cent. Data are not available on the ex- tent of the damage to Elberta in 1911, but since it ranged from T* 4 os < t ci G, *< r~< I Fig. 3. — Accumulations (in day-degrees) to blossoming at Columbia, Mo. 29.8 per cent to 79 per cent in other varieties, it must have been considerable. These dates, then, represent the opening of only a very small portion of the total number of blossoms and these, presumably, those that were least advanced during the winter and would be the last to open in the spring. With these allowances, the line connecting the blossoming dates of Elberta would become nearly horizontal, signifying a rather close agreement in totals to blossoming. In Rome the differences in day-degrees at blossoming are in the same order as the differences on March 15 with one exception. 26 Missouri Agr. Exp. Sta. Research Bulletin 53 This is on the 1909 curve where Rome seems unduly late in blos- soming. Since yield records are not available for this variety the amount of bloom this year cannot be stated. That Rome was “out of step” in this case is shown by the fact that this was the only year in the record when Ingram blossomed ahead of Rome. If this were due to scarcity of crop so that the only blossoms ap- pearing were terminals — as is sometimes the case — this discrep- ancy would be explained. However, even with this allowance, the agreement is very little greater in summations from March 15 to May 1. 5 to 0£ ei >• 4 £ £ 1 i Fig. 4. — Accumulations (in day-degrees) to blossoming at Columbia, Mo. In figure 4 are shown graphs for years in which no blossom- ing dates for the Elberta peach are available; to these 1907 is added because of its close similarity to 1918 after March 1. These graphs are strikingly similar, excepting the 1912 curve. In 1913, 1916 and 1917 Primate appears to have been retarded by a week of cool weather in early April, though its extreme lateness in 1917 can be only partly explained in this way. Here again, sparseness of the crop may be a partial explanation, since study of the spurs Relation of Temperature to Blossoming — Apple and Peach 27 of this tree in the orchard shows very little blossoming in that year. The blossoming of Rome at a comparatively low accumula- tion in 1913 may be explained by the rapid rise in temperature subsequent to April 15. The evident retarding effect of the cold weeks in early April in 1913, 1916 and 1917 and the absence of influence of these weeks on Rome, though it is clearly responsive to high temperature, raises an interesting question. Since the buds of Primate, which was retarded, were more advanced at these periods than those of Rome, which apparently was not retarded, it seems quite possible that optimum temperatures vary as the buds advance toward opening. The decrease in the rate of the temperature rise in 1907 after Primate was in blossom and Rome presumably well advanced, seems to have had a retarding effect. There is a rather strong trend toward uniformity in summa- tions to blossoming in these curves, if 1912 be disregarded. This does not, however, necessarily signify that temperatures of Jan- uary and February are effective, since the accumulations are very much alike on March 1 or even on March 15. Here again, plot- ting the curves with March 1 as the starting point or, for Rome, March 15, secures much greater agreement. Considering all graphs shown, and making the allowances in- dicated, there is a rather marked tendency for uniformity in sum- mations from January 1 to blossoming, in the Elberta peach. Where the uniformity appears in the blossoming of the apples it is accompanied by an approximate uniformity in the accumula- tions at some date subsequent to January 1. Nowhere, however, is there clear evidence pointing to uniformity or difference in the end of the rest period between the early and the late blossoming apples. The graphs in figure 3, contrasting warm springs with a very cold spring, suggest a difference in the rest period. In Table 11 are assembled data showing the fluctuating dif- ference between 30 varieties of apple for all the years of record. Included in this are all the varieties for which data are complete; in most cases the records are from the same trees throughout. The seasonal difference in dates of first bloom (range) is shown to vary from 5 to 27 days and the average deviation from 0.97 to 4.75 days. That this difference between the first and last blos- soming variety is little more — or is even less — constant when ex- pressed in terms of heat is shown by comparison of the maximum range in day-degrees (519) with the minimum (187). The gen- 28 Missouri Agr. Exp. Sta. Research Bulletin 53 Table 11. — Variations in Blossoming Among Thirty Apple Varieties at Columbia, Mo. Days Day-degrees Aver. date Duration of bloom Aver. devi- Aver. accumu- Aver. devi- Aver. daily Year of bios. Range in earliest variety ation lation Range ation acc. dur- ing bios. 1905 99.6 24 22 2.85 898.7 519 69.5 21.6 1906 111.2 13 12 1.69 926.4 415 51.1 31.9 1907 87.9 22 21 3.14 886.8 323 48.4 14.7 1908 101.4 14 12 1.87 943.6 342 45.1 24.4 1909 111.2 18 13 3.32 1191.3 419 52.3 23.3 1911 104.7 18 20 2.89 1109.8 380 61.4 21.1 1912 115.7 8 16 1.25 766.0 193 30.0 24.1 1913 109.5 12 12 1.80 964.9 315 48.6 26.3 1914 111.2 11 10 1.59 976.1 359 51.7 32.6 1915 111.0 5 6 0.97 879.6 187 36.4 37.4 1916 107.9 13 13 1.47 1059.1 279 40.3 21.5 1917 111.7 20 8 2.81 1100.0 313 63.4 15.6 1918 100.3 27 33 4.75 1087.3 418 69.1 15.5 Av. 106.4 15.8 14.5 2.34 983.8 343 51.3 23.8 eral accelerating effect of high temperature is evident in the av- erage of the daily temperature accumulations during the three shortest ranges, 31.3,° and during the three longest, 17.3.° No constant on the basis used here will measure the dif- ference between blossoming in the earliest apple and the latest. It is quite likely that an exponential or a “physiological” system would measure this brief span more closely. It is, however, quite as probable that conditions before the blossoming of the earliest apples vary from season to season and that this event may find the late blossoming variety at various stages. When the early blossoming variety is not held back by unfavorable weather the late blossoming kind will lag behind ; when the early blossoming variety is retarded the difference will be less, other things equal. This indicates a difference in date of effective temperatures. How- ever, it is noteworthy that no matter how much the season is re- tarded and how small the range between the varieties, the early- blossoming kinds bloom first and the late-blossoming varieties bloom last. This indicates a difference in temperature require- ments. Relation of Temperature to Blossoming — Apple and Peach 29 Leaf and Fruit Buds Compared. — Apparently the opening of blossoms and the unfolding of leaves respond somewhat differ- ently to a given set of conditions. Bailey 5 says that in the southern states plum flowers “tend to appear wholly in advance of the leaves, and they are borne upon short stalks, or may be nearly or quite sessile. In the North, the flowers and leaves are gen- erally coetaneous, and the flower stalks are usually longer.” Bal- lard and Volck 6 report that spraying with nitrate of soda in Feb- ruary hastened the opening of flowers but not of leaf buds, in apples and pears. Table 12 shows the variability in summations to appearance of the first fully formed leaf at Wauseon, Ohio. Since the period of record is not identical with that for blossom- ing the figures are not strictly comparable. However, the dif- ferences between the values of the tangents in Tables 4 and 12 seem considerable enough to signify some difference between leaves and blossoms in their responses. In some years blossoms preceded leaves; other years showed the opposite condition. Table 12. — Variability in Day-Degree Summations (Maximum, Above 43°) to Date oe the Appearance oe the First Fully Formed Leae in the Apple at Wauseon, Ohio. Variability in Summations Variability in summations to Tangent beginning summations average date September 1 7.30 7.47 0.98 October 1 9.05 10.23 0.88 November 1 10.21 15.74 0.65 December 1 11.74 17.69 0.66 January 1 11.90 18.60 0.64 February 1 11.28 17.73 0.64 March 1 11.41 17.63 0.65 April 1 16.42 19.09 0.86 The data for Columbia record a somewhat different phase of vegetative development, namely, the opening of the leaf buds. Very rare indeed in these records is the case where the opening of the first blossom precedes the opening of the first leaf bud ; al- most invariably the leaf buds open before the blossoms. The mar- gin of difference varies, however. In three typical early blossom- ing varieties the average difference for 13 years is 7 days; in three typical late blossoming varieties it is 11 days. The late blossoming varieties blossomed on the average 13 days after the 30 Missouri Agr. Exp. Sta. Research Bulletin 53 opening of the first bloom ; their leaves appeared, on the average, 10 days after the opening of the first leaf bud. Evidence from Microscopic Examination. — Explanation of much of the lack of agreement among the variability coefficients of the several varieties represented in Tables 8 and 10 is found through microscopic examination of flower buds at various times during the winter. Typical photomicrographs of preparations made by Mr. V. R. Boswell are shown in Plates I, II and III. Oldenburg and Primate represent the earliest blossoming varieties; Rome, Daru and Cilligos the latest. The two last are included since they have been used extensively in the apple breeding work of the Missouri Station. Daru blossoms at about the same time as Ingram ; Cilligos is the latest blossoming of all varieties under observation. Plate I shows the stages reached by several varieties on Feb- ruary 2, 1920. Oldenburg is clearly more advanced than the other varieties. In the other cases, the correspondence between develop- ment on this date and the order of blossoming is not so close. Fameuse, the second earliest in blossoming, is no farther advanced than Daru, the second latest in blossoming. Gano and York, mid- season varieties, are apparently at the same stage as Cilligos, the latest of all. In Plate II are shown the stages on three dates, November 2, 1921, January 28, 1922, and February 20, 1922, for three varie- ties, Oldenburg, Primate and Wealthy. The first two are dis- tinctly early in blossominng; Wealthy might be classed either among the last of the early blossoming or among the earliest of the mid-season varieties. Here the advancement of Oldenburg in November is marked; this appears clearly to be a factor in its early blossoming since subsequent samples show relatively slight development. The other early blossoming variety, Primate, shows a quite different condition. Its November stage is not advanced; indeed, Daru, one of the latest blossoming, shows equal or greater pistil development on this date. Its changes through the winter, however, are notable and suggest that this variety has a factor producing early blossoming quite different from or more intense than that evident in Oldenburg. Wealthy, equally or more ad- vanced in early November, does not develop as rapidly through the winter. Plate III records the development of the buds in three late blossoming varieties sampled on the same dates as those of the Relation of Temperature to Blossoming — Apple and Peach 31 early blossoming varieties shown in Plate II. Daru, the second latest in blossoming of all the varieties shown, is among the more advanced on November 2. Its lateness is due apparently to its lack of responsiveness to temperatures with which Primate de- velops. Rome presents an anomaly in that it is perhaps the least advanced in November and apparently advances little or none to February 20; nevertheless it comes into blossom ahead of Daru and Cilligos. The winter of 1921-1922 in Columbia was mild in the sense that there was little very cold weather. However, as measured in day-degrees above 43° it was not warmer than the ordinary sea- son; the monthly accumulations from November to February in- clusive being respectively 345, 167, 92 and 178. November accum- ulations were below the average (426) and December above (39), January somewhat below the average (122) and February some- what above (139). The samples shown here, however, were gath- ered on February 20, when the accumulation for the month had reached 102 day-degrees only and before the warmest weather of the month. Consequently such development as is shown to be connected with temperature for this winter may be regarded as normal for this locality. Evidently, then, early blossoming in apples involves at least two factors : first, the stage of advancement reached at the ap- proach of winter, as exemplified by Oldenburg; second, ability to develop through the winter, as shown by Primate. Late blossom- ing, presumably, is due to the absence of both these factors or to the presence of strong inhibitors of the second. The ideal late blossoming variety as represented by Cilligos is backward in de- velopment in the fall and advances little through the winter. It is plausible that mixed inheritance of these factors gives the mid- season blossoming shown by the majority of commercial var- ieties, though Daru appears to have one factor for earliness despite its late blossoming. This seems the more likely since its crosses with Delicious now growing in the Experiment Station grounds include only very few late blossoming varieties, a smaller per- centage than those shown by the majority of the crosses involving Ingram, another late blossoming variety. The behavior of the late blossoming varieties indicates either a requirement of higher temperatures for advancement or the temporary presence of a development-inhibiting factor that is ab- sent in the early blossoming kinds. If late blossoming is due to 32 Missouri Agr. Exp. Sta. Research Bulletin 53 a higher temperature requirement, the difference between late and early blossoming kinds should be diminished by forcing in a. greenhouse. If late blossoming is due to the persistence of the rest period in some form these differences should decrease as the season advances. Table 13 shows the results obtained by forc- ing twigs of Primate (hypothetically without or over the rest period )and of Rome (hypothetically still in the rest period). The stages observed in the two varieties differ, but comparison is pos- sible. Though the buds started March 3 were kept in a cooler house than that used for the two earlier lots, enough cooler ap- parently to retard Primate, Rome started in a shorter time. The lower temperatures actually retarded the early blossoming variety more than the late blossoming. This, with the progressive short- ening of the period of forcing in Rome, indicates the rest period as a factor rather than a differential temperature requirement. Table 13. — Number oe Days Involved in Forcing Blossom Buds oe Primate and Rome Apples, 1922. Date forcing started Days to blossoms open Days to buds starting in Primate in Rome February 16 17 15 February 25 14 14 March 3 20 13 Comparison of Plates IV and V shows that the difference be- tween varieties are greater when they are forced in the green- house than when the buds develop in the orchard. This points in the same direction as the evidence just cited. Other Considerations. — Analysis of the records of 42 trees for which data are complete shows no relation between the date of terminal bud formation on shoots and the date of spur blossom- ing in the spring, the correlation coefficient being 0.085—0.04. It is possible, however, that comparison of trees under different cultural conditions might show a relation of this sort, though it is doubtful if it should not be considered an associated rather than a causative condition. Incidentally the relation to cross pollination of differences in blossoming may be mentioned. Comparison of the figures in the column headed “Range” with those in that headed “Duration of bloom in earliest variety” in Table 11 shows that in eight years of the thirteen recorded the earliest variety was out of bloom be- Relation of Temperature to Blossoming — Apple and Peach 33 fore the latest blossoming came in. In two years the date of last blossom in the one and of first bloom in the other were identical. In one only was the overlapping sufficient to ensure abundant cross pollination. In this section, then, when very early blossom- ing kinds are planted with very late blossoming kinds, cross pol- lination can be ensured only by a third variety, intermediate in blossoming season. This will provide pollination for the early blooming kinds with its first blossoms and for the late blooming with its last blossoms. Most of the commercial varieties grown in Missouri fall into the intermediate class in blossoming and may be counted on with safety so far as cross pollination is con- cerned. However, it is possible that the reputation of the Rome for light bearing in Missouri, though in Ohio it has not met that objection, is due to the greater extent of the blossoming season in Missouri so that Rome may in some seasons be in bloom alone while in Ohio the difference ordinarily would be less marked. Table 14 shows the blossoming dates of several peach va- rieties, selected to permit comparison with dates for the same varieties at points with winters considerably milder than those at Columbia. For compactness these are expressed in days of the year rather than of the month. Though the list for most years at Columbia is more extensive than those given for Alabama or California, the range in blossoming represented is less in every case. In other words, just as the blossoming of the apple in dis- tinctly cold sections has a narrower range than at Columbia, so the peach at Columbia has a narrower range than at points farther south. Cool weather during the peach blossoming season at Au- burn, Alabama, may have prolonged the season of 1911 to an un- usual length, but the normal blossoming range of the varieties named in this paper is apparently as great or greater than the maxi- mum recorded for Columbia. 36 The range shown for Pomona, California, is apparently normal for that point. The slight difference between all varieties at Columbia in 1907, the year of earliest blossoming for which an approximately complete record is available, indicates that the rest period as a factor in the date of blossoming in the peach is not operative here. This was a year of rather high temperature from January on. A considerably greater number of varieties than is here reported showed almost as close agreement in blossoming in 1921, when the season was even earlier than that of 1907. Any differences in the rest period which might be concealed by the retarding effect 34 Missouri Agr. Exp. Sta. Research Bulletin 53 of an ordinary winter on the earliest varieties should become evi- dent in these seasons of exceptionally high late winter tempera- tures, as soon as growth is possible. The third year of closeness in blossoming, 1906, was characterized by rather low accumulation during winter, with a rapid advance about the time of blossom- ing. The spread of the year of greatest range is due apparently to unequal winter-killing of blossoms and to the slow accumu- lation of temperature, which brought out minor differences in re- sponse to heat or merely delayed the opening of those varieties which had fewest buds. In warmer climates it seems quite pos- sible that these differences in blossoming are due to differences in the termination of the rest period, particularly since the Peen-to peaches there blossom much earlier than those recorded in Table 14, and almonds in January or February. Table 14— Peach Blossoming Dates (In Days oe Year) at Various Points. Variety Columbia, Mo. Au- burn Ala. Pomona Calif. 1905 1906 1907 1908 1909 1911 1914 1915 1921 1911 1894 1895 1902 1903 Alexander 98 102 82 90 101 95 78 76 8S Briggs Red 98 103 83 90 96 92 78 63 91 Carman 98 102 83 85 — 95 98 107 76 52 Champion 97 103 82 86 94 94 99 107 76 52 Chinese Cling 100 102 83 86 96 93 97 106 75 46 71 64 84 77 Crawford Early 102 82 80 96 93 96 108 — 35 74 62 91 84 Crawford Late 99 102 83 86 96 94 72 60 91 69 Elberta 98 102 82 85 95 92 97 107 74 45 Family Favorite 99 102 82 85 95 93 44 Foster 101 102 82 96 95 100 109 72 63 95 74 Globe 103 82 86 96 95 46 Heath Cling 97 104 82 84 95 96 103 72 66 91 69 Henrietta 102 103 82 88 97 65 — Lemon Cling 102 83 88 96 93 103 105 73 — Mayflower 95 100 105 — 57 Mountain Rose 98 107 75 — 76 60 91 77 Oldmixon Cling 98 102 82 86 96 97 94 108 — — 72 63 94 87 Oldmixon Free 98 102 m 85 95 95 97 84 66 95 74 Salway 98 103 83 85 96 92 103 107 76 46 73 66 61 91 Smock 91 103 82 86 95 101 104 — 52 79 63 — 77 Sneed 99 102 83 87 95 93 52 Susquehanna 102 103 83 86 — 96 50 72 63 — 69 Thurber 99 102 82 86 95 43 Yellow St. John 98 103 82 86 99 96 72 63 64 79 Range 12 4 2 11 8 10 10 5 3 23 14 17 35 23 Relation of Temperature to Blossoming — Apple and Peach 35 SUMMARY 1. The amount of heat, as measured in day-degrees, received by peaches from January 1 to the time of blossoming, varies with the season and even more with the locality. 2. The agreement in temperature accumulations to blossom- ing from year to year at any one place varies with the length of the time for which they are measured indicating that ordinary temperatures are not always effective or that temperature is not always a limiting factor. 3. Variability in temperature accumulations from various dates to blossoming at Wauseon, Ohio, follows different orders in the King apple and the Late Crawford peach, indicating that the latter is responsive to high temperatures when the former is not. 4. The average temperature accumulation from January 1 to blossoming in the apple is somewhat greater at Columbia, Mo. than at Wauseon, Ohio, but much less than at Pomona, Calif. 5. Varietal differences in blossoming at Columbia, Mo., in- dicate that the early blossoming varieties of apple become re- sponsive to ordinary temperatures earlier than the late blossom- ing. There are, however, some inconsistencies which are not ex- plained by any mathematical analysis attempted. 6. Microscopic examination of blossom buds indicates that there are at least two factors governing the season of blossom- ing at Columbia, Mo. Oldenburg blossoms early chiefly because the buds are well advanced in the fall, Primate because the buds develop through the winter. Daru is well advanced in the fall but does not develop through the winter and blossoms la-te. Cilligos is backward in the fall and does not advance through the winter ; it is the latest blossoming variety observed. The mid-season va- rieties apparently have a mixed genetic constitution in this respect. 7. Observations on branches forced in the greenhouse in- dicate that late blossoming is connected with rest period influ- ences rather than with differential temperature requirements. 8. Varietal differences in the peach at Columbia appear to be masked, but may become evident farther south, in the same manner as differences apparent in the apple at Columbia are masked farther north. Relation of Temperature to Blossoming — Apple and Peach 37 ACKNOWLEDGMENTS Acknowledgments are due Professors V. R. Gardner, H. D. Hooker, Jr., and W. J. Robbins of the University of Missouri; to Dr. E. J. Kraus of the University of Wisconsin, for valuable aid and suggestions; to Mr. George Reeder of the United States Weather Bureau, for the temperature records for Columbia; to Mr. V. R. Boswell of the University of Missouri, for assistance in computations, for the sections used in the study of bud develop- ment and for the photomicrographs ; and to those whose interest and care resulted in the accumulation of the phenological records for Columbia. Relation of Temperature to Blossoming — Apple and Peach 39 LITERATURE CITED 1. Abbe, C., U. S. D. A. Weather Bur. Bui. 36 (1905). 2. Adanson, Cited by Abbe. 3. Angot, A., Cited by Abbe. 4. Aoki, S. and Tazika. Y., Journ. Met. Soc. Japan. Apr. 1921, Abs. in U. S. D. A. Monthly Weather Rev. 49. 609 (1921). 5. Bailey, L. H., The Evolution of Our Native Fruits, p. 199, N. Y. (1898). 6. Ballard, W. S. and Volck, W. H., Journ. Agr. Res. 1 :437 (1914). 7. Bigelow, J., Amer. Jour. Sci. 1:76 (1820). 8. Bradford, F. C., Oreg. Agr. Exp. Sta. Bui. 129 (1915). 9. Calif. Agr. Exp. Sta. Ann. Rept. 1894-1895. p. 379. 10. Calif. Agr. Exp. Sta. Ann. Rept. 1902-1903. pp. 187-190. 11. Calif. Agr. Exp. Sta. Ann. Rept. 1903-1904. pp. 175-188. 12. Chandler, W. H., Mo. Agr. Exp. Sta. Res. Bui. 8 (1913). 13. Chandler, W. H., Proc. Am. Soc. Hort. Sci. 12:118 (1915). — 14. Drinkard, A. W., Jr., Va. Agr. Exp. Sta. Ann. Rept. 1909-1910. pp. 159- 197. 15. Fritsch, K., Cited by Abbe. 16. Gardner, V. R., Bradford, F. C. and Hooker, H. D., Jr., Fundamentals of Fruit Production, N. Y. (1922). 17. Hedrick, U. P, N. Y. Agr. Exp. Sta. Bui. 407 (1915). 18. Ihne, E., Ueber Beziehungen zwischen Pflanzenphanologie und Landwirt- schaft p. 9, Berlin (1909). 19. Johnston, E. S., Paper before Botanical Society of America. Toronto, Ont. Dec. 29, 1921. 20. Linsser, C., Cited by Abbe. 21. Livingston, B. E., Physiol. Res. 1:8 (1916). 22. Livingston, B. E. and Livingston, G. J., Bot. Gaz. 56:5 (1913). 23. Magness, J. R., Oreg. Agr. Exp. Sta. Bui. 139 (1916). 24. Merriam, C. H., U. S. D. A. Bur. Biol. Survey, Bui. 10 (1898). 25. Mikesell, T., U. S. D. A. Mo. Weath. Rev. Sup. 2 (1915). 26. Morgan, W. M., Thesis. Cornell Univ. 1902 — Cited by Wiegand, K. M., Bot. Gaz. 41:373 (1906). 27. Palladin, V. J., Plant Physiology. Eng. ed. by Livingston. Phila. (1918). 28. Price, H. L., Va. Agr. Exp. Sta. Ann. Rept. 1909-1910. p. 206. 29. Reaumur, R. A. F. de., Mem. Acad, des Sciences. 1735 p. 545. cited by Abbe. 30. Sandsten, E. P., Wis. Agr. Exp. Sta. Bui. 137 (1906). 31. Schimper, A. F. W., Plant Geography upon a Physiological Basis p. 37. Oxford, 1903. 32. Seeley, D. A., U. S. D. A. Mo. Weather Rev. 45:354 (1917). 33. Swingle, W. T., U. S. D. A. B. P. I. Bui. 53 (1904). 34. Waugh, F. A., Vt. Agr. Exp. Sta. Ann. Rept. 11:270 (1898). 35. Weber, F., Sitzungsber. d. Akad. d. Wiss. Wien. 125, 330 (1916). 36. Williams, P. F. and Price, J. C. C., Ala. Agr. Exp. Sta. Bui. 156 (1911). 37. Yule, G. U., Introduction to the Theory of Statistics, p. 146, London (1919). Relation of Temperature to Blossoming — Apple and Peach 41 Plate I. — Blossom buds of apple on February 2 , 1920 . Oldenburg Gano Daru Fameuse York Cilligos 42 Missouri Agr. Exp. Sta. Research Bulletin 53 Plate II. — By rows: left to right, Oldenburg, Primate, Wealthy; top to bottom, November 2 , 1921 , January 28 , 1922 , February 20 , 1922 . Relation of Temperature to Blossoming — Apple and Peach 43 Plate III. — By rows: left to right, Rome, Daru, Cilligos; top to bottom, November 2 , 1921 , January 28 , 1922 , February 20 , 1922 . 44 Missouri Agr. Exp. Sta. Research Bulletin 53 Plate IV. — Buds developing out of doors, 1920. Left to right: Cilligos, Fameuse, Oldenburg. Plate V. — Buds forced in greenhouse, photographed March 6. 1922. Cilligos (1), Rome (2) (3), Daru (4), Oldenburg (5), Primate (6), Fameuse (7). Relation of Temperature to Blossoming — Apple and Peach 45 APPENDIX Methods. — In the calculations reported in this paper, the date of first blossoming has been used. Though this standard is open to some objections, as, for example, a probable fluctuation with the total quantity of blossoms in the tree, it is less subject to change through varying judgments of different observers than is the date of full bloom. Several commentators have mentioned the variability in blossoming found in young trees. This may be explained by the fact that often in young trees all the blossoms are on terminal shoots, which open markedly later than blos- soms on spurs and if recorded without qualifications may well cause a con- siderable change in the relative order of blossoming. Phenological records in the apple should distinguish clearly between blossoms on spurs and those on shoots. All samples used in microscopic study were gathered from spurs which had blossomed at least once. Since progress in phenological studies depends on the availability of data, temperature and phenological records for Columbia, Missouri, are appended. DATES OF FIRST BLOSSOMING IN APPLE AT COLUMBIA, MO. (Days of the year) Variety 1905 1906 1907 1908 1909 1911 1912 1913 1914 1915 1916 1917 1918 Alexander 114 114 94 107 115 109 116 114 113 113 109 __ Arkansas 98 111 86 101 108 104 113 108 111 111 106 112 101 Arkansas Beauty 99 110 86 100 108 101 116 109 111 111 108 108 102 Arkansas Black 101 113 88 102 116 107 115 109 111 112 107 111 104 Ashton _ _ 97 111 86 102 108 112 118 112 112 112 109 Autumn Streaked __ 99 110 99 109 106 107 112 111 106 109 97 Bailey Sweet 97 110 84 101 107 101 115 107 111 109 105 108 104 Balagh _ _ 100 111 86 100 — 103 112 112 109 113 104 Baldwin 100 111 , 103 114 107 116 110 112 112 109 96 Battyani 113 93 98 103 116 110 112 108 107 Batullen _ 98 92 98 113 102 116 109 112 112 108 114 101 Ben Davis 99 111 85 101 113 102 115 109 111 111 108 110 102 Ben Hur 87 104 113 107 116 110 112 110 107 110 96 Black Ben Davis __ 98 111 103 101 119 109 112 106 Blenheim _ _ _ 108 114 95 104 116 106 118 112 113 113 114 108 Bosnian 105 112 89 103 109 113 110 105 P.ripr 95 110 87 99 109 112 112 106 113 96 Canada Reinette 99 111 87 101 109 108 115 112 Champion 100 110 87 103 105 118 110 112 111 109 104 Cilligos — 117 120 91 113 118 115 121 117 116 117 — Clark _ _ _ _ — _ 99 111 101 113 106 116 118 113 109 Clayton _ 99 111 88 102 112 104 116 110 112 108 Collins 102 110 86 101 109 102 114 109 111 110 107 111 107 Czar Thorn 99 111 85 99 108 104 116 109 111 109 109 Daru 114 116 105 112 124 112 121 113 107 114 121 Delaware Red 116 88 104 _ 115 112 115 111 105 Delicious 113 87 104 113 104 116 113 112 111 107 113 102 Devonshire Duke — 112 87 102 106 118 110 116 111 113 — Doctor 101 113 90 101 115 105 115 112 112 111 109 111 106 Downing Blush 113 87 102 108 118 109 110 Eper - 115 116 99 103 112 113 114 113 114 113 108 Fameuse 96 110 86 98 108 104 115 107 109 111 107 108 91 46 Missouri Agr. Exp. Sta. Research Bulletin 53 DATES OF FIRST BLOSSOMING IN APPLE AT COLUMBIA, MO. (Days of the year) Variety 1905 1906 1907 1908 1909 1911 1912 1913 1914 1915 1916 1917 1918 Faust’s Rome Beauty 113 115 99 107 119 105 — 114 113 113 109 __ __ Gano 98 111 86 102 113 103 116 110 111 112 109 114 101 Ginnie 103 111 88 104 114 106 118 110 112 __ 108 111 96 Gold Medal 100 114 87 102 113 105 119 111 115 112 109 __ 102 Golden Russet 95 120 85 10'S __ 108 116 — 109 111 106 109 92 Greening 102 111 87 102 116 106 117 111 — — __ __ __ Grimes 98 112 87 101 108 106 115 109 110 112 107 111 97 Heidorn __ 115 __ 107 116 106 115 111 112 112 112 113 105 Hubbardston 99 111 86 103 114 100 115 112 114 111 108 113 97 Huntsman 101 114 86 100 114 107 115 109 110 113 107 112 101 Imperial Janet 117 117 106 110 __ 119 121 119 118 __ 115 __ — Ingram 115 120 104 109 118 118 121 119 119 114 118 128 111 Jeffries __ __ — — __ 106 116 110 112 111 107 113 96 Jonathan 97 110 86 102 112 102 116 107 111 110 108 109 97 July 100 112 87 100 „ 111 116 111 114 112 109 110 106 Kansas Greening __ 110 115 108 107 __ 111 118 111 112 112 __ __ 108 Kartacs __ 116 108 106 118 __ __ 115 115 113 110 114 104 King David __ __ __ — 117 104 115 110 112 111 109 112 103 Lady 116 115 99 108 __ „ — — — 111 114 121 __ Lady Carter 99 110 87 101 113 __ — — 110 __ 108 113 102 London __ 115 88 104 112 __ __ __ 117 112 109 — — Late Duchess 97 109 87 98 107 __ __ __ 109 112 106 121 92 Longfield 100 111 95 100 109 — — — 115 __ 112 __ 106 Lou 98 109 87 100 105 __ __ __ __ 108 __ 108 — Louise 100 111 86 101 111 __ __ __ 112 112 109 112 96 Magyar 105 111 88 103 — __ — 110 112 112 109 114 106 Maiden Blush 99 110 87 98 108 107 115 108 110 110 107 109 95 Marin 97 110 86 100 109 104 115 107 109 111 105 109 97 Melon 97 110 88 99 108 105 116 108 — — — — — Menagera 97 109 86 100 108 101 114 109 108 110 108 109 91 Metitt 100 112 84 102 110 108 115 111 112 111 __ 109 96 Miller Boy’s Favorite 104 112 87 102 115 109 115 110 110 — __ „ — Minkler 98 110 85 99 107 102 119 107 108 109 107 108 96 Minnesota - 91 107 85 96 103 100 114 107 108 110 105 108 90 Missing Link __ __ 87 104 110 104 116 108 109 — 107 __ — Missouri 98 110 85 102 112 103 116 109 111 109 106 112 97 Mosher 110 112 87 104 110 106 118 110 112 112 109 110 — Me Intosh 100 111 87 102 109 105 117 110 112 111 109 113 __ Nelson Sweet 100 112 87 103 114 106 116 110 112 113 __ — __ Noble Savar 99 111 86 100 109 __ 118 109 112 112 109 114 — Nyack 101 114 88 103 115 110 102 111 112 __ __ — — Nyari Piros 105 112 86 99 109 104 115 110 109 114 107 __ 100 Ohio Beauty 110 113 90 102 __ 105 116 111 113 111 111 112 103 Ohio Pippin 99 110 85 100 llf2 103 116 108 113 111 108 109 97 Oldenburg 94 110 86 98 107 102 114 107 110 __ 106 — 97 Olive _ 99 112 87 102 111 106 — 111 112 113 109 114 __ Ontario 102 113 86 103 117 108 118 112 113 112 __ __ — Opalescent 106 116 110 117 114 113 111 — — 107 Payne Keeper __ „ 93 105 114 106 117 113 112 113 109 114 98 Peach — 118 — 104 __ 107 112 112 113 112 110 113 — Picket 100 110 86 99 110 105 __ __ 110 112 109 109 96 Ponyik 104 115 95 105 __ __ 118 __ 114 113 110 „ 106 Primate 97 110 87 99 107 __ 114 109 107 110 106 112 90 Relation of Temperature to Blossoming — Apple and Peach 47 DATES OF FIRST BLOSSOMING IN APPLE AT COLUMBIA, MO. (Days of the year) Variety 1905 1906 1907 1908 1909 1911 1912 1913 1914 1915 1916 1917 1918 Pumpkin Russet — 99 111 88 102 117 104 116 112 109 112 __ __ 100 Pumpkin Sweet 100 111 87 101 108 __ __ „ 110 111 108 110 98 Ralls 113 117 106 110 121 118 121 118 116 114 111 128 117 Reagan 100 112 88 102 111 106 116 — — __ — — — Red Astrachan 98 110 88 99 107 102 115 108 109 110 109 __ __ Red June __ _ __ __ — 104 111 __ 112 111 109 113 94 Red Stettiner 99 110 85 100 112 103 116 108 113 111 108 109 97 Rome 105 114 99 107 119 111 115 112 115 114 111 115 108 Rutherford 96 109 85 99 — — 115 109 112 111 108 110 92 Sabadka 100 115 87 __ — 106 118 112 113 113 109 110 103 Segfu — 112 87 104 113 110 117 112 112 112 110 110 102 Sekula 104 113 94 102 __ 110 __ __ 115 __ 109 — 102 Selumes 100 112 85 102 107 107 115 114 111 112 108 — 98 Skelton 99 111 87 102 112 __ __ 110 114 112 108 __ 104 Spitzenberg 99 111 87 102 109 __ __ 111 117 „ __ __ __ Standard 93 110 86 99 104 100 __ 108 110 112 106 — 92 Stayman 98 110 86 101 112 106 115 110 111 111 109 112 103 Summer Calville __ 97 110 86 98 108 105 116 109 109 — 108 110 92 Summer King — __ __ — __ 114 113 — 113 109 112 103 Tetofski 98 110 87 103 108 106 115 109 112 110 106 110 100 Titus Pippin 102 110 86 102 114 106 115 109 111 110 108 111 105 Tudor 99 115 87 — __ 110 116 109 109 __ 106 __ 98 Wafer 94 112 88 102 113 114 __ 110 114 112 110 112 106 Wealthy __ __ __ 104 112 104 117 109 112 111 109 110 103 White Canada 98 112 87 101 109 105 117 108 111 112 108 112 97 White Pippin 97 112 87 102 __ 107 116 — 112 __ 106 — 94 Wine Rubets 104 112 87 104 117 109 117 110 109 112 109 — 102 Winesap 99 111 87 102 113 104 116 110 112 111 109 109 104 Wolf River 108 113 88 102 112 109 116 110 111 111 110 113 102 Woodmansee ~ — — — — 118 113 114 112 110 114 121 Workaroe 109 113 — 103 115 109 118 112 115 111 __ __ — Yappa 102 112 87 102 116 106 118 108 — __ __ __ — Yellow Newtown __ __ __ 104 __ 108 117 113 113 112 109 __ 103 Yellow Transparent — — — 108 113 104 117 113 113 112 108 110 105 York Imperial 99 113 88 103 113 106 116 111 — 112 __ 112 __ York Stripe 103 114 88 104 116 107 117 113 — __ — __ __ 48 Missouri Agr. Exp. Sta. Research Bulletin 53 DAILY MAXIMUM TEMPERATURES, COLUMBIA, MISSOURI January 1905 1906 1907 1908 1909 1910 1911 1912 1913 1914 1915 1916 1917 1918 1 66 39 40 47 28 46 52 26 56 42 41 62 43 44 2 43 41 49 50 49 29 7 26 38 42 33 42 50 43 3 27 41 47 49 59 23 5 19 42 29 38 48 49 31 4 35 29 46 47 60 29 34 11 36 29 41 61 50 44 5 40 42 65 42 33 25 50 13 23 28 47 63 42 42 6 25 41 72 51 4 12 48 —3 20 39 42 21 56 28 7 24 43 71 52 17 35 55 2 20 54 43 29 40 21 8 32 19 54 44 33 36 47 12 24 58 30 38 54 26 9 31 40 26 52 43 29 47 29 32 40 49 49 57 20 10 10 45 33 47 42 43 68 10 38 28 40 51 52 16 11 32 35 44 36 8 42 67 3 38 44 40 37 22 9 12 15 35 43 30 18 44 31 —5 20 34 41 34 37 0 13 11 43 56 31 32 44 34 20 32 46 52 0 16 15 14 9 48 40 39 35 30 34 22 43 52 54 20 30 18 15 17 58 28 44 34 33 22 7 56 58 61 34 23 16 16 30 37 33 26 30 37 27 29 60 53 58 11 22 23 17 39 57 34 40 27 51 28 43 59 43 28 18 30 18 18 40 40 54 43 36 44 29 33 38 50 31 23 30 13 19 37 63 60 49 37 57 41 20 64 66 34 32 40 17 20 46 72 26 58 46 44 54 27 35 47 25 55 40 16 21 34 65 49 58 61 32 39 48 32 32 20 57 60 20 22 24 17 39 44 74 42 33 49 37 54 19 52 17 24 23 33 24 35 35 72 50 45 51 42 60 14 54 36 37 24 31 43 51 32 65 44 49 34 41 41 16 62 34 42 25 10 47 23 47 40 62 55 39 54 38 26 60 45 42 26 30 48 18 42 43 61 71 50 53 62 31 63 41 28 27 37 48 25 42 57 41 60 35 43 64 26 56 42 15 28 29 47 32 42 61 42 47 34 43 65 15 33 66 18 30 25 48 33 31 10 29 40 28 54 30 42 31 51 19 31 27 46 35 34 18 32 64 41 31 43 51 27 47 5 Relation of Temperature to Blossoming — Apple and Peach 49 DAILY MAXIMUM TEMPERATURES, COLUMBIA, MISSOURI February 1905 1906 1907 1908 1909 1910 1911 1912 1913 1914 1915 1916 1917 1918 1 15 44 37 17 43 57 81 33 14 52 48 16 6 11 2 —2 27 36 31 55 51 38 21 26 55 32 17 3 30 3 5 55 14 35 63 38 54 20 32 38 38 21 36 24 4 20 42 11 37 65 43 41 18 30 32 56 36 38 11 5 20 14 12 52 60 44 46 17 15 35 37 36 24 53 6 21 21 21 36 45 31 43 23 27 32 31 22 48 50 7 23 30 25 49 43 40 41 24 25 15 34 14 47 59 8 28 40 47 41 54 41 46 22 34 16 38 35 38 65 9 29 27 50 43 53 32 40 20 35 45 45 41 24 40 10 13 29 41 46 33 43 47 34 40 37 65 44 29 63 11 27 43 49 46 56 42 48 39 36 28 64 44 21 65 12 9 58 57 62 54 25 54 30 21 21 69 35 33 54 13 —1 51 60 54 35 42 64 37 38 22 61 21 44 59 14 24 28 43 40 36 59 54 36 52 24 49 33 35 62 is 15 28 60 38 21 63 78 41 51 38 34 43 38 30 16 30 37 60 34 25 17 76 53 61 27 51 56 56 32 17 30 36 62 39 49 17 65 53 63 54 57 52 52 36 18 35 55 66 37 46 30 39 55 69 42 55 40 33 51 19 35 61 44 23 38 39 32 45 63 27 51 55 52 60 20 34 59 43 37 54 41 26 35 38 30 54 55 33 16 21 44 57 25 34 58 26 22 30 51 48 52 59 53 19 22 47 69 26 56 54 26 31 40 34 38 50 63 64 45 23 55 54 31 49 55 10 35 52 23 17 47 37 56 69 24 51 50 35 49 35 28 43 43 26 21 34 49 37 65 25 50 45 50 48 55 43 48 39 40 32 41 43 68 63 26 60 39 54 36 56 42 39 35 40 46 35 37 62 51 27 44 32 58 31 52 33 34 38 29 53 35 32 31 42 28 63 50 61 52 65 48 29 33 22 50 40 31 41 38 29 .... 71 .... .... 27 .... .... .... 38 .... 50 Missouri Agr. Exp. Sta. Research Bulletin 53 DAILY MAXIMUM TEMPERATURES, COLUMBIA, MISSOURI March 1905 1906 1907 1908 1909 1910 1911 1912 1913 1914 1915 1916 1917 1918 1 59 62 54 48 67 57 34 31 28 27 44 32 36 50 2 67 59 59 48 70 70 46 24 31 39 42 32 34 58 3 76 34 54 45 49 76 59 29 49 35 42 26 31 51 4 58 33 40 49 48 76 46 33 41 43 31 50 22 62 5 67 34 51 74 62 81 70 27 38 50 33 53 38 76 6 39 40 41 71 54 64 59 37 30 40 29 66 57 42 7 44 42 49 46 62 58 49 36 52 34 34 43 50 53 8 50 60 45 40 46 57 63 33 67 37 39 38 50 65 9 53 52 38 47 42 42 71 28 57 47 41 56 64 73 10 43 39 39 56 43 48 61 31 55 48 39 52 69 47 11 43 32 53 50 39 60 82 35 59 34 37 49 54 65 12 50 22 55 73 46 61 60 36 61 47 41 76 44 77 13 47 29 37 55 51 69 49 40 57 62 45 76 54 90 14 69 25 43 74 37 47 60 37 54 69 49 51 45 61 15 70 25 63 66 45 52 50 33 28 79 39 36 54 50 16 71 23 66 66 48 65 45 51 39 55 46 52 60 56 17 66 26 68 63 52 67 64 58 63 53 44 56 41 73 18 71 29 67 63 72 71 52 65 68 36 42 62 41 76 19 55 30 80 45 48 74 60 67 68 30 42 54 65 72 20 36 34 70 51 48 70 72 41 50 37 33 71 49 74 21 48 46 92 61 50 73 77 28 34 37 37 86 64 80 22 73 32 90 60 60 90 59 32 45 46 37 64 71 62 23 61 29 82 67 69 88 53 34 72 58 47 54 64 42 24 71 35 77 67 66 85 58 39 53 68 63 82 66 59 25 71 55 90 81 45 82 71 48 39 70 50 69 75 69 26 71 59 82 73 67 86 61 49 33 73 39 56 59 75 27 82 42 82 78 49 86 50 42 32 61 48 51 50 64 28 69 39 78 44 55 86 66 43 53 77 53 63 69 61 29 60 37 58 57 45 78 55 59 62 67 46 62 62 66 30 73 53 61 48 52 65 46 68 75 58 44 68 82 72 31 77 49 47 67 49 60 45 72 63 69 40 59 80 75 Relation of Temperature to Blossoming — Apple and Peach 51 DAILY MAXIMUM TEMPERATURES, COLUMBIA, MISSOURI April 1905 1906 1907 1908 1909 1910 1911 1912 1913 1914 1915 1916 1917 1918 1 81 55 54 57 57 71 48 50 72 57 40 52 50 79 2 72 70 63 38 56 69 69 57 79 63 46 50 54 83 3 70 74 63 60 61 74 43 65 72 46 54 61 62 51 4 58 61 70 65 85 76 47 74 50 52 71 59 54 55 5 55 56 50 65 84 56 55 80 65 60 75 56 54 54 6 55 64 50 80 76 60 63 72 69 56 76 45 59 58 7 61 73 57 69 59 73 49 55 53 46 75 36 51 60 8 86 69 47 70 48 77 57 66 46 36 77 44 46 53 9 90 72 52 54 51 75 58 67 65 46 70 50 57 48 10 74 66 51 63 59 81 62 68 48 53 69 68 70 54 11 59 78 54 59 73 63 54 77 44 49 63 82 70 53 12 62 82 47 69 61 58 78 76 43 57 58 84 57 52 13 64 66 45 79 55 72 66 74 57 62 60 70 51 64 14 53 45 52 68 64 71 53 74 68 65 71 59 59 66 15 48 54 62 68 65 68 62 65 72 72 77 68 52 61 16 46 63 45 57 80 47 70 49 77 83 81 62 73 73 17 55 69 51 61 87 42 77 44 84 83 79 65 83 65 18 61 74 44 64 78 40 61 55 79 70 76 79 83 67 19 66 77 47 81 48 42 62 53 66 51 84 76 80 55 20 75 75 56 83 51 66 61 70 63 64 78 70 62 38 21 61 81 60 80 55 78 61 73 81 84 76 55 72 50 22 64 64 65 82 54 65 58 58 82 81 78 67 81 63 23 70 61 70 79 60 40 59 68 80 67 86 65 86 63 24 73 87 78 73 69 36 60 75 60 81 84 61 81 48 25 73 80 72 75 71 42 63 65 60 81 84 57 61 45 26 59 82 57 55 82 59 75 72 64 86 78 51 53 58 27 80 86 65 48 64 77 58 67 57 81 83 61 49 58 28 85 70 81 54 80 85 76 66 66 71 85 69 54 62 29 72 77 72 52 88 91 82 57 73 58 73 73 49 55 30 73 61 47 57 51 80 72 67 84 53 73 64 54 56 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 54 Studies In Animal Nutrition II. Changes in Proportions of Carcass and Offal on Different Planes of Nutrition (Publication authorized September 1, 1922.) COLUMBIA, MISSOURI SEPTEMBER, 1922 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL the; curators of the university OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY P. E. BURTON H. J. BLANTON St. Louis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION J. C. JONES, PH. D., LL. D., PRESIDENT OF THE UNIVERSITY F. B. MUMFORD, M. S., DIRECTOR STATION STAFF SEPTEMBER. 1922 AGRICULTURAL CHEMISTRY C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, Ph. D. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. SiEveking, B. S. in Agr. AGRICULTURAL ENGINEERING J. C. Wooley, B. S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale, B. S. in Agr. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. W. P. Hays ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. C. McBride, B. S. in Agr. FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, Ph. D. O. W. Letson, B. S. in Agr. Miss Regina Schulte* RURAL LIFE O. R. Johnson, A. M. S. D. Gromer, A. M. E. L. Morgan, A.M. Ben H. Frame, B. S. in Agr. Owen Howells, B. S. in Agr. HORTICULTURE T. J. Talbert, A. M. H. D. Hooker, Jr., Ph. D. J. T. Rosa, Jr., Ph. D. H. G. Swartwout, B. S. in Agr. J. T. Quinn, B. S. in Agr. POULTRY HUSBANDRY H. L. Kempster, B. S. Earl W. Henderson, B.S. SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M W. A. Albrecht, Ph. D. F. L. Duley, A.M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr. Richard Bradfield, Ph. D. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Secretary S. B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsham, Librarian. E. E. Brown, Business Manager. In service of U. S. Department of Agriculture. STUDIES IN ANIMAL NUTRITION II. Changes in Proportions of Carcass and Offal on Different Planes of Nutrition. C. Robert Moulton, P. F. Trowbridge,* L. D. Haigh The changes experienced by beef cattle in form and weight when on different planes of nutrition were presented and discussed in a previous bulletinf. Representative animals from each of the groups were killed at intervals from birth to four years old. The data collected in the slaughter house will be presented in this bul- letin. RATION For a general discussion of the treatment of the animals the previous bulletin must be consulted. The ration included milk for several months after birth and timothy hay and grain were soon introduced. At weaning time the ration consisted of alfalfa hay and a grain mixture in the ratio of one to two. The grain consisted of six parts corn chop, three parts whole oats, and one part of old process linseed meal. PLANE OF NUTRITION The animals were early divided into three groups. Group I was fed all it would eat of the ration. Group II was fed for maxi- mum growth without permitting the laying on of much fat. Group III was fed for scanty or retarded growth. The Group II steers gained about a pound a day for the first two years while the Group III cattle gained but 0.69 pounds per day. SLAUGHTERING In the conduct of this experiment great care was exercised that all of the operations with the different animals should be carried out under similar conditions. Although the slaughtering and sub- sequent operations were necessarily carried out at different times, •Resigned September, 1918. tC. Robert Moulton, P. F. Trowbridge, L. D. Haigh, Studies in Animal Nutrition, I. Changes in Form and Weight on Different Planes of Nutrition, Mo. Agr. Expt. Station, Research Bulletin 43. 4 Missouri Agr. Exp. Sta. Research Bulletin 54 in order to make the results strictly comparable the slaughtering and cutting were done by the same experts each time*. Each animal was slaughtered on the day following the closing of its feeding period. In the morning the steer was fed and weighed as usual but no water was given. If slaughter occurred late in the morning or in the afternoon the animal was weighed again im- mediately before slaughtering. The animal was led from the feed- ing shed to the slaughter house followed by men with a shovel and long handled dipper to catch any feces or urine that might be voided. The animal was stunned with a knocking hammer, shackled by the hind legs and hoisted to swing clear of the floor. An oil cloth, funnel shaped bag had been fastened to the muzzel before hoisting so that any vomit voided could be caught and weighed. The suspended animal was stuck in the throat near the brisket so that both the jugular vein and carotid artery were severed. Com- plete bleeding was assured by pumping the fore legs up and down. The blood was caught in a tared pan and weighed. The volume of this main weighed portion was determined. A tared pan was kept under the animal to catch any blood that might drip while skinning out the head. While the blood was flowing freely the samples for analysis were taken and poured in suitable quantity into tared, covered containers and crucibles. The blood was still warm and no clotting had yet occurred. While the carcass was hanging the head was skinned out. The gullet was firmly tied with a skewer thrust through below the tie and the head was then severed permitting the dripping blood to collect in a tared container. The head was immediately placed in a large can provided with a cover and the weight obtained. The tongue with larynx and bones was removed, separated into tongue marketable, tongue base, bones, larynx, and piece of gullet. The parts were weighed and set aside in closed containers. Horns were sawed off and weighed. The skull and entire head was split accur- ately in half and the brain was removed and weighed. The lean meat and fat were removed from the right half as were the teeth. If any vomit was found it was weighed and discarded. Both halves of the head were weighed. The total lean and fat were obtained * Swift & Company of Kansas City furnished an expert butcher to do the slaughtering. Mr. Samuel Godfrey, foreman of the beef cutting department, did the cutting. Studies In Animal Nutrition — II 5 by doubling the right-side weights and the total bone by subtract- ing the sum of the lean and fat from the sum of the two halves of the head minus the teeth. As soon as the suspended carcass with the head removed had practically stopped bleeding it was lowered to the floor with the anterior end lying up the slope of the smooth cement floor. A man with a rubber window wiper and sharp edged dust pan kept all oozing blood wiped up and transferred to weighed containers. In skinning out the feet the dew claws were removed, weighed and saved and care was taken to remove the hide exactly at the hoof line. From the right feet the hoofs were separated and the remainder of the material was considered as skeleton. The usual packing house order of procedure was followed in skinning the carcass and removing the internal organs. The caul fat was re- moved while the carcass was on its back. The bladder was tied before removing and weighed with its contents and again empty. The rectum was tied as soon as it was cut loose. The tail was removed and split in two and the lean and fat meat separated and weighed. The contents of the abdominal and thoracic cavities was caught in a large tub, or tubs, and weighed. The separation and weighing of the organs was pushed as rapidly as possible to reduce to a minimum the loss of water by evaporation. A double tie was made at the end of the small intestine near the abomasum before severing one from the other. The fat was carefully cut or scraped from all four stomachs and weighed. The intestines were also carefully freed from fat, and this was generally accomplished without any portion becoming smeared with the contents. The stomachs were emptied of the contents and were cleaned by washing with water after which they were wiped dry with cloths. The contents of the intestines were removed by stripping through the fingers taking a section at a time. The stripped sections were split open and the inside was wiped lightly to remove all contents. Occasionally it was necessary to wash and wipe a smeared portion. The various organs were separated, weighed and put into closed containers. The hide was removed and the carcass was split into halves. Then the spinal cord was removed. The diaphragm was removed back to the striated muscle and composited with the internal or- gans. The carcass was allowed to chill for 48 hours and the right 6 Missouri Agr. Exp. Sta. Research Bulletin 54 side was separated into the two quarters and then into the standard wholesale cuts. (Figure 1). Each cut was separated into lean meat, fatty tissue and bone. The tendons were weighed with the bone. In the separation of the lean and fat, care was taken that the fatty tissue should contain no lean meat. Necessarily the lean meat contained small pieces of fat which could not be separated. The meat was cut from the bones as completely as could be done with a boning knife. All samples were kept in closed tin containers. The further treatment of the separated parts is of interest only in connection with the chemical analysis and the description will be deferred to a later publication. THE SLAUGHTER HOUSE DATA The detailed slaughter house data obtained are given in the Appendix in Tables 1 to 5 for the offal parts and Tables 6 to 10 for the carcass parts. Few of the weights need explanation. The warm empty weight was obtained by subtracting the contents of the stomachs, intestines, and urinary bladder and any excrement voided before death from the live weight at slaughtering. The heart and neck sweetbreads are the thymus gland. The stomach and intestinal fats are those which adhered to the respective organs. The intestinal fat is largely included in the mesentery. The caul fat is that laid on in the part of the peritoneum stretching like an apron over the stomachs and intestines. The different divisions are mutually exclusive excepting where specified otherwise. Table I. — Percent Empty Weight to Live Weight. Age Group Group Group I II III At birth . 98.41 98.41 98.41 3 months . 88.02 89.27 83.89 5% months . 84.32 85.29 86.63 8^ months . 83.16 82.28 83.19 11 months . 88.30 87.61 87.10 18 months . 88.77 90.37 21-26 months . 90.44 88.69 87.05 34 months . 90.39 91.63 38 months stunted . . 92.30 40 months . 93.26 89.19 88.95 45 months . 90.40 87.68 89.01 47-48 months . 92.24 90.12 89.09 Empty Weight. — Table I gives the proportion of empty weight in the live animal from birth to four years for each group of cattle. The figures in this table as well as in Tables II to XVIII inclusive Studies In Animal Nutrition — II 7 are taken from Tables 11 to 15 in the Appendix. The figures pre- sented for the animal at birth are for the average of Hereford calves reported in Research Bulletin 38 of this Station. The comple- ment of the percent of empty weight is of course the percent of fill. The animal at birth has the largest percent of empty weight. This is due to the lack of food and food residues in the alimentary canal. The figure decreases during the early months showing a large proportion of fill and is least at 8^2 months. It then gradually increases with some irregularity and becomes at about 3 years and thereafter a higher proportion than at any time since birth. The amount of the ration afifects the percent of the empty weight. The lighter rations show generally a smaller percent of empty weight and consequently indicate more relative fill. Since the weight of the ration is smaller and the weight of fill is smaller this can only be due to a greater difference in weight of animal re- sulting in a decreased percent of empty weight. At 8 3/2 months there is practically no difference between the three groups. Carcass. — Tables II and III show the percent of carcass in the live and empty animals respectively. The percent of carcass to live weight decreases to about 8*4 months and then increases to reach at 3 to 4 years a higher value than at any previous time. For the first six months there is little difference between the groups but thereafter the better fed animals have the greater percent of carcass. The effect of varying proportions of fill is shown in the figures discussed in the above paragraph. On the empty weight basis there is a continuous increase in percent of carcass from birth to Table II. — Percent Carcass to Live Weight. Table III. — Percent Carcass to Empty Weight. Age At birth 3 months 5 % months 8% months 11 months 18 months 21-26 months 34 months 38 months stunted . . 40 months 45 months 47-48 months Group Group Group I II III 59.30 59.30 59.30 54.19 57.32 53.63 53.71 53.00 54.39 54.17 50.46 51.09 57.52 53.85 55.23 60.46 56.02 58.24 57.71 54.27 61.49 60.49 65.69 70.42 61.18 58.08 65.32 60.28 60.34 68.95 61.80 59.63 Age At birth 3 months 5% months 8% months 11 months 18 months 21-26 months 34 months 38 months stunted . . 40 months 45 months 47-48 months Group Group Group I II III 60.39 60.39 60.39 61.56 64.21 63.92 63.70 62.14 62.78 65.14 61.33 61.41 65.14 61.47 63.41 68.11 61.98 64.39 65.07 62.34 68.03 66.02 71.17 75.52 68.59 65.29 72.25 68.76 67.80 74.75 68.58 66.63 8 Missouri Agr. Exp. Sta. Research Bulletin 54 3 or 4 years of age. The relation between the groups is the same as when the live weight formed the basis. Carcass and Offal Fat. — In Tables IV and V are shown the percents of carcass plus offal fat to live and empty weights. The offal fat is the hand separable fat on the internal organs. It is small in amount in the young animals, being entirely negligible in the calf at birth. In the very fat four year old steer it amounts to nearly five percent of the animal. The effect of adding this fat to the carcass weight is merely to increase the spread of the figures with increasing age and fatness. The general relations pointed out in the section above hold here. Table IV. — Percent Carcass and Oefal Table V. — Percent Carcass and Oeeal Fat to Live Weight. Fat to Empty Weight. Age Group Group Group Age Group Group Group I II III I II III At birth . 59.30 59.30 59.30 At birth . 60.39 60.39 60.39 3 months . 55.45 58.06 54.17 3 months . 62.99 65.03 64.57 5% months . 57.01 54.53 54.23 5% months . 67.61 63.94 63.76 8 y% months . 56.05 52.23 51.85 8% months . 67.40 63.48 62.33 11 months . 61.16 56.17 56.69 1 1 months . 69.26 64.12 65.08 18 months . 65.04 57.40 18 months . 73.27 63.49 21-26 months . 63.01 59.79 55.89 21-26 months . 69.67 67.41 64.21 34 months . 65.51 62.96 34 months . 72.47 68.71 38 months stunted . . 71.34 38 months stunted . . 77.30 40 months . 76.18 63.58 59.46 40 months . 81.69 71.29 66.84 45 months . 71.62 62.53 62.60 45 months . 79.22 71.32 70.33 47-48 months . 73.33 64.98 62.19 47-48 months . 79.49 72.11 69.80 Table VI. — Percent Offal Fat to Empty Weight. Age Group Group Group I II III At birth none none 3 months 1.43 0.83 0.65 5% months .. 3.91 1.80 0.98 8% months 2.26 2.16 0.92 1 1 months 4.12 2.65 1.68 18 months 5.16 1.51 21-26 months 5.28 2.34 1.87 34 months 4.45 2.70 38 months stunted . 6.13 40 months 6.17 2.70 1.55 46 months . 6.97 2.56 2.54 47-48 months 4.74 3.53 3.17 Table VI gives the percent of offal fat to the empty animal. There is a rather consistent increase in percent of offal fat with increasing age and fatness. A striking exception is shown by the four year old Group I steer. This is due to a great decrease in the Fig. 1. — Wholesale divisions of the beef carcass. Studies In Animal Nutrition — II 9 weight of offal fat, this animal having but 38.5 kilograms while the next younger animals had 53.6 and 48.3 kilograms respectively. The low figure should be taken with reservations as to its general applicability. Hide and Hair. — The percent of hide and hair referred to empty weight is given in Table VII. There are some variations due to individuality but in general the percent decreases from birth to four years. The percent also decreases with increasing plane of nutrition. Put in different language the percent of hide and hair decreases with increasing age and fatness of the animal. Blood. — In Table VIII are presented the figures for the per- cent of blood referred to empty weight. The maximum is at three months. At birth it is less and as age increases beyond three months it becomes less. In the poorer fed groups the percent of blood becomes fairly constant, however, while in the full fed group it continues to decrease with age. In the early stages the full fed animals have as much or more than the others but as the age in- creases the Group I steers have a materially smaller percent of blood. Table VII. — Percent Hide and Hair to Empty Weight. Table VIII. — Percent Blood to Empty Weight. Age At birth 3 months 5% months 8 Y 2 months 11 months 18 months 21-26 months 34 months 38 months stunted . . 40 months 45 months 47-48 months Group Group Group I II III 12.13 12.13 12.13 10.51 9.48 9.26 8.16 10.60 9.27 8.53 8.62 9.04 8.81 9.72 9.44 7.40 8.69 8.65 9.80 10.47 7.43 8.23 7.15 5.88 8.35 9.34 5.87 8.90 9.61 6.15 8.36 8.81 Age At birth 3 months 5^2 months SV 2 months 11 months 18 months 21-26 months 34 months 38 months stunted . . 40 months 45 months 47-48 months Group Group Group I II III 4.93 4.93 4.93 6.24 5.35 6.37 5.18 5.25 5.35 5.08 5.85 5.19 4.33 4.54 5.06 4.09 4.93 4.41 4.53 5.13 4.16 4.85 3.61 3.48 4.43 5.28 3.33 4.44 4.67 3.52 4.90 5.22 For a somewhat different presentation of these and other fig- ures bearing on the relation of blood, surface, and nitrogen to the animal the reader is referred to two earlier publications from this Station 1 Heart. — The percent of the heart referred to the empty weight J P* F. Trowbridge, C. R. Moulton, L. D. Haigh, The Maintenance Requirement of Cattle, Research Bulletin 18, Missouri Agr. Expt. Station (1915). C. R. Moulton, Units of Reference for Basal Metabolism and Their Interrelations, Four. Biol. Chem. XXIV. 299-320 (1916). 10 Missouri Agr. Exp. Sta. Research Bulletin 54 is given in Table IX. The figures are not notable but are very small and decrease with increasing age and fatness. There is, however, very little difference between Groups II and III. Lungs. — The proportion of lungs and trachea to the empty weight (Table X) increases during the first few months of life and then falls rather steadily with advancing age. The well fed group has materially less lungs than the other groups especially at the older ages. Central Nervous System. — The percent of brain and spinal cord is given in Table XI. There is a rather steady and uniform decrease with increasing age and fatness. From slightly less than 0.9 percent at birth it decreases to 0.09 percent at maturity for the fattest animal. Stomachs. — In contrast to the foregoing organs and parts the stomachs (Table XII) show an increasing proportion from birth to 8 J / 2 months for Groups I and II and from birth to two years for Table IX. — Percent Heart TO Empty Table X. — Percent Lungs TO Empty Weight. Weight. Age Group Group Group Age Group Group Group I II III I II III At birth 0.55 0.55 0.55 At birth . . . 1.01 1.01 1.01 3 months 0.53 0.43 0.44 3 months . . 1.30 1.16 1.32 5% months 0.54 0.57 0.50 5% months . 1.14 1.10 1.26 8% months 0.45 0.46 0.56 8% months . 1.09 1.21 1.23 11 months 0.38 0.54 0.49 11 months . 0.87 1.12 1.09 18 months 0.45 0.53 18 months . 0.84 1.00 21-26 months 0.34 0.44 0.44 21-26 months 0.76 1.00 0.62 34 months 0.40 0.49 34 months . . 0.54 0.90 38 months stunted . 0.32 38 months stunted . 0.67 40 months 0.41 0.39 0.49 40 months . 0.55 0.89 1.07 45 months 0.30 0.44 0.42 45 months . 0.63 0.83 0.84 47-48 months 0.27 0.40 0.36 47-48 months 0.47 0.79 0.92 Table XI. — Percent Brain and Cord to Table XII.— Percent Stomachs to Empty Empty Weight. Weight. Age Group Group Group Age Group Group Group I II III I II III At birth 0.86 0.86 0.86 At birth . . . 0.99 0.99 0.99 3 months 0.41 0.51 0.50 3 months . . . 2.01 1.55 1.88 5 x /• t- r 30 E Id k 0 20 H r Id £ K hi 0. 0 LEAN 20 lo 4 — r-i SKELETON E 3 3*4 8% 11 18 21 35- 39* 44*47 2 AGE IN MONTHS. Fig. 3. — Distribution of tissues in empty animal — Group I. increases very markedly from an amount so small as to be insep- arable from the lean at birth to 40 percent at four years. The or- gans of the animal are about 11 percent from birth to 8 months. They then decrease to about 6 percent at four years. The bloo r l and the hide have been discussed above. 16 Missouri Agr. Exp. Sta. Research Bulletin 54 In Group II (Figure 4) less change with advancing age is shown. This is largely due to a smaller amount of fatty tissue be- ing formed. The skeleton decreases from about 28 percent at birth to 16 percent at four years. The lean flesh is about 39 percent at birth and increases to 46 percent at 45 months. The four year old 10 0 DI5 TR BU TIOM or TISSUES •" EMPTY AMIMAL HIPE. $ o rr'BLix* ti n n mi 10 o r-i ORGANS. 10 0 n — FATTY TISSUE. £' 240 I— 1 [“1 j— 1 5! r 5« & H,. tU LEAN FLESH. £ 2d r-j 10 — e SKELETON. i ca 3 9% 8* 11 26 34 40 45 48 AGE 1 N MONTHS. Fig. 4. — Distribution of tissues in empty animal — Group II. steer shows but 41 percent, probably an abnormal figure due to an increase in percent of fatty tissue. The fatty tissue increases from practically nothing to 19 percent at four years. The organs are about 11 percent at birth and 8.5 percent at four years. Studies In Animal Nutrition — II 17 Group III (Figure 5) shows still less change. The skeleton decreases from about 28 percent at birth to 18 percent at four years. The lean flesh increases from 39 percent at birth to 45 to 48 percent at the end. The fatty tissue is small in amount running from prac- tically nothing at birth to only a little over 11 percent at four years. The organs decrease from 11 percent at birth to 9 percent at four years. 1«J P DISTRIBUTION of TI 55 UCS »*E.MPTY ANIMALcroopei. 0 NIPE. 3 -__ IT inn 1 1 ~tt BL00P - 1 1 ni i 10 __ i 0 ORGANS. 10 1 5 FATTY ^ o nnn TISSUE pp X o taJ >40 r-j — _ 2 >- 30 K O. z Id 20 k Q 1- Z 14 LEAN ui £ Id 0> FLESH. CL to 1* — r— I Q SKELETON. - 3 5 V Z 11 18 26 40 «fc 43 40 t AG L IN MONTHS. J® - J Fig. 5. — Distribution of tissues in empty animal — Group III. Loss on Cooling and Cutting. — The weights and percents of the main divisions of the empty animal and the calculated loss on cooling and cutting are given in the appendix in Tables 16 to 20. The losses vary from practically nothing to over 6 percent in the case of one animal. Five animals show about 5 percent. The other 18 Missouri Agr. Exp. Sta. Research Bulletin 54 25 animals show less than 3 percent loss. This loss is largely moisture lost from the animal during slaughter, cooling, and cut- ting. While the differences between the groups are not large, in general the greater percent losses are from the thin animals. The covering of fat on the Group I steers protects the carcass and even the offal from as great a moisture loss. PISTRIBUTION or CUTS •" CARCASS -group*. S' , , Mn nn an shin. n nn nn ^ „ _ IS is r-i r-i - X CHUCK. IS 1A — o — plate:. !• — a m. — H * A OS < u — - — k. 10 o 1- £ « LOIN. us , a. * 1 — li 1 r— i anp kipney. n mn ril 1 ij * r— _nnnn nf flank r . rr 1 nnnri nfT~ ~ nm i nn" Vr— «« UJ t- ROUMP SC n n Pirn rrrr ~n r-i mm 3 5*4 8'4 11 18 21 35 36* 33^ 44*47. AGE IN MONTHS. Fig. 6. — Distribution of cuts in carsass — Group I. DISTRIBUTION OF WHOLESALE CUTS Tables 21 to 25 in the appendix give the detailed figures for the distribution of the wholesale cuts in the carcass. Figures 6 to 8 show these figures graphically. In Group I (Figure 6) the shank (hind shank) decreases from about 5 percent at three months to 2 percent at four years. The Studies In Animal Nutrition — II 19 round decreases from 20 percent to 13 percent. The rump in- creases from 3 to a little over 4 percent. The flank increases from 3 to 6 percent. The kidney fat increases from about 1 percent at three months to 4 percent in the baby beef (18 to 21 months) and then decreases again to 2 to 3 percent. The loin increases from 16.5 percent to 21 percent at four years. The rib cut does not vary DISTRIBUTION or CUTS »* CARCASS. «- CROUP®. Jnnnn sum. n n mm S'J’rn— i r—i i— i NtCK. r— i i — i i—i r— i i — i 2o r-i - 10 O CHUCK. £ 10 5 as 3 - 10 -° PUTL. O o RIB. £ 15 __ — Z y to E u * LOIN. 5 KEDHEY FAT 5 * °i — 1< — i i — i r—i AMP KIDNEY. | — | r i r— i i — i i — i Q-5 1 — 1 1 i n n flank. n LrU a.o*o — — RUMP |“l n is . 10 0 ROUND. 3 ornnn SHAf1K - n n n n n 3 3‘4 8V 2 11 26 34 40 45 48 AGE lb MONTHS. Fig. 7. — Distribution of cuts in carcass — Group II. much being 8 percent at three months, and 9 to 10 percent at four years. The plate increases from 10 percent to 17 percent. The chuck decreases from 25 percent to 20 percent. The neck decreases from about 2 percent to 1 percent and the shin (fore shank) de- creases from 6 percent to 3 percent. 20 Missouri Agr. Exp. Sta. Research Bulletin 54 The Group II cattle show much less marked changes in distri- bution of the cuts from three months to four years. The shank decreases from 5 to 3 percent, the round from 20 to 18 percent, the shin from 6 to 4 percent, and the neck from nearly 2 to 1 percent. The flank increases from 2 to 4 percent, the loin increases from 16 to 18 percent at Sy 2 months and then decreases again to less than 17 percent, and the plate increases from less than 9 percent to nearly 14 percent. The rump, kidney and kidney fat, rib, and chuck remain fairly constant. In Group III (Figure 8) there is less change with age. The shank, kidney and kidney fat, and shin decrease with age. The loin and plate show fair increases while the rump and flank show slight increases. The round, rib, chuck, and neck show individual variations but on the whole are constant. The fatter animal, then, increases its proportion of loin, the most expensive cut, of rump, a less expensive cut, and of flank and plate very cheap cuts. The rib cut, a rather expensive cut, in- creases but slightly. On the average for all three groups it is con- stant. The round, a valuable cut, decreases with increasing fatness as do the chuck and the neck. The shin and shank decrease in all cases with increasing age irrespective of fattening. Summing up the changes with respect to the effect of fattening on the three ex- pensive cuts — loin, rib, and round — it is seen that the first increases with fattening, the second remains fairly constant, and the third decreases. LEAN, FAT, AND BONE IN THE CARCASS Figure 9 (Tables 26 to 30 in the appendix) shows the propor- tions of skeleton, lean flesh and fatty tissue in the entire carcasses of the steers from three months to four years for all three groups. In the Group I steers the skeleton decreases from about 25 percent to 10 percent, and the lean flesh decreases from 67 percent to 42 percent. The fatty tissue of the carcass, on the other hand, increases from 6 percent to nearly 48 percent. In Group II the changes are not so marked since relatively less fattening occurs. The skeleton decreases from nearly 28 percent to a little over 18 percent and the lean flesh decreases from 66 per- cent at three months and 68 percent at 11 and 26 months to 58 per- cent at four years. The fatty tissue increases from 5 percent to 22 percent. Studies In Animal Nutrition — II 21 The changes are still less marked in Group III. The skeleton decreases from 27 percent to a little over 20 percent while the lean flesh increases from 67 percent at three months to about 70 percent 5_| D13T r~ip-u RIBUT10M or CUTS •« CARCASS ! - oroup-bi. s-o X n r i ii shin, ii n n o . — .i — i < — .i — i rn m ^E-CK. . — , 12 pi — p 16 o CHUCK. 16 0 — PLATE. 1* 1 — — — RIB. 2 U ct * i pi p < u M LOIN. „ KlPHETFAr.-KiPN^ _ £ £-6 1 — ii — i r-i rn m n FLAMK. r— i run H . nn nn □ CL Rump n nn £ faj — ft. 15 RQUNP. • If inn J1 n SHANK, n | nn 3 3*8*11 18* 16 V AGE IN MONTNS. k 45 A 8. Fig. 8. — Distribution of cuts in carcass — Group III. from 18*/2 months on to 45 months. The four-year-old steer shows a decrease to a little over 66 percent. The fatty tissue increases from a little over 3 percent to over 12 percent. 22 Missouri Agr. Exp. Sta. Research Bulletin 54 PISTRiBUTIOM of lea « . FAT, ais* f ONE •« CAF IS ?CA 55. • _a i — -f— 1 m FATTY TISSUE V} \n ao — n r-i o CCAo LEAN FLESH p : I c. O ROUNP s _°nn nn m m n n n 5 o nn run flank. n I— I n n r-i 7.0 10 — — O LGiM. •r to . 5 0 RIB -» 10 2 o 0 PLATE Ik 0 30 — h- z ul J-J 20 — — i 0£ Ui CL to CHUCK ANP HECK s. — S o 1 shinampshank) o i — II — 1 Pr-nHEAPANPTAIL. , i— i i— i 3 5%.8H.ll ZG 3+ 40 44*148 AGE. IN MONTHS. Fig. 11. — Distribution of lean in cuts of carcass — Group II. In Group II (Figure 11) the head and tail, shin and shank, rib, flank, and rump have about the same percent of the total lean as shown by Group I. The chuck and neck and the round contain relatively more in the Group II steers, while the plate and loin con- tain relatively less. The differences are, however, not very great in any case being generally within 2 percent. The plate shows Studies In Animal Nutrition — II 25 an increase with increasing age while the round shows a decrease. The other cuts show little effect of age. With the Group III animals (Figure 12) the head and tail, shin and shank, flank, and rump have about the same proportion of total lean as shown by the other groups. The head and tail in 20 PI STf OB UTION o >r LU iti FLESH 00 ip j m. 10 o ROUNP. F onr-inn n n Rump. n n n 5 o nm nn m n flank. , n n 10 10 pi p p 0 LOIN. 10 i J RI3. -» lo -J f . — PLATE! . »- 30 w 0 \~ Z. 10 tel u or u 10 CHUCK AHQ £L 0 NECK. 5 i— SHINahp SHANP S o . nnnrn r-n 1 TAIL., — , „ 3 5 0\ U Tgw 26 4 0\ 45 46 AGC IN MONTHS. Fig. 12. — Distribution of lean in cuts of carcass — Group III. some cases runs 1 percent higher and the shin and shank 2 percent higher. The plate, rib, loin, and round vary over wider limits while the chuck and neck keep within narrower limits. The round varies from 20 to 26 percent, the loin from 14.5 to 19 percent, the plate from 7 to 12 percent, and the rib from 7 to 10 percent. There seems to be no consistent effect of age excepting a reduction of the head and tail and the shin and shank. 26 Missouri Agr. Exp. Sta. Research Bulletin 54 DISTRIBUTION OF TOTAL FAT FLESH The distribution of the total fatty tissue exclusive of the offal fat is shown in Figures 13 to 15 (Tables 36 to 40 in the Appendix). There is much greater variation, and age and fatness affect the distribution of fatty tissue quite markedly. In Group I (Figure 13) the head and tail contains 3 to 4 per- cent of the fatty tissue in the young animal and less than 0.5 per- cent in the four-year-old steer. The shin and shank start with about 6 percent and have less than 2 percent at four years. The chuck and neck have about 19 percent at three months and 13 to 15 percent at four years. The plate increases from 7.5 percent to 21 percent, and the rib increases from 2 to 10 percent. The kidney fat increases from 10 percent at three months to 16 percent in the baby beef (11 to 18 months) and drops again to 6 or 7 percent in the old animals. The loin increases its proportion from 20 percent to 25 percent. The flank decreases from 11 to 9 percent and the round decreases markedly from 17 to about 7 percent. The rump increases slightly from 3.5 to 5 or 6 percent. With increasing age and fatness, therefore, a greater part of the fatty tissue is found in the plate, rib, loin, and rump and a smaller part in the head and tail, shin and shank, chuck and neck, flank, and round. The kidney fat at first increases and then decreases. In Group II (Figure 14) the head and tail have 5 to 6 percent of the total fat in the young animal and only 1 percent in the old animal. The shin and shank decrease from 8 to 2 percent. The chuck and neck starts at about 18 percent. It then rises to 21 per- cent at 11 months and falls to about 13 percent at 34 months. It then rises again to 21 percent at 44^4 months and decreases to 15 percent at four years. The plate increases its proportion of fat from 9 to 20 percent with advancing age. The rib has none of the total fat at three months and 13 percent at four years. The kidney fat is 9 to 10 percent of the total from three months to 34 months with the exception of the 11-month-old steer which has but 5.5 percent. From 40 months on there is but 6 percent of the total here. The loin, like the chuck and neck, shows two maxima. It increases from 16 percent at three months to 26 percent at 8^4 months. It then decreases to 21 percent. At 34 to 40 months it is 23 percent, only to decrease to less than 21 percent at four years. The flank in general shows a decrease in the proportion of total fat found here. The variations in the rump are not a function of PERCENT or TOTAL FAT FLESH. Studies In Animal Nutrition — II FMSTRI&UTION Of FAT FLESH GROUP I, 10 -£L _l ROVNR o nn nn n RUMP. n FLANK. 15. i — n RIB. —l £ - — t n o 0 PLATE. & o t- C 10 PI pi CHUCK. Ui p K U o AMP HECK. 5 SHIM AMO 0 1 In nn r-,n SHANK, n i — > i ii — i 5 HEAP amp o. nn fin: n — 3 5^ 8’4 11 18 Zl ACE IN TAIL. MONTHS. Fig. 13. — Distribution of fat in cuts of carcass — Group 1. 28 Missouri Agr. Exp. Sta. Research Bulletin 54 age, while the round decreases from 19 percent to 10 percent at 40 months and then increases again to 13 percent. With increasing fatness, then, a greater part of the total fatty tissue is found in the plate and rib. This is true in general of the loin, although PISTRIBUTIOM or FAT FLESH :• groupie. 14- 10 — 10-0 ROUNP. o rinnn n fl n nn 10 0 FLANK x» — 10 o LOIN £ 10 c: 10-0 |-| KIP HEY FAT. n nn 2 o i — i nn RIB □ n •4 zo I .. __ n L * 9 , _ 0 I PLATE. 5 20 U OS kJ a. jo rn 0 CHUCKAHP NECK 5 SHIN ANP 0 n I— II— I SHANK | — , m i l ~~1 r~ i . 5 HEAP ANP ol in rim TA,L ^ n i — i i — i i — i- 3 5*4 26 34 40 44^ 48. ACE IN MONTHS Fig. 14. — Distribution of fat in cuts of carcass — Group II. there are cycles which have higher maxima at younger ages. The chuck and neck show cycles also but have less at the end. The rump and flank show little change with age, while the head and tail, shin and shank, and round decrease. Studies In Animal Nutrition — II 29 Group III (Figure 15) shows greater individual variations than do the other groups. Practically every wholesale cut shows irregu- 2o 0 STRIBUTIOfl of FAT FLESH*- oroup m. to o ROUNP. c » rii-inn rr RUMP. 10 i— 1 o — FLANK. as 20 10 — 0 LOIN IS o D KIPNEY fat. n n r s eia — uj o i — 1 1 — ii l rn rn ' r~\ rr _i **'2© K io _n PLATE.. 1- u 20 0 1- £ 10 llJ U tc uJ o CHUCK AMP NECK. a. 10 0 m SHIM AMP 1 =. O n 5HAm - n nn 10 0 n HEAP AMP n n n TA,L - n I ™ s. a*. 8^ 11. l»v 2 . 26 . 4«< AGE ’IN MONTHS. it Al 48. Fig. 15. — Distribution of fat in cuts of carcass — Group III. lar changes. The head and tail and the shin and shank decrease rather uniformly while the plate and rib increase uniformly. The round has less at four years than at three months while the loin 30 Missouri Agr. Exp. Sta. Research Bulletin 54 has more, but there are many ups and downs in between. The changes in the other cuts do not consistently follow age. DISTRIBUTION OF TOTAL SKELETON Figures 16 to 18 (Tables 41 to 45 in the Appendix) show the changes in the distribution of the total skeleton for the three groups PISTRISUTIOM of SKELETON •* croup x. 10 0 ROUMP 5 nrinn nfl iwmr fl l~l fin o FLANK. 10 0 LOIN. 10 s — r-i o RIR. 10 • 0 — PLATE. z: o 20 1- kJ -1 UJ to CHUCK X S? o ANP NECK. cT 20 1— u o io j-i r-i SHIN rn H E UJ o ANP SHANK. o tc UJ a. io [-» 0 FEET. 10 __ j-j o HEAP anp TAIL. 3 5\ 8\ 11 18 21 35 Z9\ 4^49 AGE IN MONTHS. Fig. 16. — Distribution of bone in cuts of carcass — Group 1. with advancing age. The values are much more constant than for either the lean flesh or the fatty tissue. In Group I (Figure 16) the head and tail have about 13 percent Studies In Animal Nutrition — II 31 of the total skeleton with variations from about 11.5 to 14.5. The feet bones increase from about 12.5 percent at three months to 15 percent at 8 months and then decrease to 9.5 percent at four years. The shin and shank decrease rather uniformly from 17.5 PISTRJSUTiON ©? SKELETON *• oroupst. 10 0 ~ s . nn nd □ □ n n n 1 FLANK. 10 . LOiN. 18 _SJ j— 1 o RS9. 10 8 — o PLATE 20 £ g CHUCK AMP id X 0 NECK. 10 . ?o -1 2 Q 10 pi p| SHIN 1- W « n AMP SHANK. h r u Id FEET. n L 10 pi pi HEAP 0 AMP TAIL. 3 5*4 8\U 26 34 40 44k 48 AGE IN M0NTN5. Fig. 17. — Distribution of bone in cuts of carcass — Group II. to 15.5 percent. The chuck and neck bones increase from 18 to 21 percent at 18 months and then decrease to 19.5 percent. The plate bones vary from 7.5 to 9 percent increasing slightly with age. The rib varies around 8 percent and the loin around 11 percent. The flank has but a tip of rib bone in it and varies from a few hun- 32 Missouri Agr. Exp. Sta. Research Bulletin 54 dredths of a percent to about 0.2 percent increasing with age. The rump increases slightly from 3 to 5 percent, while the round de- creases very slightly from 9.5 to 9 percent. PI 5 TRIBUTIC 10 IN or SKELETON » growpES. 0 ROUNP. S o nn nm n n RuriP - n (in “ ' 1 FLAnK. 5 i—| 0 LOIN. 10 6 i—i rn 0 RIB. 10 s r- 1 S 1 1 PLATE. 6 * _ bl X 40 to chuck g . AMP NECK. zo 1* e E IP — SHIN Id U 0 t Id o AN? SHANK. 0. to __ 0 FEET. to — — HEAP 9 AMP TAIL. J 5 ^ 8V t 11 18*4 26 40 l / z 45 AGE. IN MONTHS. 48 Fig. 18. — Distribution of bone in cuts of carcass — Group III. Groups II and III (Figures 17 and 18) show somewhat greater variations but the range of the figures is much the same. The in- crease in the proportion of skeleton found in the feet at 8^2 months is not seen. In other respects the figures are much as those for Group I. Studies In Animal Nutrition — II 33 DISTRIBUTION OF LEAN, FAT, AND BONE IN THE WHOLESALE CUTS The Shin (Fore Shank). — With the Group I animals (Figure 19) the lean forms about 50 percent of the shin, the fat 5 to 18 per- cent and the bone 48 to 34 percent. The percent of fat increases with age and fatness and the bone decreases, while the lean varies. With the Group II animals the lean forms 44 to 53 percent of the cut, the fat 3 to 6 percent, and the bone 50 to 40 percent. The lean increases with age while the bone decreases. The fat changes but little with age. With the thinnest animals — Group III — the lean is 46 to 57 percent of the cut, the fat 3 to 5 percent, and the bone 50 to 40 per- cent. There are more irregularities shown in this group, but in general the lean increases in percent with age while the bone de- creases. The Neck. — The neck of the Group I animals (Figure 20) con- tains from 66 to 40 percent lean, 12 to 33 percent fat, and 18 to 30 percent bone. The fat increases with increasing age and fatness while the lean decreases. The bone varies between the limits given without respect to age. The amount and composition of this cut will depend much upon the general conformation and fatness of the carcass. There will then be considerable differences between individuals. In the Group II animals the neck is 50 to 67 percent lean, 4 to 14 percent fat, and 23 to 39 percent bone. The percent of lean, of fat, or of bone, does not seem to depend on age. The Group III animals show even greater variations and no better correlation between age and composition. The Chuck. — The chuck (Figure 21) of the Group I animals has 72 to 58 percent lean, 4 to 30 percent fat and 23 to 12 percent bone. The lean and bone decrease with age while the fat increases. With the Group II animals the lean is 70 to 75 percent of the cut, the fat 3 to 12 percent and the bone 26 to 17 percent. The fat increases with age, the bone decreases, and the lean is about con- stant. In the case of the Group III animals the lean runs 70 to 75 percent, the fat 2 to 8 percent and the bone 24 to 19 percent. The fat increases slightly with age, the bone decreases slightly while the lean is fairly constant. The Plate. — The plate (Figure 22) becomes rather a fat cut 34 Missouri Agr. Exp. Sta. Research Bulletin 54 Fig. 19. — Distribution of lean, fat and bone in the shin. PERCENT of WHOLESALE CUT. Studies In Animal Nutrition — II 35 LEAN FAT AMD BONE IN THE NECK. 10 ” PI — 0 BONE. D a FAT. n 40 — 20 GROUP IL. o LEAN. 3 55t 8J411AGEIMMOMTHS.2.6 34 40 44\4& o c£ xo — UJ Q. to 0 BOME. 30 xo GROUP I 10 o FAT. 60 40 — r-j 10 0 LEAN. 3 SK 8’* U A&£ IQ 21 1(1 MOMTMS- 35 33\ 44S 47 Fig. 20. — Distribution of lean, fat and bone in the neck. 36 Missouri Agr. Exp. Sta. Research Bulletin 54 10 L£ IA M F AT AMD B0ME IM THE CHUCK. 0-10 BONE. r 0 < — >1 — 1 1 — n n FAT ‘ r — i r 65 60 4Q_ GROUPIE. 0 LEAN. 2o. 3 5H 8 * 11 18* AGE 26 IM MONTHS.40* 45 43 10 0 BONE. L 10 r a o ^nnn FAT - n n Mil o Ui 60 — -i _l 55 u 40 GROUP H. J O 5 20 > Ll o 0 LEAN. \r 5 3 5* 8*11 ASE IN M0NTHS.26 34 40 44^48 £ io i o i — il i FAT. < 60 10 GROUP n. o z ?„ u o o LEAN. h Z sn 3 5%8% UAGE IN M0MTHS26 34 40 44% 4 6 U O 20 * Id 0. o BONE. 40 X ^ A i—i r-i FAT. 60 r— i 40 GROUP I. i/t o LEAN. 3 5% 854 U AGE 18 Z1 IN MONTHS 35 39% 44% 47 Fig. 23. — Distribution of lean, fat and bone in the rib. 40 Missouri Agr. Exp. Sta. Research Bulletin 54 LEA? 20 1 FAT AMO BONE IN THE LOIN. 0 BONE. 10 o r~ir~i f~1 FAT n 6o — r-j 40 GROupnr. 20 0 LEAN. . 10 3 5^ 8'^ 11 r 18'* AGE U IN MC »N THS.- n 1 4! n ■ 46 t; L BONE. E L_ 1 ° 20 *i 10 , , — < 01 — ll 1 FAT. hi -1 O 60 i-i — — — 1 i 40 GROUPIE O £ - Id O £ o LEAN. 0. 10 A 3 5>t ffill A6E 111 M0NTHS.26 34 40 44! nfln QD Bone, n n n 4 r 48 40 20 GROUP I. O — FAT. 60 __ 1 r— | 40 20 0 LEAN. 3 5\ St U AGE 18 21 IN MONTHS. 35 39*1 44k4-7 Fig. 24. — Distribution of lean, fat and bone in the loin. Studies In Animal Nutrition — II 41 Fig. 25. — Distribution of lean, fat and bone in the flank. 42 Missouri Agr. Exp. Sta. Research Bulletin 54 II steers the lean decreases from 72 percent to 33 percent while the fat increases from 25 to 65 percent. With the Group III steers the lean decreases from about 83 to 60 percent, while the fat in- creases from 14 to 38 percent. In all groups then the fat increases with age while the lean decreases. The Rump . — The changes in the composition of the rump are shown in Figure 26. With the Group I animals the lean decreases from 59 to about 30 percent while the fat increases from 8 to 56 percent. The bone decreases from 33 to 14 percent. With the Group II animals with one striking exception the lean decreases with age running from 50 to 56 percent down to 44 percent. The fat increases from 8 to 32 percent, while the bone decreases from 41 to 23 percent. In the case of the Group III steers the lean decreases from about 58 percent to 49 percent, and the fat increases from 5 to 21 percent. The bone increases at first from 36 to 39 percent and then decreases to 28 percent. The Round. — The round shows rather small relative changes (Figure 27). The Group I steers have from 78 to 63 percent lean, from 5 to 28 percent fat, and from 16 to 9 percent bone. The Group 11 animals have about 80 percent lean in all cases but the oldest. The fat runs from 5 to 16 percent and the bone from 17 to about 12 percent. In Group III the lean runs between 77 and 81 percent, the fat increases from 4 to 9 percent and the bone decreases from 18 to about 12 percent. The Shank. — The shank also shows small relative changes in make up. Figure 28 gives the distribution. The Group I steers have about 30 percent lean with a few animals running higher. The fat increases from 2 to 16 percent, but the next to the oldest shows 22.5 percent. The bone decreases from 66 to about 50 percent. With the Group II steers the lean is fairly constant after 11 months at about 36 percent. At 3 months it is 29 percent. The fat is small and increases irregularly with age. The bone decreases from 69 percent to a value running around 60 percent from 11 months on. The Group III animals show much the same thing as the Group II steers. This is a very bony cut and after 8^4 months it is not much af- fected by age. PERCENT OF WHOLESALE CUT. Studies In Animal Nutrition — II 43 LEAN FAT AND BONE IN THE RUMP. — iO o BONE. 20 to n i or~i r— | n 1 i 1 FAT. JO 44 10 group nr. LEAN. O M. ^ 3 5^8 11 18iiA6E 26 IN MONTHS. 40\ 4? 48 7a A BONE. so ao 10 «n FAT. 40 pi p| 20 GROUP n. 0 LEAN. 30 3 SK 8M1AGEJN MONTHS.Zfe 34 40 44 *l 48 20 — |— | to 0 BONE. 40 — xo r—, eo.oI3 FAT. > — 1 GROUP I. ip LEAN. O l_t.Mii. 3 5V8*UA6E18 21 IN M0NTHS.3S 39’^ 44V47 Fig. 26. — Distribution of lean, fat and bone in the rump. 44 Missouri Agr. Exp. Sta. Research Bulletin 54 lo LEAM , 1 ri fin "AT , * mi => BO I s IE m THE R :oi UMP. o BONE. zn: i2 T or-ii-i i — ii — i n n fax n M L fio 6o GROUP m. 40 20 LEAM. O 7Q AGE IM MOUTHS. IQ — 0 BOME. r 10 8 0-0 n — — 1 FAT. 9 O j 60 -J 40 GROUP XT. O E * 2 o LEAM. w o K 0 Z 3 5*4 6'£ 11 Ate IM MONTHS 26 34 40 ^ 20 14* 48 u S to — “■ 0 1 1 1 DOME. nn ?o n 10 n 8«-o 1 1 FAT. 6o — GROUP I. 40 ?o LEAM. o 3 5\ 8^ 11 AGE 18 21INM0nTtt5. 35 39^ 44*47 Fig. 27. — Distribution of lean, fat and bone in the round. PERCENT of WHOLESALE CUT. Studies In Animal Nutrition — II 45 LEAN FAT ANP BONE. IN THE SHANK. 60 — Aq GROUP m. 2o OlO BONE. o f 1 | 1 I — i a n ^ 1—3 1— 1 I~1 30 20 LEAN. 0 6 o 3 5% 11 18* AGE 26 ir 40* 45 1 MONTHS. 48 40 20 GROUP xc. Olo BONE. o. — ■! 1 1 ii — i fat. r — i rn r — -i □ i i 3o i— 1 n r- 20 0 LEAN. 6 o 3 5^ i av 2 11 26 34 AO 44* 48 AGE IN MONTHS. AO pi GROUP X. 20 0 BONE. 10 0 i — 1 1 1 n rn FAT. 3o — 2q_ 0 LEAN. 1 ■ ft m i a- AGE. IN MONTHS. Fig. 28. — Distribution of lean, fat and bone in the shank. 46 Missouri Agr. Exp. Sta. Research Bulletin 54 SUMMARY Hereford-Shorthorn beef steers were fed on three planes of nutrition : Group I, full fed from birth ; Group II, fed for maximum growth without fattening giving gains of one pound per day for the first two years ; Group III, fed for scanty or retarded growth gaining about .69 pounds per day for the first two years. The slaughter house data for 31 animals are presented. With these animals the ratio of empty weight to live weight is greatest at birth, least at 8 y 2 months, and intermediate at 4 years. A lower plane of nutrition gives a lower ratio. The ratio of carcass to live weight decreases to 8 y 2 months then increases to a maximum at 3 to 4 years. The better fed animals have the greater percent. On the empty weight basis the carcass continuously increases to 3 to 4 years. The proportions of hide and hair, heart, brain and spinal cord, and intestinal length to empty weight decrease with increasing age and fatness. The blood, lungs, stomachs, intestines (weight), liver, kidneys, spleen, and pancreas increase relatively during the early months and then decrease. The maxima occur generally at 8 y 2 months. The stomachs and liver show marked retardation on the low planes of nutrition. The proportions of skeleton and of total organs are greatest at birth and the total fleshy parts at 4 years. In the carcass the proportions of loin, rump, flank, and plate increase with increasing age and fatness of the steers. The rib changes but little while the round, chuck and neck, and shin and shank decrease. The distribution of the total lean flesh is but little affected by age and fatness excepting that there is a slight reduction in the proportion found in the shin, shank, head and tail and in some cases in the round. The plate shows a larger part of the total as the animal grows and fattens. The proportion of the total fat flesh found in the plate, rib, loin, and in some cases in the rump increases with increasing fat- ness. Age and fatness influence the distribution of the total skele- ton but slightly. The feet bones tend to increase relatively up to 8 Yz months (Group I) and then decrease. The chuck and neck Studies In Animal Nutrition — II 47 bones show a similar phenomenon with the maximum at 18 months. The rump shows a slight increase. The composition of the wholesale cuts of meat is affected by increasing age and fatness. In general the percent of fatty tissue increases and the percent of bone decreases. The percent of lean flesh may increase, remain constant, or decrease, but on the aver- age it decreases. The increases in percent of fat are greatest in the plate, loin, and flank. The rib and rump show rather large in- creases. 48 Missouri Agr. Exp. Sta. Research Bulletin 54 APPENDIX Table 1. — Slaughter House Weights of Offal Parts (.in Grams). Steer 556 554 555 557 552 548 3 3 3 5 Months 5 Months 5 Months Age Months Months Months 17 Days 7 Days 9 Days Group 1 2 3 1 2 3 Live weight 111,489 87,456 84,725 204,934 116,491 99,255 Blood 6,124 4,197 4,529 8,952 5,219 4,603 Heart, pericardium, arteries 993 591 554 2,342 1,056 753 Heart, marketable 521 335 315 928 561 427 Heart, lean 432 319 295 760 561 385 Lungs and trachea 1,272 907 937 1,972 1,096 1,084 Brain 275 273 252 351 332 374 Spinal cord 123 122 101 200 134 152 Tongue, including bones and larynx 884 660 432 908 673 571 Tongue, marketable 466 358 247 650 464 428 Tongue bones, including larynx 106 142 53 131 96 88 Gullet 209 146 193 189 355 204 Stomachs 1,975 1,208 1,338 4,341 1,938 1,885 Rumen 999 616 745 2,614 1,100 975 Reticulum 260 147 147 358 188 203 Omasum 391 161 176 724 374 363 Abomasum 325 284 270 645 276 344 Intestines, small 1,944 1,474 1,651 2,863 1,781 1,660 Intestines, large 750 603 792 1,233 1,003 861 Intestines, small; length, cm 2,878 2,452 2,080 3,139 2,630 2,806 Intestines, large; length, cm 540 450 380 667 491 550 Neck sweetbread 173 193 129 359 143 109 Heart sweetbread 155 143 30 264 122 59 Spleen 300 202 167 485 284 215 Pancreas 96 76 106 225 101 85 Liver 1,760 1,166 1,240 3,003 1,181 1,104 Gall bladder and gall 25 23 8 148 55 34 Gall 87 40 Kidneys 338 530 439 649 316 353 Urinary bladder 46 58 63 76 41 53 Penis 110 116 59 155 94 79 Diaphragm 166 70 51 199 92 98 Caul fat 417 170 107 1,981 363 Stomach fat 146 73 20 1,486 469 275 Intestinal fat 839 401 335 3,290 952 570 Hide and hair 10,314 7,400 6,580 14,100 10,532 8,328 Dewclaws 32 28 24 64 66 36 Teeth 218 225 190 261 278 264 Horns 9 46 36 40 Hoofs 358 301 274 585 448 359 Right fore foot and hoof 761 666 610 1,035 828 728 Left fore foot and hoof 760 653 630 1,007 807 728 Right hind foot and hoof 719 651 596 1,005 882 740 Left hind foot and hoof 707 734 595 1,096 792 659 Fore quarter, right 15,436 13,096 12,075 28,483 16,023 13,944 Bind quarter, right 14,849 12,358 10,596 27,438 15,135 12,764 Left half 30,129 24,675 22,764 54,150 30,580 27,275 Studies In Animal Nutrition — II 49 Table 2. — Slaughter House Weights of Offal Parts (in Grams). Steer ._, >r • 547 550 558 541 538 540 Age 8 Months 5 Days 8 Months 14 Days 8 Months 12 Days 10 Months 22 Days 10 Months 26 Days 11 Months 2 Days Group 1 2 3 1 2 3 Live weight 206,175 147,202 108,191 323,836 180,930 158,131 Blood 8,711 7,080 4,666 12,470 7,219 6,967 Heart, pericardium, arteries 1,624 1,163 971 2,646 1,670 1,436 Heart, marketable 771 556 506 1,087 873 673 Heart, lean 666 481 425 940 698 572 Lungs and trachea 1,868 1,460 1,111 2,387 1,825 1,501 Brain 346 319 373 383 343 361 Spinal cord 113 124 147 185 144 151 Tongue, including bones and larynx 800 834 650 2,336 1,513 1,547 Tongue, marketable 586 523 464 1,209 779 811 Tongue bones, including larynx 161 139 111 309 221 170 Gullet 384 209 331 538 400 324 Stomachs 5,190 3,833 2,550 7,876 4,521 3.894 Rumen 3,127 1,914 1,329 4,050 2,209 1,966 Reticulum 373 366 253 866 751 470 Omasum 1,072 1,015 579 1,885 1,013 878 Abomasum 618 538 389 1,075 548 480 Intestines, small 2,649 2.303 1,599 3,922 2,327 1,954 Intestines, large 1,826 1,321 1,018 1,322 1,319 1,305 Intestines, small; length, cm 2,900 2,755 2,568 3,343 3,393 2,510 Intestines, large; length, cm 663 620 531 716 610 772 Neck sweetbread 372 189 73 396 252 194 Heart sweetbread 319 147 59 344 229 214 Spleen 391 291 193 596 331 331 Pancreas 200 200 153 390 208 180 Liver 2,851 1,992 1,352 3,832 1,978 1,593 Gall bladder and gall 138 117 28 275 153 83 Gall 103 86 7 202 122 58 Kidneys 450 379 318 645 487 363 Urinary bladder 83 60 48 153 123 134 Penis 181 112 149 226 146 131 Diaphragm 198 170 116 176 131 136 Caul fat 1,118 532 150 4,212 939 578 Stomach fat 851 781 248 1,803 799 469 Intestinal fat 1,910 1,297 431 4,322 1,714 1,260 Hide and hair 14,618 10,440 8,138 26,576 15,342 12,994 Dewclaws 86 53 38 99 65 49 Teeth 310 228 274 304 240 278 Horns 77 112 31 468 250 304 Hoofs 574 384 374 770 570 432 Right fore foot and hoof 1,177 820 764 1,270 931 803 Left fore foot and hoof 999 780 719 1,303 935 813 Right hind foot and hoof 1,185 778 771 1,368 937 805 Left hind foot and hoof 1,054 765 719 1,340 935 819 Fore quarter, right 27,711 19,194 14,089 46,507 25,401 21,605 Hind quarter, right 28,561 18,582 13,921 47,873 24,824 20,820 Left half 55,406 36,500 27,259 95,368 46,859 44,904 50 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 3. — Slaughter House Weights of Offal Parts (in Grams). Steer 505 503 532 531 504 523 525 Age 10 mo. 11 mo. 1 yr. 5 mo. 1 yr. 6 mo. lyr.8mo. 2yr.2mo. 2yr.2mo. 18 da. 11 da. 20 da. 12 da. 26 da. 6 da. 8 da. Group 1 2 1 3 1 2 3 Live weight 313,317 270,566 517,093 212,466 526,164 380,880 305,112 Blood 13,810 13,058 18,752 9,457 21,005 15,287 13,614 Heart, pericardium, arteries 2,106 2,845 4,843 1,971 3,065 3,004 1,320 Heart, marketable 1,056 1.251 2,058 1,011 1,632 1,477 1,169 Heart, lean 938 1,063 1,685 791 1,428 1,188 1,005 Lungs and trachea 2,498 2,549 3,870 1,915 3,628 3,371 1,658 Brain 376 442 406 373 423 475 467 Spinal cord 161 224 237 200 301 322 244 Tongue, including bones and larynx 1,989 1,365 3,195 1,728 3,904 3,702 2,554 Tongue, marketable 1,115 789 1,543 894 1,587 1,678 1,276 Tongue bones, including larynx 190 204 340 207 367 404 252 Gullet 311 442 744 494 455 803 525 Stomachs 8,818 5,765 10,893 5,343 12,820 9,342 8,849 5,943 2,740 1,272 593 Omasum 2,208 1,117 Abomasum 1,470 893 Intestines, small | 4,852 4,736 | 3,792 2,346 | 9,526 5,226 5,960 Intestines, large 2,132 1,342 Intestines small; length, cm 4,460 4,039 4,389 2,926 4,125 3,797 Intestines large; length, cm 909 975 762 1,054 866 Neck sweetbread 425 348 321 236 396 272 150 Heart sweetbread 337 399 120 236 532 341 145 Spleen 599 661 884 481 1,317 847 556 Pancreas 300 289 630 297 295 300 151 Liver 3,983 3,646 5,694 2,205 4,754 3,268 2,876 Gall bladder and gall 215 201 284 114 697 146 126 Gall 185 86 Kidneys 718 655 868 506 877 703 664 Urinary bladder 80 108 308 223 254 254 193 Penis 251 288 302 240 310 196 157 Diaphragm 204 917 626 270 1,819 362 343 Gaul fat 8,735 686 2,156 1,132 Stomach fat 6,344 3,771 4,940 586 12,272 2,156 1,180 Intestinal fat 6,437 3,614 10,022 1,627 12,833 3,603 2,649 Hide and hair 23,026 23,120 33,988 16,693 41,144 33,097 27,813 Dewclaws 142 112 158 90 198 115 88 Teeth 268 253 494 426 338 766 690 Horns 342 285 228 1,272 1,167 1,298 Hoofs 722 661 1,248 700 1,062 948 852 Right fore foot and hoof 1,218 1,456 1,937 1,095 1,767 1,609 1,380 Left fore foot and hoof 1,218 1,469 1,968 1,096 1,815 1,664 1,380 Right hind foot and hoof 1,251 1,350 1,932 1,136 1,736 1,649 1,320 Left hind foot and hoof 1,251 1,383 1,878 1,137 1,805 1,608 1,289 Fore quarter, right 45,336 37,450 79,605 29,683 76,580 56,114 43,204 Hind quarter, right 43,630 36,954 76,799 29,429 81,101 53,785 39,581 Left half 87,885 71,832 156,225 59,900 148,728 Studies In Animal Nutrition — II 51 Table 4. — Slaughter House Weights of Offal Parts (in Grams). Steer 515 507 529 527 526 524 2yr. 9 mo. 2 yr. 9 mo. 3 yr. 2 mo. 3 yr. 3 mo. 3 yr. 4 mo. 3 yr. 4 mo. Age 19 days 16 days 21 days 15 days 13 days Group 1 2 1 1 2 3 Live weight 743,361 457,155 690,704 842,841 479,846 362,260 Blood 27,856 20,316 23,028 27,382 18,957 17,019 Heart, pericardium, arteries 5,787 4,278 5,713 8,891 3,998 2,953 Heart, marketable 2,664 2,061 2,045 3,246 1,673 1,585 Heart, lean 1,890 1,556 1,419 2,370 1,246 1,369 Lungs and trachea 3,653 3,768 4,272 4,326 3,797 3,455 Brain 483 482 443 407 469 508 Spinal cord 245 262 261 294 191 250 Tongue, including bones and larynx 4,229 3,697 4,021 5,734 3,626 3,602 Tongue, marketable 2,309 1,791 1,603 1,904 1,769 1,666 Tongue bones, including larynx 484 467 441 420 438 435 Gullet 627 709 829 1,090 777 1,340 Stomachs 13,171 9,620 12,101 10,347 10,985 8,096 6,007 5,790 6,091 4,602 1,069 889 896 775 3,179 2,361 2,460 1,457 Abomasum 1,846 1,307 1,538 1,262 Intestines, small j> 6,823 2,985 2,539 2,257 1,964 Intestines, large OylJ&O \ 2,304 2,508 1,877 2,032 Intestines, small; length, cm j. 4 694 q qaoJ 3,818 3,861 3,515 2,911 Intestines, large; length, cm 1,036 1,072 925 747 Neck sweetbread 484 325 514 464 269 267 Heart sweetbread 447 318 552 573 170 274 Spleen 1,482 1,132 1,049 1,226 831 757 Pancreas 424 231 824 849 498 435 Liver 5,982 3,788 5,374 5,720 3,531 3,019 Gall bladder and gall 382 130 280 149 231 296 Gall 190 10 143 229 Kidneys 1,065 752 1,006 1,244 922 766 Urinary bladder 347 282 257 295 312 361 Penis 225 239 436 300 250 365 Diaphragm 1,107 1,036 645 1,276 807 532 Caul fat 9,263 3,859 13,993 14,263 2,763 1,068 Stomach fat 6,965 3,097 10,951 9,669 2,815 1,244 Intestinal fat 13,649 4,351 14,112 24,585 5,973 2,695 Hide and hair 49,943 34,473 45,567 46,240 35,732 30,092 Dewclaws 201 152 241 240 227 194 Teeth 786 712 872 782 806 Horns 1,804 1,600 2,200 1,266 1,427 1,227 Hoofs 1,692 1,338 1,934 1,648 1,300 Hight fore foot and hoof 2,346 1,936 2,213 2,293 2,031 1,864 Left fore foot and hoof 2,312 1,938 2,230 2,321 1,984 1,867 Right hind foot and hoof 2,473 1,884 2,331 2,265 1,862 1,791 Left hind foot and hoof 2,607 1,900 2,362 2,306 1,863 1,815 Fore quarter, right 115,806 72,574 111,434 147,674 75,917 54,798 Hind quarter, right 112,738 65,693 112,704 141,372 70,858 50,394 Left half 229,567 304,508 52 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 5. — Slaughter House Weights of Offal Parts (in Grams). Steer 513 502 509 501 512 500 3 yr. 8 mo. 3 yr. 8 mo. 3 yr. 8 mo. 3 yr. 3yr. 11 mo. . 3yr. 11 mo. Age 15 days 19 days 22 days 11 mo. 21 days 26 days Group 1 2 3 1 2 3 Live weight 854,650 506,878 439,814 883,480 548,050 457,786 Blood 25,680 19,728 18,291 28,710 24,176 21,269 Heart, per cardium, arteries 7,427 4,451 3,645 6,177 5,158 3,414 Heart, marketable 2,311 1,946 1,660 2,214 1,955 1,467 Heart, lean 2,003 1,680 1,444 1,882 1,555 1,284 Lungs and trachea 4,858 3,696 3,283 3,838 3,881 3,747 Brain 413 465 389 459 466 422 Spinal cord 335 335 350 298 200 410 Tongue, including bones and larynx 4,171 4,107 3,317 5,294 4,215 3,541 Tongue, marketable 1,942 2,198 1,555 2,153 1,766 1,619 Tongue bones, including larynx 455 499 425 500 526 439 Gullet 1,152 1,098 1,081 1,051 973 909 Stomachs 13,446 10,832 9,851 14,185 11,089 10,995 Rumen 7,704 7,216 5,274 8,769 5,867 6,446 1,431 1,064 933 1,068 1 045 Omasum 2,951 2,218 2,337 2,976 2,643 2,419 Abomasum 1,360 1,398 1,176 1,507 1,511 1.085 Intestines, small 3,543 2,324 2,186 2,796 3,067 2,645 Intestines, large 2,327 1,829 1,703 2,079 2,255 1,890 Intestines, small; length, cm 4,480 3,545 3,255 3,848 4,481 3,388 Intestines, large; length, cm 1,054 970 901 1,001 1,067 945 Neck sweetbread 581 338 267 335 235 250 Heart sweetbread 753 164 363 449 276 288 Spleen 1,114 921 1,304 1,178 1,255 1,054 Pancreas 873 581 562 836 736 625 Liver 5,920 3,716 3,875 6,161 4,416 4,634 Gallbladder and gall 127 334 225 266 300 300 Gall 37 241 110 176 212 241 Kidneys 1,015 838 774 1,037 1,074 1,019 Urinary bladder 227 283 253 275 366 331 Penis 330 305 274 339 302 271 Diaphragm 811 561 661 779 672 692 Caul fat 24.621 2,738 2,937 14,688 5,545 3,930 Stomach fat 7,860 3,256 2,240 7,432 3,658 2,968 Intestinal fat 21,290 5,383 4,745 16,503 8,251 6.042 Hide and hair 45,286 39,556 37,614 50,090 41,268 35,938 Dewclaws 308 218 196 331 234 247 Teeth 874 1,038 838 778 710 852 Horns 2,144 1,949 3,354 1,810 Hoofs 1,872 1,792 1,394 2,192 1,490 1,848 Right fore foot and hoof 2,452 2,272 1,941 2,466 2,210 2,226 Left fore foot and hoof 2,426 2,277 1,904 2,467 2,184 2,141 Right hind foot and hoof 2,283 2,115 1,828 2,502 2,043 2,117 Left hind foot and hoof 2,432 2,115 1,875 2,569 2,008 2,126 Fore quarter, right 146,424 80,313 70,406 152.353 90,208 71,295 Hind quarter, right 132,093 73,106 63,182 151,476 79,228 64,330 Left half | 278,645 152,147 131,803 305,356 169,239 136,107 Studies In Animal Nutrition— II 53 Table 6. — Slaughter House Weights of Carcass Parts (in Grams). Steer 556 554 555 557 552 548 Age 3 months 3 months 3 months 5 mo. 17 da. 5 mo. 7 da. 5 mo. 9 da Group [l 2 3 1 2 3 Head, total 3,338 2,895 2,833 5,614 4,059 4,033 Lean, total 702 667 765 1,048 795 996 Fat, total 132 141 143 728 308 118 Bone, total 2,504 2,087 1,925 3,838 2,956 2,901 Shin, right 1,765 1,468 1,637 2,465 1,700 1,592 Lean, right 840 645 829 1,159 864 735 Fat, right 81 81 71 194 70 51 Bone, right 851 732 727 1,123 775 792 Neck, right 517 414 502 769 428 527 Lean, right 344 231 287 423 266 287 71 51 149 54 55 Bone, right 100 133 215 206 114 191 Chuck, right 7.600 6,744 5,863 13,088 7,851 6,866 Lean, right 5,465 4,795 4,298 8,783 5,613 5,033 Fat, right 323 188 116 1,718 409 193 Bone, right 1,767 1,733 1,448 2,613 1,840 1,624 Plate, right 3,048 2,245 2,107 6,739 2,940 2,525 Lean right 2,096 1,475 1,461 3,710 1,887 1,655 Fat, right 156 125 42 1,769 288 131 Bone, right 773 628 605 1,241 750 741 Rib right 2.485 2,144 1,915 5,276 3,120 2,412 Lean, right 1,611 1,457 1,256 3,186 1,995 1,570 Fat, right 39 808 57 20 Bone, right 823 677 628 1,286 1,042 807 Loin, right 5,029 4,088 3,102 9,367 5,147 3,864 Lean, right 3,469 2,906 2,309 5,857 3,517 2,834 Fat, right 410 214 137 2,236 521 192 Bone, right 1,134 934 632 1,239 1,096 826 Kidney, Fat, right 210 120 65 1,614 250 142 Flank, right 868 533 549 2,169 935 630 Lean, right 599 384 453 1,117 597 445 Fat, right 236 134 78 1,051 330 170 Bone, right 16 11 13 8 8 16 Rump, right 926 891 703 1,779 987 811 Lean, right 548 453 401 814 551 485 Fat, right 74 73 37 477 112 36 Bone, right 304 366 253 480 320 285 Round, right 6,106 5,064 4,600 10,090 6,230 5,811 Lean, right 4,736 3,959 3,563 7,480 4,915 4,604 Fat, right 343 260 188 1.266 454 260 Bone, right 987 851 826 1,343 813 949 Shank, right 1,478 1,395 1,309 2,064 1,429 1,305 Lean, right 447 400 417 628 495 415 Fat, right 36 26 27 117 59 30 Bone, right 976 959 852 1,305 871 856 Tail, total 172 200 122 332 237 147 Lean, total 80 75 48 106 106 53 Fat, total 33 Bone, total 92 125 74 193 131 94 54 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 7. — Slaughter House Weights of Carcass Parts (in Grams). Steer 547 550 558 541 538 540 Age 8 mo. 5 da. 8 mo. 14 da. 8 mo. 12 da. 10mo.22da. 10mo.26da. 11 mo. 2 da. Group 1 2 3 1 2 3 Head, total 6,461 5,108 4,764 6,939 5,178 4,747 Lean, total 2,532 1,704 1,388 1,388 1,118 890 Fat, total 808 384 312 460 280 344 Bone, total 3,121 3,020 3,064 5,091 3,780 3,513 Shin, right 2,487 1,951 1,705 3,712 2,400 2,228 Lean, right 1,378 991 778 1,927 1,197 1,167 Fat, right 169 65 51 318 96 69 Bone, right 928 876 864 1,473 1,104 976 Neck, right 525 451 343 1,500 856 689 Lean right 303 259 209 976 503 452 Fat, right 115 40 20 183 164 20 Bone, right 109 147 103 338 200 213 Chuck, right 13,902 9,962 7,557 22,782 13,036 10,763 Lean, right 10,100 6,880 5,343 16,871 9,442 7,771 Fat, right 1,300 728 269 2,210 1,033 746 Bone, right 2,451 2,331 1,925 3,310 2,506 2,235 Plate, right 5,804 3,745 2,055 10,165 5,120 4,374 Lean right 3,574 2,424 1,386 6,285 3,197 2,798 Fat, right 1,240 448 136 2,463 829 646 Bone, right 969 762 542 1,415 1,062 923 Rib, right 4,932 3,085 2,380 8,630 3,886 3,418 Lean, right 3,280 2,084 1,593 5,794 2,598 2,361 Fat, right 551 120 28 1,217 213 162 Bone, right 1,086 870 758 1,581 1,077 912 Loin, right 10,373 6,840 4,359 17,833 8,957 7,803 Lean, right 6,788 4,536 2,981 11,708 6,366 5,350 Fat, right 1,986 1,067 300 4,044 1,180 1,154 Bone, right 1,566 1,232 1,069 1,958 1,348 1,275 Kidney, Fat, right 815 378 113 3,028 311 341 Flank, right 2,317 1,101 575 3,725 1,434 973 Lean, right 1,372 656 462 1,670 961 657 Fat, right 927 420 112 2,037 531 311 Bone, right 10 8 4 17 10 8 Rump, right 1,531 1,180 866 2,765 1,436 1,300 Lean, right 740 631 463 1,378 780 764 Fat, right 388 200 52 849 268 218 Bone, right 380 350 339 528 366 311 Round, right 10,871 7,391 6,223 17,065 10,294 8,305 Lean, right 8,546 5,860 4,382 13,500 8,162 6,728 Fat, right 1,075 472 322 1,927 851 405 Bone' right 1,244 1,050 1,047 1,625 1,278 1,175 Shank, right 2,285 1,502 1,537 3,042 2,106 1,810 Lean, right 856 435 432 981 754 629 Fat, right 148 56 42 189 48 16 Bone, right 1,267 1,011 1,053 1,872 1,304 1,152 Tail, total 292 186 98 381 229 217 Lean, total 138 88 18 160 82 36 Fat, total 9 4 54 16 4 Bone, total 144 94 80 206 135 138 Studies In Animal Nutrition — II 55 Table 8. — Slaughter House Weights of Carcass Parts (in Grams). Steer 505 503 532 531 504 523 525 Age 10 mo. 11 mo. 1 yr. 5 mo. 1 yr. 6 mo. lyr.8 mo. 2yr.2 mo. 2yr.2 mo 18 da. 11 da. 20 da. 12 da. 26 da. 6 da. 8 da. Group 1 2 1 3 1 2 3 Head, total 6,977 8,321 10,510 6,251 11,035 9,848 8,536 Lean, total 1,494 1,728 2,486 1,310 2,656 2,500 2,078 Fat, total 814 886 1,406 280 648 306 508 Bone, total 4,669 5,707 6,514 4,487 7,675 6,832 5,758 Shin, right 3,703 3,330 6,742 3,107 6,123 4,941 4,328 Lean, right 2,017 1,604 3,544 1,602 3,176 2,671 2,421 Fat, right 205 147 468 112 751 263 199 Bone, right 1,479 1,568 2,703 1,393 2,164 1,995 1,681 Neck, right 1,044 1,159 1,476 1,292 1,536 1,391 1,552 Lean, right 641 548 839 930 844 943 1,025 Fat, right 135 181 277 118 233 58 117 Bone, right 264 430 370 257 454 378 398 Chuck, right 21,957 18,821 38,288 15,303 34,645 28,898 20,626 Lean, right 15,877 14,040 27,000 11,524 23,092 21,723 15,605 Fat, right 2,519 1,198 4,986 879 5,735 2,436 1,165 Bone, right 3,497 3,384 6,137 2,792 5,134 4,515 3,827 Plate, right 9,943 7,580 18,479 5,483 18,905 11,995 8,556 Lean right 5,861 4,398 10,017 3,585 9,617 7,704 5,651 Fat, right 2,586 1,475 5,657 609 6,963 2,028 1,174 Bone, right 1,478 1,676 2,689 1,248 2,249 1,914 1,695 Rib, right 8,607 6,625 14,483 4,483 15,286 8,753 8,134 Lean, right 5,650 4,466 8,678 3,137 9,253 6,016 5,833 Fat, right 1,320 420 3,097 169 3,385 761 332 Bone, right 1,596 1,691 2,611 1,140 2,546 1,922 1,944 Loin, right 15,622 13,475 27,366 9,641 29,121 18,844 13,241 Lean, right 9,843 8,603 18,068 7,039 16,838 12,917 9,355 Fat, right 3.779 2,873 7,477 1,132 9,170 3,188 1.879 Bone, right 1,937 1,917 3,123 1,417 2,925 2,661 1,964 Kidney, Fat, right 2,877 1,063 5,867 363 5,700 1,555 629 Flank, right 3,665 2,330 6,692 1,253 8,280 3,795 2,598 Lean, right 1,912 1,480 2,934 861 3,778 2,279 1,704 Fat, right 1,738 792 3,710 372 4,467 1,481 852 Bone, right 20 42 50 25 37 27 32 Rump, right 2,786 2,146 5,190 1,783 6,631 3,627 2,876 Lean, right 1,300 1,012 2,570 1,001 2,884 1,799 1,589 Fat, right 914 591 1,459 252 2,539 910 488 Bone, right 553 525 1,141 530 1,214 885 771 Round, right 15,380 14,600 25,006 13,349 25,995 21,613 16,843 Lean, right 11,592 11,499 19,032 10,748 18,619 16,950 13,762 Fat, right 2,071 1,109 3,032 745 4,909 2,278 981 Bone, right 1,698 1,928 2,812 1,820 2,404 2,315 2,023 Shank, right 2,885 3,043 4,682 2,656 4,817 3,874 3,024 Lean, right 1,019 991 1,773 971 1,541 1,324 1,020 Fat, right 162 171 219 55 772 84 152 Bone, right 1,693 1,874 2,675 1,606 2,469 2,436 1,820 Tail, total 520 385 675 280 691 630 468 Lean, total 214 150 318 120 222 256 178 Fat, total 58 24 50 12 64 38 24 Bone, total 240 190 266 148 316 316 224 56 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 9. — Slaughter House Weights of Carcass Parts (in Grams). Steer 515 507 529 527 526 524 2 yr. 9 mo. 2 yr. 9 mo. 3 yr. 2 mo. 3 yr. 3 mo. 3 yr. 4 mo. 3 yr. 4 mo. 19 da. 16 da. 21 da. 15 da. 13 da. Group 1 2 1 1 2 3 Head, total 14,869 11,874 13,249 12,838 11,643 10,915 2,990 2,370 3,362 2,426 2,382 2,484 1,352 1,246 822 740 9,395 8,152 8,032 8,395 7,593 Shin, right 8,370 6,313 8,275 8,824 6,789 5,333 3,822 3,519 4,371 3,704 2,631 1,538 258 1,281 248 189 3,009 2,510 3,170 2,820 2,475 Neck, right 2,288 1,611 1,587 1,832 1,812 1,473 1,110 998 729 1,047 902 667 179 603 333 38 508 417 508 425 526 Chuck, right 49,301 36,460 47,355 62,001 35,906 28,236 31,074 27,311 36,160 25,478 21,342 11,216 2,777 18,728 3,756 911 6.756 6,184 6,927 6,496 5,922 Plate, right 34,875 16,603 35,901 46,616 18,103 10,486 Lean right 13,189 10,064 17,706 10,740 6,548 Fat, right .... 18,346 3,390 25,650 4,110 1,023 Bnne right 3,124 3,066 3,019 3,217 2,895 Rib, right 21,008 11,635 18,316 28,401 13,371 9,216 Lean, right 9,508 7,894 8,995 12,930 8,632 6,403 Fat, right 8,142 1,216 6,691 12,139 1,860 169 Bone, right 3,232 2,525 2,576 3,273 2,845 2,655 Loin, right 43,844 23,374 43,272 55,188 25,010 16,656 Lean right 20,810 14,862 25,070 15,720 12.100 Fat right 19,162 5,094 26,362 5,817 1,222 Bone right 3,892 3,253 3,570 3,422 3,293 Kidney, Fat, right 4,961 2,188 5,112 9,482 1,612 383 Flank, right 12,112 4,697 13,514 16,167 4,933 2,407 Lean right 3,281 2,459 4,823 2,241 1,563 Fat, right . 8,753 2,152 11,310 2,671 760 Bone right . 60 73 22 77 68 Rump, right 10,005 5,140 9,344 13,721 5,748 3,232 Lean right 3,443 2,521 4,450 2,804 1,616 Fat, right 4,462 1,347 7,560 1,493 399 Bone, right 2,037 1,268 1,630 1,419 1,212 Round, right 34,172 25,274 40.959 39,746 28,102 23,013 Lean, right 21,471 19,651 25,698 22,307 18,857 Fat, right 9,529 2,689 10,733 2,508 1,263 Bone, right 3,172 2,932 3,223 3,176 2,939 Shank, right 6,92(2 4,865 * 6,434 4,865 4,202 Lean, right 1,873 1,864 2,190 1,772 1,422 Fat, right 1,105 289 837 98 05 Bone right 3,941 2,665 3,398 2,986 2,656 Tail, total 882 730 857 778 700 553 Lean total 312 292 362 328 218 Fat, total 132 24 48 40 24 Bone total 354 354 370 332 290 ^Weighed with round. Studies In Animal Nutrition — II 57 Table 10. — Slaughter House Weights of Carcass Parts (in Grams). Steer 513 502 509 501 512 500 Age 3 yr. 8 mo. 15 da. 3 yr. 8 mo. 19 da. 3 yr. 8 mo. 22 da. 3yr. 11 mo. 3yr. 11 mo. 21 da. 3yr. 11 mo. 26 da. Group 1 2 3 1 2 3 Head, total 13,287 12,902 10,642 14,702 13,233 11,824 Lean, total 3,690 3,386 2,556 4,004 3,266 2,930 Fat, total 1,294 438 264 606 706 380 Bone, total 7,865 9,078 7,822 9,962 9,139 8,514 Shin, right 8,748 7,032 6,523 9,018 7,437 7,355 Lean, right 4,639 3,719 3,769 4,670 3,928 4,199 Fat, right 1,039 355 238 1,223 461 316 Bone, right 3,060 2,970 2,523 3,085 3,037 2,805 Neck, right 1,944 1,585 1,635 1,841 1,600 1,660 Lean, right 987 944 925 833 800 874 Fat, right 616 188 171 558 175 198 Bone, right 357 462 537 447 620 589 Chuck, right 61,419 41,503 35,550 61,734 43,567 35,199 Lean, right 36,491 29,998 26,143 35,518 30,271 26,178 Fat, right 17,376 4,242 2,849 18,586 5,510 2,209 Bone, right 7,406 7,084 6,304 7,442 7,648 6,636 Plate, right 46,181 16,244 14,923 51,977 23,006 16,798 Lean right 18,269 10,056 9,499 18,093 11,581 10,651 Fat, right 24,127 3,412 2,896 30,290 7,474 3,052 Bone, right 3,623 2,713 2,512 3,487 3,827 3,094 Rib, right 27,830 13,821 11,725 27,783 14,527 10,321 Lean right 12,372 9,128 8,180 10,417 8,454 6,801 Fat, right 11,804 1,669 989 14,161 2,699 902 Bone, right 3,548 2,960 2,493 3,194 3,469 2,596 Loin, right 52,503 26,154 22,766 63,056 28,051 22,159 Lean, right 22,255 17,552 15,418 22,998 16,031 14,846 Fat, right 24,964 4,572 3,785 35,679 7,654 3,415 Bone, right 4,268 3,933 3,436 4,307 4,374 3,886 Kidney, Fat, right 7,245 1,458 788 9,772 2,370 1,216 Flank, right 15,560 4,390 3,558 18,768 5,502 4,586 £ Lean, right 4,237 2,331 2,117 4,544 1,847 2,805 Fat, right 11,254 1,998 1,383 14,146 3,571 1,697 ? Bone, right 96 82 50 47 67 81 Rump, right 11,288 5,243 5,142 13,018 6,870 5,041 Lean, right 3,609 2,998 2,631 3,816 3,054 2,471 i Fat, right 5,932 1,052 1,054 7,297 2,188 1,058 Bone, right 1,732 1,181 1,430 1,841 1,632 1,494 Round, right 38,655 28,023 25,862 39,970 30,725 26,073 Lean, right 25,391 22,213 20,188 25,065 21,704 19,949 Fat, right 9,554 2,310 2,553 11,142 4,970 2,468 Bone, right 3,522 3,352 3,010 3,632 3,913 3,559 Shank, right 6,245 5,077 4,639 6,381 5,071 4,657 Lean, right 1,813 1,815 1,708 1,837 1,717 1,548 Fat, right 1,408 293 176 980 247 185 Bone, right 3,029 2,989 2,749 3,564 3,078 2,875 Tail, total 675 893 774 854 875 842 Lean, total 304 384 324 496 366 406 Fat, total 102 42 68 118 74 68 Bone, total 297 441 386 304 416 386 58 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 11. — Distribution of Carcass and Offal Parts. Steer 556 554 555 557 552 548 Age 3 mo. 3 mo. 3 mo. 5 mo. 17 da. 5 mo. 7 da. 5 mo. 9 da. Group 1 2 3 1 2 3 Live weight 111,489 87,456 84,725 204,934 116,491 99,255 Warm empty weight 98,133 78,071 71,078 172,797 99,349 85,988 Percent empty weight to live weight 88.020 89.269 83.893 84.318 85.285 86.633 Percent carcass to live weight 54.188 57.319 53.626 53.710 52.998 54.388 Percent carcass to empty weight 61.563 64.210 63.923 63.700 62.143 62.780 Percent carcass + offal fat to live weight. . . . 55.446 58.055 54.172 57.008 54.530 54.232 Percent carcass + offal fat to empty weight. . 62.992 65.034 64.573 67.610 63.938 63.762 Percent offal fat to empty weight 1.429 0.825 0.650 3.910 1.796 0.983 Percent hide and hair to empty weight 10.510 9.479 9.257 8.160 10.601 9.720 Percent blood to empty weight 6.241 5.353 6.372 5.181 5.253 5.353 Percent heart market to empty weight 0.531 0.429 0.443 0.537 0.565 0.497 Percent lungs and trachea to empty weight. . 1.295 1.162 1.318 1.141 1.103 1.261 Percent brain and spinal cord to empty weight 0.406 0.506 0.497 0.319 0.469 0.612 Percent stomach to empty weight 2.013 1.547 1.882 2.512 1.951 2.192 Percent intestines to empty weight 2.745 2.660 3.437 2.370 2.802 2.932 Cm. intestines per kilo empty weight 34.83 37.17 34.61 22.03 31.41 39.03 Percent of liver to empty weight 1.793 1.494 1.745 1.838 1.189 1.284 Percent gall bladder and gall to empty weight 0.025 0.029 0.011 0.086 0.055 0.040 Percent of kidneys to empty weight 0.344 0.679 0.618 0.376 0.318 0.411 Percent spleen to empty weight 0.306 0.259 0.235 0.281 0.286 0.250 Percent pancreas to empty weight 0.098 0.097 0.149 0.130 0.102 0.099 Table 12. — Distribution of Carcass and Offal Parts. Steer 547 550 558 541 538 540 Age 8 mo. 5 da. 8 mo. 14 da. 8 mo. 12 da. 10 mo. 22 da. 10 mo. 26 da. 11 mo. 2 da. Group 1 2 3 1 2 3 Live weight 206,175 147,202 108,191 323,836 180,930 158,131 Warm empty weight 171,448 121,112 89,999 288,297 158,911 137,726 Percent empty weight to live weight 83.157 82.277 83.185 89.025 87.830 87.096 Percent carcass to live weight 54.167 50.459 51.085 58.594 53.658 55.226 Percent carcass to empty weight 65.138 61.328 61.411 65.817 61.093 63.408 Percent carcass + offal fat to live weight. . . . 56.048 52.232 51.851 61.786 55.566 56.685 Percent carcass + offal fat to empty weight. . 67.401 63.483 62.332 69.403 63.266 65.083 Percent offal fat to empty weight 2.262 2.155 0.921 3.586 2.172 1.675 Percent hide and hair to empty weight 8.526 8.620 9.042 9.219 9.654 9.435 Percent blood to empty weight 5.081 5.846 5.185 4.325 4.543 5.059 Percent heart market to empty weight 0.450 0.459 0.562 0.377 0.549 0.489 Percent lungs and trachea to empty weight . . 1.090 1.205 1.234 0.828 1.148 1.090 Percent brain and spinal cord to empty weight 0.268 0.366 0.578 0.197 0.306 0.372 Percent stomach to empty weight 3.027 3.165 2.833 2.732 2.845 2.827 Percent intestines to empty weight 2.610 2.992 2.908 1.819 2.294 2.366 Cm. intestines per kilo empty weight 20.78 27.87 34.43 14.08 25.19 23.83 Percent of liver to empty weight 1.663 1.645 1.502 1.329 1.245 1.157 Percent gall bladder and gall to empty weight 0.080 0.097 0.031 0.095 0.096 0.060 Percent of kidneys to empty weight 0.262 0.313 0.353 0.223 0.306 0.264 Percent spleen to empty weight 0.228 0.240 0.214 0.207 0.208 0.240 Percent pancreas to empty weight 0.117 0.165 0.170 0.135 0.131 0.131 Studies In Animal Nutrition — II 59 Table 13. — Distribution of Carcass and Offal Parts. Steer 505 503 532 531 504 523 525 10 mo. 11 mo. lyr. lyr. lyr. 2yr. 2 yr. 18 da. 11 da. 5 mo. 6 mo. 8 mo. 2 mo. 2 mo. 20 da. 12 da. 26 da. 6 da. 8 da. Group 1 2 1 3 1 2 3 Live weight 313,317 270,566 517,093 212,466 526,164 380,880 305,112 Warm empty weight 274,357 236,429 459,025 192,005 475,854 337,803 265,587 Percent empty weight to live weight 87.565 87.383 88.770 90.370 90.438 88.690 87.046 Percent carcass to live weight 56.445 54.048 60.459 56.015 58.235 57.708 54.265 Percent carcass to empty weight 64.460 61.852 68.107 61.984 64.391 65.067 62.341 Percent carcass +, offal fat to live weight 60.524 56.778 65.042 57.379 63.006 59.786 55.891 Percent carcass + offal fat to empty weight 69.119 64.976 73.270 63.494 69.667 67.410 64.209 Percent offal fat to empty weight 4.659 3.124 5.162 1.510 5.276 2.343 1.868 Percent hide and hair to empty weight 8.393 9.779 7.404 8.694 8.646 9.798 10.472 Percent blood to empty weight 5.034 5.523 4.085 4.925 4.414 4.525 5.126 Percent heart market to empty weight 0.385 0.529 0.448 0.527 0.343 0.437 0.440 Percent lungs and trachea to empty weight 0.910 1.078 0.843 0.997 0.762 0.998 0.624 Percent brain and spinal co^d to empty weight. . 0.196 0.282 0.140 • 0.298 0.152 0.236 0.268 Percent stomach to empty weight 3.214 2.439 2.373 2.783 2.694 2.766 3.332 Percent intestines to empty weight 1.768 2.003 1.291 1.921 2.002 1.547 2.244 Cm. intestines per kilo empty weight 16.26 20.93 11.69 19.21 10.88 13.80 Percent of liver to empty weight 1.452 1.542 1.240 1.148 0.999 0.967 1.083 Percent gall bladder and gall to empty weight. . . 0.078 0.085 0.062 0.059 0.146 0.043 0.047 Percent of kidneys to empty weight 0.262 0.277 0.189 0.264 0.184 0.208 0.250 Percent spleen to empty weight 0.218 0.280 0.193 0.251 0.277 0.251 0.209 Percent pancreas to empty weight 0.109 0.122 0.131 0.155 0.062 0.089 0.057 Table 14. — Distribution of Carcass and Offal Parts. Steer 515 507 529 527 526 524 Age 2 yr. 9 mo. 19 da. 2 yr. 9 mo. 16 da. 3 yr. 2 mo. 21 da. 3 yr. 3 mo. 15 da. 3 yr. 4 mo. 3 yr. 4 mo. 13 da. Group 1 2 1 1 2 3 Live weight 743,361 457,155 690,704 842,841 479,846 362,260 Warm empty weight 671,917 418.896 637,507 786,005 427,995 322,234 Percent empty weight to live weight 90.389 91.631 92.298 93.257 89.194 88.951 Percent carcass to live weight 61.489 60.490 65.687 70.423 61.176 58.075 Percent carcass to empty weight 68.028 66.015 71.169 75.515 68.587 65.289 Percent carcass + offal fat to live weight. . . . 65.509 62.964 71.342 76.179 63.583 59.458 Percent carcass + offal fat to empty weight. . 72.474 68.714 77.295 81.688 71.286 66.843 Percent offal fat to empty weight 4.447 2.699 6.126 6.173 2.699 1.554 Percent hide and hair to empty weight 7.433 8.229 7.148 5.883 8.349 9.339 Peroent blood to empty weight 4.161 4.850 3.612 3.484 4.429 5.282 Percent heart market to empty weight 0.396 0.492 0.321 0.413 0.391 0.492 Percent lungs and trachea to empty weight . 0.544 0.900 0.670 0.550 0.887 1.072 Percent brain and spinal cord to empty weight 0.108 0.178 0.110 0.089 0.154 0.235 Percent stomach to empty weight 1.960 2.297 1.898 1.316 2.567 2.512 Percent intestines to empty weight 1.015 1.343 0.830 0.642 0.966 1.240 Cm. intestines per kilo empty weight 6.99 9.46 6.05 6.28 10.37 11 35 Percent of liver to empty weight 0.890 0.904 0.843 0.728 0.825 0.937 Percent gall bladder and gall to empty weight 0.057 0.031 0.044 0.019 0.054 0.092 Percent of kidneys to empty weight 0.159 0.180 0.158 0.158 0.215 0.238 Percent spleen to empty weight 0.221 0.270 0.165 0.156 0.194 0.235 Percent pancreas to empty weight 0.063 0.055 0.129 0.108 0.116 0.135 60 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 15. — Distribution of Carcass and Offal Parts. Steer 513 502 509 501 512 500 Age 3 yr. 8 mo. 15 da. 3 yr. 8 mo. 19 da. 3 yr. 8 mo. 22 da. 3yr. 11 mo. 3yr. 11 mo. 21 da. 3yr. 11 mo. 26 da. Group 1 2 3 1 2 3 Live weight 853,007 506,878 439,814 883,480 548,050 457,786 Warm empty weight 771,142 444,424 391,461 814,914 493,877 407,833 Percent empty weight to live weight 90.403 87.679 89.006 92.239 90.115 89.088 Percent carcass to live weight 65.317 60.284 60.342 68.953 61.796 59.358 Percent carcass to empty weight 72.252 68.756 67.795 74.755 68.575 66.628 Percent carcass + offal fat to live weight. . . 71.621 62.528 62.598 73.325 64.981 62.185 Percent carcass + offal fat to empty weight. 79.224 71.315 70.330 79.494 72.109 69.801 Percent offal fat to empty weight 6.973 2.560 2.535 4.740 3.534 3.173 Percent hide and hair to empty weight 5.873 8.901 9.609 6.147 8.356 8.812 Percent blood to empty weight 3.330 4.439 4.672 3.523 4.895 5.215 Percent heart market to empty weight 0.300 0.438 0.424 0.272 0.396 0.360 Percent lungs and trachea to empty weight. . 0.630 0.832 0.839 0.471 0.786 0.919 Percent brain and spinal cord to empty weight 0.097 0.180 0.189 0.093 0.135 0.204 Percent stomach to empty weight 1.744 2.438 2.516 1.741 2.245 2.696 Percent intestines to empty weight 0.761 0.934 0.994 0.598 1.078 1.112 Cm. intestines per kilo empty weight 7.18 10.16 10.62 5.95 11.23 10.62 Percent of liver to empty weight 0.768 0.836 0.990 0.756 0.894 1.136 Percent gall bladder and gall to empty weight 0.016 0.075 0.057 0.033 0.061 0.074 Percent of kidneys to empty weight 0.132 0.189 0.198 0.127 0.217 0.250 Percent spleen to empty weight 0.144 0.207 0.333 0.145 0.254 0.258 Percent pancreas to empty weight 0.113 0.131 0.144 0.103 0.149 0.153 Table 16. — Main Divisions of Empty Animal and Loss on Cooling. Steer 556 554 555 557 552 548 Age 3 mo. 3 mo. 3 mo. 5 mo. 17 da. 5 mo. 7 da. 5 mo. 9 da. Group 1 2 3 1 2 3 Weights of parts in grams Warm empty weight 98,133 78,071 71,078 172,797 99,349 85,988 Carcass 60,414 50,129 45,435 110,071 61,738 53,983 Offal fat 1,402 644 462 6,757 1,784 845 Hide and hair 10,314 7,400 6,580 14,100 10,532 8,358 Head, tail, feet, etc 6,813 6,203 5,653 10,591 8,081 7,463 Blood 6,124 4,197 4,529 8,952 5,219 4,603 Organs 11,488 8,419 8,489 19,831 10,701 9,645 Loss on cooling 1,422 300 23 724 716 1,658 Percent of parts to warm empty weight Carcass 61.563 64.210 63.923 63.700 62.143 62.780 Offal fat 1.429 0.825 0.650 3.910 1.796 0.983 Hide and hair 10.510 9.479 9.257 8.160 10.601 9.720 Head, tail, feet, etc 6.943 7.945 7.953 6.129 8.134 8.679 Blood 6.241 5.353 6.372 5.181 5.253 5.353 Organs 11.707 10.784 11.943 11.476 10.771 11.217 Loss on cooling 1.449 0.384 0.032 0.419 0.721 1.928 Studies In Animal Nutrition — II 61 Tables 17 , 18 , 19 . — Main Divisions of Empty Animal and Loss on ? Cooling. Steer 547 550 558 541 538 __ 540 Age 8 mo. 8 mo. 8 mo. 10 mo. 10 mo. 11 mo 5 da. 14 da. 12 da. 22 da. 26 da. 2 da. Group 1 2 3 1 2 3 Weights of i parts in grams Warm empty weight 171,448 121,112 89,999 288,297 158,911 137,726 Carcass 111,678 74,276 55,269 189,748 97,084 87,329 Offal fat 3,879 2,610 829 10,337 3,452 2,307 Hide and hair 14,618 10,440 8,138 26,576 15,342 12,994 Head, tail, feet, etc 11,802 8,969 8,289 13,781 9,921 9,005 Blood 8,711 7,080 4,666 12,470 7,219 6,967 Organs 19,822 15,084 11,128 28,319 17,879 15,662 Loss on cooling 72 1,377 929 8,054 4,648 5,941 Percent of parts to warm empt y weight Carcass 65.138 61.328 61.411 65.817 61.093 63.408 Offal fat 2.262 2.155 0.921 3.586 2.172 1.675 Hide and hair 8.526 8.620 9.042 9.219 9.654 9.435 Head, tail, feet, etc 6.884 7.406 9.210 4.780 6.243 6.538 Blood 5.081 5.846 5.185 4.325 4.543 5.059 Organs 11.562 12.455 12.365 9.823 11.251 11.372 Loss on cooling 0.042 1.137 1.032 2.794 2.925 4.314 Steer 505 503 532 531 504 523 525 Age 10 mo. 11 mo. 1 yr. 5 mo. 1 yr. 6 mo. 1 yr. 8 mo. 2 yr. 2 mo. 2 yr. 2 mo . 18 da. 11 da. 20 da. 12 da. 26 da. 6 da. 8 da. Group 1 2 1 3 1 2 3 Weights of parts in grams Warm empty weight 274,357 236,429 459,025 192,005 475,854 337,803 265,587 Carcass 176,851 146,236 312,629 119,012 306,409 219,798 165,570 Offal fat 12,781 7,385 23,697 2,899 25,105 7,915 4,961 Hide and hair 23,026 23,120 33,988 16,693 41,144 33,097 27,813 Head, tail, feet., etc 13,377 15,218 20,120 11,718 21,024 23,460 16,701 Blood 13,810 13,058 18,752 9,457 21,005 15,287 13,614 Organs 28,033 25,676 39,809 20,313 45,006 32,570 26,676 Loss on cooling 5,398 3,164 10,030 12,701 7,208 5,676 10,252 Percent of parts to warm empt y weight Carcass 64.460 61.852 68.107 61.984 64.391 65.067 62.341 Offal fat 4.659 3.124 5.162 1.510 5.276 2.343 1.868 Hide and hair 8.393 9.779 7.404 8.694 8.646 9.798 10.472 Head, tail, feet, etc 4.876 6.437 4.383 6.103 4.418 6.945 6.288 Blood 5.034 5.523 4.085 4.925 4.414 4.525 5.126 Organs 10.218 10.860 8.673 10.579 9.458 9.642 10.044 Loss on cooling 1.968 1.338 2.185 6.615 1.515 1.680 3.860 Steer 515 507 529 527 526 524 Age 2 yr. 9 mo. 2 yr. 9 mo. 3 yr. 2 mo. 3 yr. 3 mo. 3 yr. 4 mo. 3 yr. 4 ma 19 da. 16 da. 21 da. 15 da. 13 da. Group 1 2 1 1 2 3 Weights of parts in grams Warm empty weight 671,917 418,896 637,507 786,005 427,995 322,234 Carcass 457,088 276,534 453,705 593,554 293,550 210,384 Offal fat 29,877 11,307 39,056 48,517 11,551 5,007 Hide and hair 49,943 34,473 45,567 46,240 35,732 30,092 Head, tail, feet, etc 28,764 23,193 26,124 25,599 22,957 21,467 Blood 27,856 20,316 23,028 27,382 18,957 17,019 Organs 51,173 36,207 43,425 47,812 35,360 30,837 Loss on cooling 27,216 16,866 12,031 12,363 9,888 7,428 Percent of parts to warm empty Carcass Offal fat Hide and hair Head, tail, feet, etc Blood Organs 68.028 4.447 7.433 4.281 4.161 7.616 66.015 2.699 8.229 5.537 4.850 8.643 4 n9« weight 71.169 6.126 7.148 4.098 3.612 6.812 1 75.515 6.173 5.883 3.257 3.484 6.083 1 K73 68.587 2.699 8.349 5.364 4.429 8.262 o Qin 65.289 1.554 9.339 6.662 5.282 9.570 62 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 20 . — Main Divisions of Empty Animal and Loss on Cooling. Steer 513 502 509 501 512 500 Age 3 yr. 8 mo. 15 da. 3 yr. 8 mo. 19 da. 3 yr. 8 mo. 22 da. 3 yr.ll mo. 3 yr.ll mo. 21 da. 3 yr.ll mo. 26 da. Group 1 2 3 1 2 3 Weights of parts in grams Warm empty weight 771,142 444,424 391,461 814,914 493,877 407,833 Carcass 557,162 305,566 265,391 609,185 338,675 271,732 Offal fat 53,771 11,377 9,922 38,625 17,454 12,940 Hide and hair 45,286 39,556 37,614 50,090 41,268 35,938 Head, tail, feet, etc 27,336 26,278 20,423 30.523 25,833 22,814 Blood 25,680 19,728 18,291 28,710 24,176 21,269 Organs 48,968 36,679 33,938 47,332 40,410 36,998 Loss on cooling 13,067 3,968 4,097 11,976 5,864 6,624 Percent of parts to warm empty weight Carcass 72.252 68.756 67.795 74.755 68.575 66.628 Offal fat 6.973 2.560 2.535 4.740 3.534 3.173 Hide and hair 5.873 8.901 9.609 6.147 8.356 8.812 Head tail feet etc 3.545 5.913 5.217 3.746 5.231 5.594 5.215 Blood 3.330 4.439 4.672 3.523 4.895 Organs 6.350 8.253 8.670 5.808 8.182 9.072 1.624 Loss on cooling 1.694 0.893 1.047 1.470 1.187 Table 21 . — Proportion of Cuts to Empty Weight and to Carcass. Steer 556 554 555 557 552 548 Age 3 mo. 3 mo. 3 mo. 5 mo. 17 da. 5 mo. 7 da. 5 mo. 9 da. Group 1 2 3 1 2 3 Warm empty weight 98,133 78,071 71,078 172,797 99,349 85,988 Weight of carcass 60,570 50,908 45,342 111,842 62,316 53,416 Percent forequarters to empty weight 31.459 33.549 33.977 32.967 32.256 32.432 Percent forequarters to carcass 50.969 51.450 53.262 50.934 51.425 52.209 Percent hindquarters to empty weight 30.263 31.658 29.815 31.757 30.468 29.688 Percent hindquarters to carcass 49.031 48.550 46.738 49.065 48.575 47.791 Percent shins to empty weight 3.597 3.761 4.606 2.853 3.422 3.703 Percent shins to carcass 5.828 5.767 7.221 4.408 5.456 5.961 Percent of neck to empty weight 1.054 1.061 1.413 0.890 0.862 1.226 Percent of neck to carcass 1.707 1.626 1.214 1.375 1.374 1.973 Percent chucks to empty weight 15.489 17.277 16.497 15.148 15.805 15.970 Percent chucks to carcass 25.095 26.489 25.861 23.404 25.197 25.708 Percent of plates to empty weight 6.212 5.751 5.929 7.800 5.919 5.873 Percent of plates to carcass 10.064 8.820 9.294 12.051 9.436 9.454 Percent ribs to empty weight 5.065 5.492 5.388 6.107 6.281 5.610 Percent ribs to carcass 8.205 8.423 8.447 9.435 10.013 9.031 Percent of loin to to empty weight 10.249 10.473 8.728 10.842 10.361 8.987 Percent loin to carcass 16.606 16.060 13.683 16.750 16.519 14.468 Percent kidney fat + kidney to empty weight 0.772 0.986 0.801 2.244 0.821 0.741 Percent kidney fat + kidney to carcass 1.251 1.512 1.255 3.466 1.309 1.193 Percent flanks to empty weight 1.769 1.365 1.432 2.510 1.882 1.465 Percent flanks to carcass 2.866 2.094 2.245 3.879 3.001 2.359 Percent rumps to empty weight 1.887 2.283 1.978 2.059 1.987 1.886 Percent rumps to carcass 3.058 3.500 3.101 3.181 3.168 3.037 Percent rounds to empty weight 12.444 12.973 12.944 11.678 12.542 13.516 Percent rounds to carcass 20.162 19.893 20.290 18.043 19.995 21.758 Percent shanks to empty weight 3.012 3.574 3.683 2.390 2.877 3.035 Percent shanks to carcass 4.880 5.480 5.774 3.691 4.586 4.886 Percent head to empty weight 3.402 3.708 3.986 3.249 4.086 4.690 Percent tail to empty weight 0.175 0.256 0.172 0.192 0.239 0.171 Studies In Animal Nutrition — II 63 Table 22. — Proportion of Cuts to Empty Weight and to Carcass. Steer 547 550 558 541 538 540 Age 8 mo. 5 da. 8 mo. 14 da. 8 mo. 12 da. 10 mo. 22 da. 10 mo. 26 da. 11 mo. 2 da. Group 1 2 3 1 2 3 Warm empty weight 171,448 121,112 89,999 288,297 158,911 137,726 Weight of carcass 112,544 75,552 56.020 188,760 100,450 84,850 Percent forequarters to empty weight 32.326 31.696 31.309 32.263 31.969 31.374 Percent forequarters to carcass 49.245 50.810 50.300 49.276 50.574 50.925 Percent hindquarters to empty weight 33.317 30.686 30.963 33.211 31.243 30.234 Percent hindquarters to carcass 50.755 49.190 49.700 50.724 49.426 49.075 Percent shins to empty weight 2.901 3.222 3.789 2.575 3.021 3.235 Percent shins to carcass 4.420 5.165 6.087 3.933 4.778 5.252 Percent of neck to empty weight 0.612 0.745 0.762 1.041 1.077 1.001 Percent of neck to carcass .933 1.194 1.225 1.589 1.704 1.624 Percent chucks to empty weight 16.217 16.451 16.794 15.805 16.407 15.630 Percent chucks to carcass 24.705 26.371 26.980 24.139 25.955 25.369 Percent of plates to empty weight 6.771 6.184 4.567 7.052 6.444 6.352 Percent of plates to carcass 10.314 9.914 7.337 10.770 10.194 10.310 Percent ribs to empty weight 5.753 5.094 5.289 5.987 4.891 4.964 Percent ribs to carcass 8.766 8.167 8.497 9.144 7.737 8.057 Percent of loin to empty weight 12.100 11.295 9.687 12.371 11.273 11.331 Percent loin to carcass 18.434 18.107 15.562 18.895 17.834 18.392 Percent kidney fat + kidney to empty weight 1.213 0.937 0.604 2.324 0.698 0.759 Percent kidney fat + kidney to carcass 1.848 1.502 0.971 3.550 1.104 1.232 Percent flanks to empty weight 2.703 1.818 1.278 2.584 1.805 1.413 Percent flanks to carcass 4.118 2.915 2.053 3.947 2.855 2.293 Percent rumps to empty weight 1.786 1.949 1.924 1.918 1.807 1.888 Percent rumps to carcass 2.721 3.124 3.092 2.930 2.859 3.064 Percent rounds to empty weight 12.681 12.205 13.829 11.838 12.956 12.060 Percent rounds to carcass 19.319 19.565 22.217 18.081 20.496 19.576 Percent shanks to empty weight 2.666 2.480 3.416 2.110 2.651 2.628 Percent shanks to carcass 4.061 3.976 5.487 3.223 4.193 4.266 Percent head to empty weight 3.768 4.218 5.293 2.407 3.260 3.447 Percent tail to empty weight 0.170 0.154 0.109 0.132 0.144 0.158 Table 23. — Proportion of Cuts to Empty Weight and to Carcass. Steer 505 503 532 531 504 523 525 10 mo. 18 da. 11 mo. 11 da. 1 yr. 5 mo. 20 da. 1 yr. 6 mo. 12 da. 1 yr. 8 mo. 26 da. 2 yr. 2 mo. ' 6 da. 2 yr. 2 mo. 8 da. Group 1 2 1 3 1 2 3 Warm empty weight 274,357 236,429 459,025 192,005 475,854 337,803 265,587 Weight of carcass 177,932 148,805 312,808 118,224 315,362 219,798 165,570 Percent forequarters to empty weight 33.049 31.680 34.684 30.919 32.187 33.223 32.535 Percent foreauarters to carcass 50.959 50.334 50.897 50.215 48.566 51.060 52.188 Percent hindquarters to empty weight 31.805 31.260 33.462 30.654 34.087 31.844 29.806 Percent hindquarters to carcass 49.041 49.688 49.103 49.785 51.434 48.940 47.812 Percent shins to empty weight 2.699 2.817 2.938 3.236 2.574 i 2 . 925 3.259 Percent shins to carcass 4.162 4.476 4.311 5.256 3.883 4.496 5.228 Percent of neck to empty weight 0.761 0.980 0.643 1.346 0.646 0.824 1.169 Percent of neck to carcass 1.173 1.558 0.944 2.186 0.974 M.266 1.875 Percent chucks to empty weight 16.006 15.921 16.682 15.940 14.561 17.109 15.532 Percent chucks to carcass 24.680 25.296 24.480 25.888 21.972 26.295 24.915 Percent of plates to empty weight 7.248 6.412 8.051 5.711 7.946 7.102 6.443 Percent of plates to carcass 11.176 10.188 11.815 9.276 11.990 10.915 10.335 Percent ribs to empty weight 6.274 5.604 6.310 4.670 6.424 { 5.182 i 6. 125 Percent ribs to carcass 9.674 8.904 9.260 7.584 9.694 17.965 9.825 Percent of loin to empty weight 11.388 11.399 11.924 10.042 12.239 11.156 9.971 Percent loin to carcass 17.560 18.111 17.497 16.310 18.468 17.147 15.994 Percent kidney fat + kidney to empty weight. . 2.360 1.176 2.745 0.642 2.580 1.129 0.724 Percent kidney fat -|- kidney to carcass 3.637 1.869 4.029 1.042 3.893 1.735 1.161 Percent flanks to empty weight 2.672 1.971 2.916 1.305 3.480 2.247 1.956 Percent flanks to carcass 4.120 3 . 132 4.279 2.120 5.251 3.453 3.138 Percent rumps to empty weight 2.031 1.815 2.261 1.857 2.787 2.147 2.166 Percent rumps to carcass 3.132 2.884 3.318 3.016 4.205 3.300 3.474 Percent rounds to empty weight 11.212 12.350 10 895 13.905 10.926 12.796 12.684 Percent rounds to carcass 17.288 19.623 15.988 22.583 16.486 19.666 20.345 Percent ehanks to empty weight 2.103 2.574 2 040 2.767 2.025 2.294 2.277 Percent shanks to carcass 3.243 4.090 2.994 4.493 3.055 3.525 3.653 Percent head to empty weight 2.543 3.519 2 290 3.256 2.319 2.915 3.214 Percent tail to empty weight 0.190 0.163 0.147 0.146 0.145 0.186 0.176 64 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 24. — Proportion of Cuts to Empty Weight and to Carcass. Steer 515 507 529 527 526 524 2 yr. 9 mo. 2 yr. 9 mo. 3 yr. 2 mo. 3 yr. 3 mo. 3 yr. 4 mo. 3 yr. 4 mo. Age 19 da. 16 da. 21 da. 15 da. 13 da. Group 1 2 1 1 2 3 Warm empty weight 671,917 418,896 637,507 786,005 427,995 322,234 Wt. of carcass 457,088 276,534 448,276 578,092 293,550 210,384 Percent forequarters to empty weight 34.470 34.650 34.959 37.576 35.476 34.011 Percent forequarters to carcass 50.671 52.488 49.717 51.090 51.723 52.093 Percent hindquarters to empty weight 33.557 31.365 35.358 35.972 33.112 31.278 Percent hindquarters to carcass 49.329 47.512 50.283 48.910 48.277 47.907 Percent shins to empty weight 2.491 3.014 2.596 2.245 3.172 3.310 Percent shins to carcass 3.662 4.566 3.692 3.053 4.625 5.070 Percent of neck to empty weight 0.681 0.769 0.498 0.466 0.847 0.914 Percent of neck to carcass 1.001 1.165 0.708 0.634 1.235 1.400 Percent chucks to empty weight 14.675 17.408 14.856 15.776 16.779 17.525 Percent chucks to carcass 21.572 26.369 21.128 21.450 24.463 26.842 Percent of plates to empty weight 10.381 7.927 11.263 11.862 8.459 6.508 Percent of plates to carcass 15.260 12.008 16.017 16.128 12.334 9.968 Percent ribs to empty weight 6.253 5.555 5.746 7.227 6.248 5.720 Percent of loin to empty weight 9.192 8.415 8.172 9.826 9.110 8.761 Percent ribs to carcass 13.050 11.160 13.575 14.043 11.687 10.338 Percent loin to carcass 19.184 16.905 19.306 19.093 17.040 15.834 Percent kidney fat + kidney to empty weight 1.635 1.224 1.762 2.571 0.969 1.762 Percent kidney fat + kidney to carcass 2.404 1.854 2.505 3.496 1.412 2.699 Percent flanks to empty weight 3.605 2.243 4.240 4.114 2.305 1.494 Percent flanks to carcass 5.300 3.397 6.029 5.593 3.361 2.288 Percent rumps to empty weight 2.978 2.454 2.931 3.491 2.686 2.006 Percent rumps to carcass 4.378 3.717 4.169 4.747 3.916 3.072 Percent rounds to empty weight 10.171 12.067 12.850 10.113 13.132 14.283 Percent rounds to carcass 14.952 18.279 18.274 13.751 19.146 21.877 Percent shanks to empty weight 2 060 2 323 1 637 2.273 2.608 Percent shanks to carcass 3.029 3.519 2.226 3.315 3.995 Percent head to empty weight 2.213 2.835 2.078 1.633 2.720 3.387 Percent tail to empty weight 0.131 0.174 0.134 0.099 0.164 0.172 Table 25. — Proportion of Cuts to Empty Weight and to Carcass. Steer ' 513 502 509 501 512 500 Age 3 yr. 8 mo. 15 da. 3 yr. 8 mo. 19 da. 3 yr. 8 mo. 22 da. 3yr. 11 mo. 3yr. 11 mo. 21 da. 3yr. 11 mo. 26 da. Group 1 2 3 1 2 3 Warm empty weight 771,142 444,424 391,461 814,914 493,877 407,833 Weight of carcass 557,034 306,838 267,176 607,658 338,872 371,250 Percent forequarters to empty weight 37.976 36.143 35.971 37.391 36.531 34.963 Percent forequarters to carcass 52.573 52.349 52.704 50.144 53.240 52.568 Percent hindquarters to empty weight 34.259 32.899 32.280 37.176 32.084 31.547 Percent hindquarters to carcass 47.427 47.651 47.296 49.856 46.760 47.432 Percent shins to empty weight 2.269 3.165 3.333 2.213 3.012 3.607 Percent shins to carcass 3.141 4.584 4.883 2.968 4.389 5.423 Percent of neck to empty weight 0.504 0.713 0.835 0.452 0.648 0.814 Percent of neck to carcass 0.698 1.033 1.224 0.606 0.944 1.224 Percent chucks to empty weight 15.929 18.677 18.163 15.151 17.643 17.261 Percent chucks to carcass 22.052 27.052 26.612 20.319 25.713 25.953 Percent of plates to empty weight 11.977 7.310 7.624 12.756 9.316 8.238 Percent of plates to carcass 16.581 10.588 11.171 17.107 13.578 12.386 Percent ribs to empty weight 7.218 6.220 5.990 6.819 5.883 5.061 Percent ribs to carcass 9.992 9.009 8.777 9.144 8.574 7.610 Percent of loin to empty weight 13.617 11.770 11.631 15.476 11.360 10.867 Percent loin to carcass 18.851 17.047 17.042 20.754 16.556 16.338 Percent kidney fat + kidney to empty weight 2.011 0.845 0.600 2.526 1.177 0.846 Percent kidney fat + kidney to carcass 2.783 1.223 0.880 3.387 1.716 1.272 Percent flanks to empty weight 4.036 1.976 1.818 4.606 2.228 2.249 Percent flanks to carcass 5.587 2.861 2.663 6.177 3.247 3.381 Percent rumps to empty weight 2.928 2.359 2.627 3.195 2.782 2.472 Percent rumps to carcass 4.053 3.417 3.849 4.285 4.055 3.717 Percent rounds to empty weight 10.025 12.611 13.213 9.810 12.442 12.786 Percent rounds to carcass 13.879 18.266 19.360 13.139 18.134 19.224 Percent shanks to empty weight 1.620 2.285 2.370 1.566 2.054 2.284 Percent shanks to carcass 2.242 3.309 3.473 2.100 2.993 3.434 Percent head to empty weight 1.723 2.903 2.719 1.804 2.679 2.899 Percent tail to empty weight 0.087 0.201 0.198 0.105 0.177 0.206 Studies In Animal Nutrition — II 65 Table 26. — Distribution of Lean, Fat and Bone. Steer 556 554 555 557 552 548 Age 3 mo. 3 mo. 3 mo. 5 mo. 17 da. 5 mo. 7 da. 5 mo. 9 da. Group 1 2 3 1 2 3 Proportion of lean, fat and bone in empty animal Percent skeleton 21.148 24.090 23.364 17.020 21.443 22.973 Percent lean flesh 41.874 43.747 44.125 39.045 42.579 43.234 Percent fatty tissue (excl. offal fat) 4.168 3.440 2.334 13.634 5.552 3.114 Percent total fatty tissue 5.596 4.265 2.994 17.545 7.348 4.097 Percent offal and kidney fats to total fatty tissue 33.176 26.547 27.820 32.935 31.288 32.047 Proportion of iean, fat and bone in carcass Percent skeleton 25.527 27.595 27.343 19.392 24.485 26.535 Percent lean flesh 66.551 65.628 67.372 59.293 66.436 67.631 Percent fatty tissue 6.535 4.997 3.357 20.384 6.357 4.793 Percent kidney fat to fatty tissue in carcass. . 10.611 9.434 8.541 14.159 9.601 11.094 Table 27. — Distribution of Lean, Fat and Bone. Steer 547 550 558 541 538 540 Age 8 mo. 8 mo. 8 mo. 10 mo. 10 mo. 11 mo. 5 da. 14 da. 12 da. 22 da. 26 da. 2 da. Group 1 2 3 1 2 3 Proportion of lean, fat and bone in empty animal Percent skeleton 16.096 19.271 23.734 13.301 17.502 18.126 Percent lean flesh 44.646 42.361 42.627 42.917 43.496 42.316 Percent fatty tissue (excl. offal fat) 10.642 6.916 3.558 12.988 7.139 6.189 Percent total fattv tissue Percent offal and kidney fats to total fatty 12.905 9.071 4.479 16.574 9.311 7.864 tissue 24.899 30.639 26.172 34.309 27.534 27.597 Proportion of lean, fat and bone in carcass Percent skeleton 17.789 22.864 27.504 14.958 20.418 21.638 Percent lean flesh 65.640 65.534 65.973 64.728 67.616 67.595 Percent fatty tissue 15.485 10.573 5.159 19.565 10.999 9.636 Percent kidney fat to fatty tissue in carcass. . 9.353 9.464 7.820 16.399 5.630 8.341 Table 28. — Distribution of Lean, Fat and Bone. Steer 505 503 532 531 504 523 Age 10 mo. 11 mo. 1 yr. 5 mo. 1 yr. 6 mo. 1 yr. 8 mo. 2 yr. 2 mo. 18 da. 11 da. 20 da. 12 da. 26 da. 6 da. Group 1 2 1 3 1 2 Proportion of lean, fat and bone in empty animal Percent skeleton 13.758 17.393 13.557 17.218 12.082 15.161 Percent lean flesh 41.235 41.941 41.765 43.867 38.281 44.821 Percent fatty tissue (excl. offal fat) 13.662 8.861 16.111 5.158 18.905 9.008 Percent total fatty tissue 18.321 11.985 21.274 6.668 24.181 11.351 Percent offal and kidney fats to total fatty tissue 36.875 33.566 36.283 28.314 31.726 38.754 Proportion of lean, fat and bone in carcass Percent skeleton 15,978 20.207 15.544 20.686 18.696 17.332 Percent lean flesh 62 . 622 65.374 60.392 70.033 56.850 67.631 Percent fatty tissue 20.576 13 407 23.177 8.130 28.300 13.687 Percent kidney fat to fatty tissue in carcass. . 15.716 10.609 16.185 7.553 12.773 10.338 66 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 29. — Distribution of Lean, Fat and Bone. Steer 525 515 507 527 526 524 Age 2 yr. 2 mo. 8 da. 2 yr. 9 mo. 19 da. 2 yr. 9 mo. 16 da. 3 yr. 3 mo. 15 da. 3 yr. 4 mo. 3 yr. 4 mo. 13 da. Group 3 1 2 1 2 3 Proportion of lean, fat and bone in empty animal Percent skeleton 16.230 11.555 15.532 9.382 16.138 19.740 Percent lean flesh 44.504 33.109 44.151 34.603 44.777 46.354 Percent fatty tissue (excl. offal fat) 6.201 26.547 10.631 31.891 11.653 4.242 Percent total fatty tissue Percent offal and kidney fats to total fatty 8.069 30.994 13.331 38.063 14.352 5.795 tissue 29.021 19.111 26.085 22.555 24.054 30.913 Proportion of lean, fat and bone in carcass Percent skeleton 19.522 13.009 18.004 9.943 18.316 23.425 Percent lean flesh 69.778 47.947 65.918 46.403 64.347 69.762 Percent fatty tissue 9.625 38.452 15.607 43.137 16.696 6.134 Percent kidney fat to fatty tissue in carcass . . 7.894 5.645 10.139 7.605 6.578 5.936 Table 30. — Distribution of Lean, Fat and Bone. Steer 513 502 509 501 512 500 Age 3 yr. 8 mo. 15 da. 3 yr. 8 mo. 19 da. 3 yr. 8 mo. 22 da. 3yr. 11 mo. 3yr. 11 mo. 21 da. 3yr. 11 mo. 26 da. Group 1 2 3 1 2 3 Proportion of lean, fat and bor.e in empty animal Percent skeleton 9.991 16.302 16.570 9.891 16.285 17.509 Percent lean flesh 34.251 46.190 47.013 31.915 40 . 983 45.112 Percent fatty tissue (excl. offal fat) 30.090 9.806 8.710 35.389 15.271 8.307 Percent total fatty tissue 37.063 12.365 11.245 40.129 18.805 11.480 Percent offal and kidney fats to total fatty tissue 23.884 26.009 26.121 17.788 23.897 32.832 Proportion of lean, fat and bone in carcass Percent skeleton 11.001 18.072 18.747 10.218 18.688 20.361 Percent lean flesh 46.698 65.672 67.804 42.060 58.658 66.597 Percent fatty tissue 41.405 14.046 12.637 47.340 22.025 12.325 Percent kidney fat to fatty tissue in carcass. . 6.283 6.766 4.668 6.794 6.351 7.274 Table 31. — Distribution of Lean Flesh in the Animal. Steer 556 554 555 557 552 548 Age 3 mo. 3 mo. 3 mo. 5 mo. 5 mo. 5 mo. 17 da. 7 da. 9 da. Group 1 2 3 1 2 3 Weight of lean in animal 20,546 17,077 15,681 33,734 21,151 18,588 Percent of total lean in head 1.708 1.956 2.442 1.553 1.882 2.679 Percent of total lean in shin 4.088 3.777 5.287 3.436 4.085 3.954 Percent of total lean in neck 1.674 1.353 1.830 1.254 1.258 1.544 Percent of total lean in chuck 26.599 28.079 27.409 26.036 26.538 27.077 Percent of total lean in plate 10.201 8.637 9.317 10.998 8.922 8.904 Percent of total lean in rib 7.841 8.532 8.010 9.444 9.432 8.446 Percent of total lean in loin 16.S84 17.017 14.725 17.362 16.628 15.246 Percent of total lean in flank 2.915 2.249 2.889 3.311 2.823 2.394 Percent of total lean in rump 2.667 2.653 2.557 2.413 2.605 2.609 Percent of total lean in round 23.051 23.183 22.722 22.173 23.238 24.769 Percent of total lean in shank 2.176 2.342 2.659 1.862 2.340 2.233 Percent of total lean in tail 0.195 0.223 0.153 0.157 0.251 0.145 Studies In Animal Nutrition — II 67 Table 32. — Distribution of Lean Flesh in the Animal. Steer 547 550 558 541 538 540 Age 8 mo. 8 mo. 8 mo. 10 mo. 10 mo. 11 mo. 5 da. 14 da. 12 da. 22 da. 26 da. 2 da. Group 1 2 3 1 2 3 Weight of lean in animal 38.272 25,652 19,182 61,864 34,560 29,140 Percent of total lean in head 3.308 3.321 3.618 1.122 1.617 1.527 Percent of total lean in shin 3.601 3.863 4.056 3.115 3.464 4.005 Percent of total lean in neck 0.792 1.010 1.090 1.578 1.455 1.551 Percent of total lean in chuck 26.390 26.820 27.854 27.271 27.321 26.668 Percent of total lean in plate 9.338 9.450 7.226 10.159 9.251 9.602 Percent of total lean in rib 8.570 8.124 8.305 9.366 7.517 8.102 Percent of total lean in loin 17.736 17.683 15.541 18.925 18.420 18.360 Percent of total lean in flank 3.585 2.557 2.409 2.699 2.781 2.255 Percent of total lean in rump 1.934 2.460 2.414 2.227 2.257 2.622 Percent of total lean in round 22.330 22.844 25.190 21.822 23.617 23.089 Percent of total lean in shank 2.237 1.696 2.252 1.586 2.182 2.159 Percent of total lean in tail 0.180 0.172 0.047 0.129 0.119 0.062 Table 33. — Distribution of Lean Flesh in the Animal. Steer 505 503 532 531 504 523 m 525 Age 10 mo. 11 mo. 1 yr. 5 mo. 1 yr. 6 mo. 1 yr. 8 mo. 2 yr. 2 mo. 2 yr. 2 mo. 18 da. 11 da. 20 da. 12 da. 26 da. 6 da. 8 da. Group 1 2 1 3 1 2 3 Weight of lean in animal 56,566 49,580 95,857 42,113 91,081 75,704 59,099 Percent of total lean in head. . . 1.321 1.743 1.297 1.555 1.458 1.651 1.758 Percent of total lean in shin. . . 3.566 3.235 3.697 3.804 3.487 3.528 4.097 Percent of total lean in neck. . . 1.133 1.105 0.875 2.208 0.927 1.246 1.734 Percent of total lean in chuck. . 28.068 28.318 28.167 27.364 25.353 28.695 26.405 Percent of total lean in plate. . . 10.361 8.871 10.450 8.513 10.559 10.176 9.562 Percent of total lean in rib 9.988 9.008 9.053 7.449 10.159 7.947 9.870 Percent of total lean in loin 17.401 17.352 18.849 16.715 18.487 17.063 15.829 Percent of total lean in flank. . . 3.380 2.985 3.061 2.044 4.148 3.010 2.883 Percent of total lean in rump . . 2.298 2.041 2.681 2.377 3.166 2.376 2.689 Percent of total lean in round . . 20.493 23.193 19.855 25.522 20.442 22.390 23.286 Percent of total lean in shank. . 1.801 1.999 1.850 2.306 1.692 1.749 1.736 Percent of total lean in tail. . . . 0.189 0.151 0.166 0.143 0.122 0.169 0.151 Table 34. — Distribution of Lean Flesh in the Animal. Steer 515 507 529 527 526 524 Age 2 yr. 9 mo. 19 da. 2 yr. 9 mo. 16 da. 3 yr. 2 mo. 21 da. 3 yr. 3 mo. 15 da. 3 yr. 4 mo. 3 yr. 4 mo. 13 da. Group 1 2 1 1 2 3 Weight of lean in animal Percent of total lean in head Percent of total lean in shin Percent of total lean in neck Percent of total lean in chuck Percent of total lean in plate Percent of total lean in rib Percent of total lean in loin Percent of total lean in flank Percent of total lean in rump Percent of total lean in round Percent of total lean in shank Percent of total lean in tail 111,232 1.344 3.436 0.998 27.936 11.857 8.548 18.709 2.950 3.095 19.303 1.684 0.140 92,474 1.281 3.805 1.079 29.534 10.883 8.536 16.072 2.659 2.726 21.250 2.016 0.158 135,989 1.236 3.214 0.536 26.590 13.020 9.508 18.435 3.547 3.272 18.897 1.610 0.133 95,822 1.266 3.866 1.093 26.589 11.208 9.008 16.405 2.339 2.926 23.280 1.849 0.171 74,684 1.595 3.523 1.208 28.576 8.768 8.573 16.201 2.093 2.164 25.249 1.904 0.146 68 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 35. — Distribution of Lean Flesh in the Animal. Steer 513 502 509 501 512 500 3 yr. 8 mo. 3 yr. 8 mo. 3 yr. 8 mo. 3yr. 11 mo. 3yr. 11 mo .3yr. 11 mo 15 da. 19 da. 22 da. 21 da. 26 da. Group 1 2 3 1 2 3 Weight of lean in animal 132,060 102,639 92,018 130,041 101,203 91,990 Percent of total lean in head 1.397 1.649 1.389 1.540 1.613 1.593 Percent of total lean in shin 3.513 3.623 4.096 3.591 3.881 4.565 Percent of total lean in neck 0.747 0.920 1.005 0.641 0.790 0.950 Percent of total lean in chuck 27.632 29.227 28.411 27.313 29.911 28.457 Percent of total lean in plate 13.834 9.797 10.323 13.913 11.443 11.578 Percent of total lean in rib 9.368 8.893 8.890 8.011 8.354 7.393 Percent of total lean in loin 16.852 17.101 16.755 17.685 15.840 16.139 Percent of total lean in flank 3.208 2.271 2.301 3.494 1.825 3.049 Percent of total lean in rump 2.733 2.921 2.859 2.934 3.018 2.686 Percent of total lean in round 19.227 21.642 21.939 19.275 21.446 21.686 Percent of total lean in shank 1.373 1.768 1.856 1.413 1.697 1.683 Percent of total lean in tail 0.115 0.187 0.176 0.191 0.181 0.221 Table 36. — Distribution of Fat Flesh in the Animal. Steer 556 554 555 557 552 548 Age 3 mo. 3 mo. 3 mo. 5 mo. 17 da. 5 mo. 7 da. 5 mo. 9 da. Group 1 2 3 1 2 3 Weight of fat in animal Percent of total fat in head Percent of total fat in shin Percent of total fat in neck Percent of total fat in chuck Percent of total fat in plate Percent of total fat in rib 2,045 3.227 3.961 3.472 15.795 7.628 1.907 10.269 20.049 11.540 3.619 16.773 1.760 1,343 5.287 6.031 3.797 13.999 9.308 833 8.643 8.523 13.926 5.042 11,780 3.090 1.647 1.265 14.584 15.017 6.859 13.701 18.981 8.922 4.049 10.747 0.993 0.144 2.758 5.584 2.538 1.958 14.822 10.442 2.067 9.065 18.891 11.965 4.061 16.461 2.139 1,339 4.406 3.809 4.108 14.414 9.783 1.494 10.605 14.339 12.696 2.689 19.417 2.240 Percent of total fat in kidney fat Percent of total fat in loin Percent of total fat in flank Percent of total fat in rump Percent of total fat in round Percent of total fat in shank Percent of total fat in tail . 8.935 15.934 9.978 5.436 19.360 1.936 7.803 16.447 9.364 4.442 22.569 3.241 Table 37. — Distribution of Fat Flesh in the Animal. Steer 547 550 558 541 538 540 8 mo. 8 mo. 8 mo. 10 mo. 10 mo. 11 mo. 5 da. 14 da. 12 da. 22 da. 26 da. 2 da. Group 1 2 3 1 2 3 Weight of fat in animal 9,123 4,188 1,601 18,722 5,672 4,262 Percent of total fat in head 4.428 4.585 9.744 1.229 2.468 4.036 Percent of total fat in shin 1.852 1.552 3.186 1.699 1.693 1.619 Percent of total fat in neck 1.261 0.955 1.249 0.977 2.891 0.469 Percent of total fat in chuck 14.250 17.383 16.802 11.804 18.212 17.500 Percent of total fat in plate 13.592 10.697 8.495 13.155 14.616 15.157 Percent of total fat in rib 6.040 2.865 1.749 6.500 3.755 3.801 Percent of total fat in kidney fat 8.933 9.026 7.058 16.174 5.483 8.001 Percent of total fat in loin 21.769 25.478 18.738 21.600 20.804 27.076 Percent of total fat in flank 10.161 10.029 6.996 10.880 9.362 7.297 Percent of total fat in rump 4.253 4.776 3.248 4.535 4.725 5.115 Percent of total fat in round 11.783 11.270 20.112 10.293 15.004 9.503 Percent of total fat in shank 1.622 1.337 2.623 1.010 0.846 0.375 Percent of total fat in tail 0.055 0.048 0.144 0.141 0.047 Studies In Animal Nutrition — II 69 Table 38. — Distribution of Fat Flesh in the Animal. Steer 505 503 532 531 504 523 525 Age 10 mo. 11 mo. 1 yr. 5 mo. 1 yr. 6 mo. 1 yr. 8 mo. 2 yr. 2 mo. 2 yr. 2 mo. 18 da. 11 da. 20 da. 12 da. 26 da. 6 da. 8 da. Group 1 2 1 3 1 2 3 Weight of fat in animal 18,742 10 475 36.977 4,952 44,980 15,214 8,234 Percent of total fat in head 2.172 4.229 1.901 2.827 0.720 1.006 3.085 Percent of total fat in shin 1.094 1.403 1.266 2.262 1.670 1.729 2.417 Percent of total fat in neck 0.720 1.728 0.749 2.383 0.518 0.381 1.421 Percent of total fat in chuck. . . . 13.440 11.437 13.484 17.750 12.750 16.012 14.149 Percent of total fat in plate 13.798 14.081 15.299 12.298 15.480 13.330 14.258 Percent of total fat in rib 7.043 4.010 8.375 3.413 7.526 5.002 4.032 Percent of total fat in kidney fat 15.351 10.148 15.867 7.330 12.672 10.221 7.639 Percent of total fat in loin 20.163 27.427 20.221 22.859 20.387 20.954 22.820 Percent of total fat in flank 9.273 7.561 10.033 7.512 9.931 9.734 10.347 Percent of total fat in rump .... 4.877 5.642 3.946 5.089 5.645 5.981 5.927 Percent of total fat in round .... 11.050 10.587 8.200 15.044 10.914 14.973 11.914 Percent of total fat in shank 0.864 1.632 0.592 1.111 1.716 0.552 1.846 Percent of total fat in tail 0.155 0.115 0.068 0.121 0.071 0.125 0.146 Table 39. — Distribution of Fat Flesh in the Animal. Steer 515 507 529 527 526 524 Age 2 yr. 9 mo. 16 da. 3 yr. 2 mo. 21 da. 3 yr. 3 mo. 15 da. 3 yr. 4 mo. 3 yr. 4 mo. 13 da. 19 da. Group 1 2 1 1 2 3 Weight of fat in animal Percent of total fat in head Percent of total fat in shin Percent of total fat in neck Percent of total fat in chuck Percent of total fat in plate Percent of total fat in rib Percent of total fat in kidney fat Percent of total fat in loin Percent of total fat in flank Percent of total fat in rump Percent of total fat in round Percent of total fat in shank Percent of total fat in tail 89,188 1.393 1.724 0.748 12.576 20.570 9.129 5.562 21.485 9.814 5.003 10.684 1.239 0.074 22,267 3.036 1.159 0.804 12.471 15.224 5.461 9.826 22.877 9.665 6.099 12.076 1.298 0.054 125,332 0.497 1.022 0.481 14.943 20.466 9.685 7.566 21.034 9.024 6.032 8.564 0.668 0.019 24,937 1.648 0.995 1.335 15.062 16.482 7.459 6.464 23.327 10.711 5.987 10.057 0.393 0.080 6,834 5.414 2.766 0 556 13.330 14.969 2.473 5.604 17.881 11.121 5.838 18.481 1.390 0.176 Table 40. — Distribution of Fat Flesh in the Animal. Steer 513 502 509 501 512 500 Age 3 yr. 8 mo. 15 da. 3 yr. 8 mo. 19 da. 3 yr. 8 mo. 22 da. 3yr. 11 mo. 3yr. 11 mo. 21 da. 3 yr. 1 1 mo 26 da. Group 1 2 3 1 2 3 Weight of fat in animal 116,017 21,789 17,048 144,196 37,709 16,940 Percent of total fat in head 0.558 1.005 0.774 0.210 0.936 1.122 Percent of total fat in shin 0.896 1 629 1.396 0.848 1.223 1.865 Percent of total fat in neck 0.531 0.863 1.003 0.387 0.464 1.169 Percent of total fat in chuck 14.977 19.469 16.712 12.890 14.612 13.040 Percent of total fat in plate 20.796 15.659 16.987 21.006 19.820 18.017 Percent of total fat in rib 10.174 7.660 5.801 9.821 7.157 5.325 Percent of total fat in kidney fat 6.245 6.691 4.622 6 777 6.285 7.178 Percent of total fat in loin 21.518 20.983 22.202 24.743 20.298 20.159 Percent of total fat in flank 9.700 9.170 8.112 9.810 9.470 10 018 Percent of total fat in rump 5.113 4.828 6.183 5.060 5.802 6.246 Percent of total fat in round 8.235 10.602 14.975 7.727 13.180 14.569 Percent of total fat in shank 1.214 1.345 1.032 0.680 0.655 1.092 Percent of total fat in tail 0.044 0.096 0.199 0.041 0.098 0.201 70 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 41. — Distribution of Skeleton in the Animal. Steer 556 554 555 557 552 548 Age 3 mo. 3 mo. 3 mo. 5 mo. 17 da. 5 mo. 7 da. 5 mo. 9 da. Group 1 2 3 1 2 3 Wt. of skeleton in animal 10,377 9,404 8,304 14,705 10,652 9,877 Percent of total skeleton in head 12.576 11.857 11.911 13.499 14.327 15.136 Percent of total skeleton in shin 8.201 7.784 8.755 7.637 7.276 8.019 Percent of total skeleton in neck 0.964 1.414 2.589 1.401 1.070 1.934 Percent of total skeleton in chuck 17.029 18.429 17.438 17.769 17.275 16.442 Percent of total skeleton in plate 7.450 6.678 7.286 8.439 7.041 7.502 Percent of total skeleton in rib 7.931 7.199 7.563 8.745 9.783 8.170 Percent of total skeleton in loin 10.929 9.932 7.611 8.426 10.290 8.363 Percent of total skeleton in flank 0.154 0.117 0.157 0.054 0.075 0.162 Percent of total skeleton in rump 2.930 3.892 3.047 3.264 3.004 2.885 Percent of total skeleton in round 9.512 9.050 9.948 9.133 7.633 9.608 Percent of total skeleton in shank 9.406 10.198 10.261 8.875 8.177 8.667 Percent of total skeleton in tail 0.443 0.670 0.446 0.660 0.620 0.476 Percent of total skeleton in feet 12.475 12.777 12.988 12.098 13.430 12.635 Table 42. — Distribution of Skeleton in the Animal. Steer 547 550 558 541 538 540 Age 8 mo. 5 da. 8 mo. 14 da. 8 mo. 12 da. 10 mo. 22 da. 10 mo. 26 da. 11 mo. 2 da. Group 1 2 3 1 2 3 Weight of skeleton in animal 13,798 11,670 10,680 19,173 13,906 12,482 Percent of total skeleton in head 11.893 13.539 14.869 14.082 14.389 14.749 Percent of total skeleton in shin 6.726 7.506 8.090 7.683 7.939 7.819 Percent of total skeleton in neck 0.790 1.260 0.964 1.763 1.438 1.706 Percent of total skeleton in chuck 17.763 19.974 18.024 17.264 18.021 17.906 Percent of total skeleton in plate 7.023 6.530 5.075 7.380 7.637 7.395 Percent of total skeleton in rib 7.871 7.455 7.097 8.246 7.745 7.307 Percent of total skeleton in loin 11.349 10.557 10.009 10.212 9.694 10.215 Percent of total skeleton in flank 0.072 0.069 0.037 0.089 0.072 0.064 Percent of total skeleton in rump 2.754 2.999 3.174 2.754 2.632 2.492 Percent of total skeleton in round 9.016 8.997 9.803 8.475 9.190 9.414 Percent of total skeleton in shank 9.182 8.663 9.860 9.764 9.377 9.229 Percent of total skeleton in tail 0.522 0.403 0.375 0.537 0.489 0.553 Percent of total skeleton in feet 15.038 12.048 12.622 11.751 11.384 11.152 Table 43. — Distribution of Skeleton in the Animal. Steer 505 503 532 531 504 523 525 10 mo. 11 mo. 1 yr. 5 mo. 1 yr. 6 mo. 1 yr. 8 mo. 2 yr. 2 mo. 2 yr. 2 mo. Age 18 da. 11 da. 20 da. 12 da. 26 da. 6 da. 8 da. Group 1 2 1 3 1 2 3 Weight of skeleton in animal 18,873 20,561 31,116 16,539 28,747 25,608 21,552 Percent of total skeleton in head . 12.876 14.377 11.014 14.198 13.988 14.128 13.943 Percent of total skeleton in shin. . 7.837 7.626 8.687 8.427 7.528 7.791 7.800 Percent of total skeleton in neck. . 1.399 2.091 1.189 1.555 1.579 1.476 1.847 Percent of total skeleton in chuck. 18.529 16.458 19.723 16.891 17.859 17.631 17.757 Percent of total skeleton in plate. 7.831 8.151 8.642 7.550 7.823 7.474 7.865 Percent of total skeleton in rib. . . 8.457 8.224 8.391 6.897 8.857 7.505 9.020 Percent of total skeleton in loin . . 10.263 9.323 10.037 8.572 10.175 10.391 9.113 Percent of total skeleton In flank . 0.106 0.204 0.161 0.151 0.129 0.105 0.148 Percent of total skeleton in rump. 2.930 2.553 3.667 3.206 4.223 3.456 3.577 Percent of total skeletonin round . 8.997 9.377 9.037 11.010 8.363 9.040 9.387 Percent of total skeleton in shank. 8.970 9.114 8.597 9.716 8.589 9.513 8.473 Percent of total skeleton in tail . . . 0.636 0.462 0.427 0.448 0.550 0.617 0.520 Percent of total skeleton in feet. . 11.169 12.037 10.429 11.379 10.338 10.872 10.551 Studies In Animal Nutrition — II 71 Table 44. — Distribution of Skeleton in the Animal . 515 507 529 527 526 524 2 yr. 9 mo. 19 da. 2 yr. 9 mo. 16 da. 3 yr. 2 mo. 21 da. 3 yr. 3 mo. 15 da. 3 yr. 4 mo. 3 yr. 4 mo 13 da. Group 1 2 1 1 2 3 Weight of skeleton in animal Percent of total skeleton in head Percent of total skeleton in shin Percent of total skeleton in neck Percent of total skeleton in chuck Percent of total skeleton in plate Percent of total skeleton in rib Percent of total skeleton in loin Percent of total skeleton in flank Percent of total skeleton in rump Percent of total skeleton in round Percent of total skeleton in shank Percent of total skeleton in tail Percent of total skeleton in feet 38,821 12.725 7.751 1.309 17.403 8.047 8.325 10.026 0.155 5.247 8.171 10.152 0.456 10.234 32,531 13.249 7.716 1.282 19.010 9.425 7.762 10.000 0.224 3.898 9.013 8.192 0.544 9.686 36,872 11.491 8.597 1.378 18.787 8.188 8.877 9.682 0.060 4.421 8.741 9.216 0.502 10.092 34,535 12.790 8.166 1.231 18.810 9.315 8.238 9.909 0.223 4.109 9.196 8.646 0.481 8.887 31,805 12.621 7.782 1.654 18.620 9.102 8.348 10.354 0.214 3.811 9.241 8.351 0.456 9.448 Table 45. — Distribution of Skeleton in the Animal. Steer 513 502 509 501 512 500 Age 3 yr. 8 mo. 15 da. 3 yr. 8 mo. 19 da. 3 yr. 8 mo. 22 da. 3yr. 11 mo. 3yr. 11 mo. 21 da. 3yr. 11 mo. 26 da. Group 1 2 3 1 2 3 Weight of skeleton in animal 38,521 36,226 32,433 40,301 40,214 35,704 Percent of total skeleton in head 10.799 13.220 12.715 12.980 12.018 12.539 Percent of total skeleton in shin 7.944 8.199 7.779 7.655 7.552 7.856 Percent of total skeleton in neck 0.927 1.275 1.656 1.109 1.542 1.650 Percent of total skeleton in chuck 19.226 19.555 19.437 18.466 19.018 18.586 Percent of total skeleton in plate 9.405 7.489 7.745 8.652 9.517 8.666 Percent of total skeleton in rib 9.211 8.171 7.687 7.925 8.626 7.271 Percent of total skeleton in loin 11.080 10.857 10.594 10.687 10.877 10.884 Percent of total skeleton in flank 0.249 0.226 0.154 0.117 0.167 0.227 Percent of total skeleton in rump 4.496 3.260 4.409 4.568 4.058 4.184 Percent ot total skeleton in round 9.143 9.253 9.281 9.012 9.730 9.968 Percent of total skeleton in shank 7.863 8.251 8.476 8.843 7.654 8.052 Percent of total skeleton in tail 0.387 0.610 0.595 0.377 0.517 0.541 Percent of total skeleton in feet 9.862 9.637 9.472 9.608 8.723 9.576 72 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 46. — Proportion of Lean, Fat, and Bone in Cuts of the Carcass. Steer 556 554 555 557 552 548 3 mo. 3 mo. 3 mo. 5 mo. 5 mo. 5 mo. 17 da. 7 da. 9 da. Group j 2 3 1 2 3 47.592 43.937 50.641 47.018 50.824 46.168 4.589 5.518 4.337 7.870 4.118 3.204 48.215 49.864 44.411 45.558 45.588 49.749 66.538 55.797 57.171 55.007 62.150 54.459 13.733 12.319 19.376 12.617 10.436 Percent of bone in neck 19.342 32.126 42.829 26.788 26.636 36.243 71.908 71 . 100 73.307 67.107 71.494 73.303 Percent of fat in chuck 4.250 2.788 1.979 13.127 5.210 2.811 Percent of bone in chuck 23.250 25.697 24.697 19.965 23.437 23.653 Percent of lean in plate 68.766 65.702 69.340 55.053 64 . 184 65.545 Percent of fat in plate 5.118. 5.568 1.993 26.250 9.796 5.188 Percent of bone in plate 25.361 27.973 28.714 18.415 25.510 29.347 Percent of lean in rib 64.829 67.957 65.587 60.387 63.942 65.091 Percent of fat in rib 1.569 15.315 1.827 0.829 Percent of bone in rib 33.119 31.576 32.794 24.375 33.397 33.458 Percent of lean in loin 68 . 980 71.086 74.436 62.528 68.331 73.344 Percent of fat in loin 8.153 5.235 4.417 23.871 10.122 4.969 Percent of bone in loin 22.549 22.847 20.374 13.227 21.294 21.377 Percent of lean in flank 69.009 72.045 82.514 51.498 63.850 70.635 Percent of fat in flank 27.189 25.141 14.208 48.456 35.294 26.984 Percent of bone in flank 1.843 2.064 2.368 0.369 0.856 2.540 Percent of lean in rump 59.179 50.842 57.041 45.756 55.826 59.803 Percent of fat in rump 7.997 8.193 5.263 28.813 11.348 4.439 Percent of bone in rump 32.829 41.077 35.989 26.981 32.421 35 142 Percent of lean in round 77.563 78.179 77.457 74.133 78.892 79.229 Percent of fat in round 5.617 5.134 4.087 12.547 7.287 4.474 Percent of bone in round 16.164 16.805 17.957 13.310 13.050 16.331 Percent of lean in shank 30.244 28.674 31.856 30.426 34.640 31.801 Percent of fat in shank 2.436 1.804 2.063 5.669 4.129 2.299 Percent of bone in shank 66.035 68.746 65.088 63.227 60.952 65.594 Studies In Animal Nutrition — II 73 Table 47. — Proportion of Lean, Fat, and Bone in Cuts of the Carcass. 547 550 558 541 538 540 8 mo. 8 mo. 8 mo. 10 mo. 10 mo. 11 mo. 5 da. 14 da. 12 da. 22 da. 26 da. 2 da 1 2 3 j 2 3 55.408 50.794 45.630 51.913 49.875 52.379 6.795 3.332 2.991 8.567 4.000 3.097 37.314 44.900 50.674 39.682 46.000 43.806 57.714 57.428 60.933 65.067 58.762 65.602 21.905 8.869 5.831 12.200 19.159 2.903 20.762 32.594 30.029 22.533 23.364 30.914 72.651 69.062 70.703 74.054 72.430 72.201 9.351 7.308 3.560 9.701 7.924 6.931 Percent of bone in chuck 17.631 23.399 25.473 14.529 19.224 20.766 Percent of lean in plate 61.578 64.726 67.445 61.830 62.441 63.969 Percent of fat in plate 21.365 11.963 6.618 24.230 16.191 14.769 Percent of bone in plate 16.695 20.347 26.375 13.920 20.742 21.102 Percent of lean in rib 66.504 67.553 66.933 67.138 66.855 69.075 Percent of fat in rib 11.172 3.890 1.176 14.102 5.481 4.740 Percent of bone in rib 22.019 28.201 31.849 18.320 27.715 26.682 68.563 Percent of lean in loin 65.439 66.316 68.387 65.654 71.073 Percent of fat in loin 19.146 15.599 6.882 22.677 13.174 14 . 789 Percent of bone in loin 15.097 18.012 24.524 10.980 15.050 16.340 Percent of lean in flank 59.215 59.582 80.348 44.832 67.015 67.523 Percent of fat in flank 40.009 38.147 19.478 54.685 37.029 31.963 Percent of bone in flank 0.432 0.727 0.696 0.456 0.697 0.822 Percent of lean in rump 48.334 53.475 53.464 49.837 54.318 58 . 769 Percent of fat in rump 25.343 16.949 6.005 30.705 18.663 16.769 Percent of bone in rump 24.820 29.661 39.145 19.096 25.487 23 . 923 Percent of lean in round 78.613 79.286 77.647 79.109 79.289 81.011 Percent of fat in round 9.889 6.386 5.174 11.292 8.267 4.877 Percent of bone in round 11.443 14.206 16.825 9.522 12.415 14.148 Percent of lean in shank 37.462 28.961 28 107 32.249 35.802 34.751 Percent of fat in 6hank 6.477 3.728 2.733 6.213 2.279 0.884 Percent of bone in shank 55.449 67.310 68.510 61.538 61.918 63.646 74 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 48. — Proportion of Lean, Fat, and Bone in Cuts of the Carcass. Steer 505 503 532 531 504 523 525 Age 10 mo. 11 mo. 1 yr. 5 mo. 1 yr. 6 mo. 1 yr. 8 mo. 2 yr. 2 mo. 2 yr. 2 mo. i§ 18 da. 11 da' 20 da. 12 da. 26 da. 6 da. 8 da. Group 1 2 1 3 1 2 3 Percent of lean in shin 54.469 48.168 52.566 51.561 51.870 54.058 55.938 Percent of fat in shin 5.536 4.414 6.942 3.605 12.265 5.323 4.598 Percent of bone in shin 39.941 47.087 40.092 44.834 35.342 40.376 38.840 Percent of lean in neck 61.398 47.282 56.843 71.981 54.948 67.693 66.044 Percent of fat in neck 12.931 15.617 18.767 9.133 15.169 4.170 7.539 Percent of bone in neck 25.287 37.101 25.068 19.892 29.557 27.175 25.644 Percent of lean in chuck 72.310 74.598 70.518 75.305 66.653 75.017 75.657 Percent of fat in chuck 11.472 6.365 13.022 5.744 16.554 8.430 5.648 Percent of bone in chuck 15.927 17.980 16.029 18.245 14.819 15.624 18.554 Percent of lean in plate 58.946 58.021 54.207 65.384 50.870 64.227 66.047 Percent of fat in plate 26.008 19.459 30.613 11.107 36.832 16.907 13.721 Percent of bone in plate 14.865 22.111 14.552 22.761 11.896 15.957 19.811 Percent of lean in rib 65.644 67.411 59.919 69.975 60.532 68.731 71.711 Percent of fat in rib 15.336 6.340 21.384 3.770 22.144 8.694 4.082 Percent of bone in rib 18.543 25.525 18.028 25.429 16.656 21.958 23.900 Percent of lean in loin 63.007 63.844 66.024 73.011 57.821 68.547 70.652 Percent of fat in loin 24.190 21.321 27.322 11.742 31.489 16.918 14.191 Percent of bone in loin 12.399 14.266 11.412 14.698 10.044 14.121 14.833 Percent of lean in flank 52.169 63.519 43.843 68.715 45.628 60.053 65.589 Percent of fat in flank 47.422 33.991 55.439 29.689 53.949 39.025 32.794 Percent of bone in flank 0.546 1.803 0.747 1.995 0.447 0.711 1.232 Percent of lean in rump 46.662 47.158 49.518 56.141 43.493 49.600 55.250 Percent of fat in rump 32.807 27.540 28.112 14.133 38.290 25.090 16.968 Percent of bone in rump 19.849 24.464 21.985 29.725 18.308 24.400 26.808 Percent of lean in round 75.371 78.760 76.110 80.515 71.625 78.425 81.708 Percent of fat in round 13.466 7.596 12.125 5.581 18.884 10.540 5.824 Percent of bone in round 11.040 13.205 11.245 13.634 9.248 10.711 12.011 Percent of lean in shank 35.321 32.567 37.868 36.559 31.991 34.177 33.929 Percent of fat in shank 5.615 5.619 4.677 2.071 16.027 2.168 5.026 Percent of bone in shank 58.683 61.584 57.134 60.467 51.256 62.881 60.384 Studies In Animal Nutrition — II 75 Table 49. — Proportion of Lean, Fat, and Bone in Cuts of the Carcass. Steer 515 507 529 527 526 521 Age 2 yr. 9 mo. 2 yr. 9 mo. 3 yr. 2 mo. 3 yr. 3 mo. 3 yr. 4 mo. 3 yr. 4 mo 19 da. 16 da. 21 da. 15 da. 13 da. Group 1 2 1 1 2 3 45.663 55 . 742 49.535 54.559 49.334 18.375 4.087 14.517 3.653 3.544 35.950 39.759 35.925 41.538 46.409 48.514 61 . 949 39.792 57.781 61.236 29.152 11.111 32.915 18.377 2.580 22.203 25.885 27.729 23.455 35.709 63.029 74.907 58.273 70.957 75.584 Percent of fat in chuck 22.750 7.617 30.206 10.461 3.510 Percent of bone in chuck 13.704 16.961 11.172 18.092 20.973 Percent of lean in plate 37.818 60.616 37.983 59.327 62.445 Percent of fat in plate 52.605 20.418 55.024 22.703 9.756 Percent ot bone in plate 8.958 18.467 6.476 17.771 27.608 Percent of lean in rib. 45.259 67.847 49.110 45.527 64.558 69.477 Percent of fat in rib 38.757 10.451 36.531 42.742 13.911 1.834 Percent of bone in rib 15.385 21.702 14.064 11.524 21.277 28.809 Percent of lean in loin 47.464 63.583 45.427 62.855 72 . 646 Percent of fat in loin 43.705 21.793 47.768 23.259 7.337 Percent of bone in loin 8.877 13.917 6.469 13.683 19.771 Percent of lean in fla nk 27.089 52.353 29.832 45.429 64.936 Percent of fat in flank 72.267 45.816 69.957 54.146 31.575 Percent of bone in fl \nk 0.495 1.554 0.136 1.561 2.825 Percent of lean in rump 34.413 49.047 32.432 48.782 50.000 Percent of fat in rump 44.598 26.206 55.098 25.974 12.345 Percent of bone in rump 20.360 24.669 11.880 24 . 687 37.500 Percent of lean in round 62.832 77.752 64.656 79.379 81.941 Percent of fat in round 27.885 10.639 27.004 8.925 5.488 Percent of bone in round 9.282 11.601 8.109 11.302 12.771 Percent of lean in shank 27.059 38.314 34.038 36.423 33.841 Percent of fat in shank 15.964 5.940 13.009 2.014 2.261 Percent of bone in shank 56.934 54.779 52.813 61.377 63.208 76 Missouri Agr. Exp. Sta. Research Bulletin 54 Table 50. — Proportion of Lean, Fat, and Bone in Cuts of the Carcass. 513 502 509 501 512 500 3 yr. 8 mo. 19 da. 3 yr. 8 mo. 22 da. 3yr. 11 mo. 3yr. 11 mo. 21 da. 3yr. 11 mo 26 da. 15 da. j 2 3 1 2 3 Percent of lean in shin » 53.029 11.900 52.887 5.048 57.780 3.649 51.785 13.562 52.817 6.199 57.090 4.297 34.979 42.235 38.679 34.154 40.836 38.138 50.772 59.557 56.575 45.247 50.000 52.651 31 . 687 11.861 10.459 30.310 10.938 11.928 18.364 29.148 32.844 24.280 38.750 35.482 59.413 72.279 73.539 57.534 69.481 74.371 Percent of fat in chuck 28.291 10.221 8.014 30.106 12.647 6.276 Percent of bone in chuck 12.058 17.069 17.735 12.055 17.555 18.853 Percent of lean in plate 39.560 61.906 63.653 34.810 50.339 63.406 Percent of fat in plate 52.244 21.005 19.406 58.276 32.487 18.169 Percent of bone in plate 7.845 16.702 16.833 6.709 16.635 18.419 Percent of lean in rib 44.456 66.044 69.765 37.494 58.195 65.894 Percent of fat in rib 42.415 12.076 8.435 50.970 18.579 8.7-39 Percent of bone in rib 12.749 21.417 21.262 11.496 23.880 25.153 Percent of lean in loin 42.388 67.110 67.724 36.472 57.149 66.998 Percent of fat in loin 47.548 17.481 16.626 56.583 27.286 15.411 Percent of bone in loin 8.129 15.038 15.093 6.830 15.593 17.537 Percent of lean in flank 27.230 53.098 59.500 24.211 33.570 61.164 Percent of fat in flank 72.326 45.513 38.870 75.373 64.904 37.004 Percent of bone in flank 0.617 1.868 1.405 0.250 1.218 1.766 Percent of lean in rump 31.972 57.181 51.167 29.313 44.454 49.018 Percent of fat in rump 52.551 20.065 20.498 56.053 31.849 20.988 Percent of bone in rump 15.344 22.525 27.810 14.142 23.755 29.637 Percent of lean in round 65.686 79.267 78.060 62.710 70.640 76.512 Percent of fat in round 24.716 8.243 9.872 27.876 16.176 9.466 Percent of bone in round 9.111 11.962 11.639 9.087 12.736 13.650 Percent of lean in shank 29.031 35.749 36.818 28.789 33.859 33.240 Percent of fat in shank 22.546 5.771 3.794 15.358 4.871 3.973 Percent of bone in shank 48.503 58.873 59.258 55.853 60.698 61.735 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 55 Studies In Animal Nutrition III. Changes in Chemical Composition on Different Planes of Nutrition COLUMBIA, MISSOURI OCTOBER, 1922 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY P. E. BURTON H. J. BLANTON St. Louis Joplin Paris ADVISORY COUNCIL THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION J. C. JONES, PH. D., LL. D., PRESIDENT OF THE UNIVERSITY F. B. MUMFORD, M. S., DIRECTOR STATION STAFF OCTOBER, 1922 AGRICULTURAL CHEMISTRY RURAL LIFE O. R. Johnson, A. M. C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, Ph. D. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. Sieveking, B. S. in Agr. AGRICULTURAL ENGINEERING J. C. Wooley, B. S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale, B. S. in Agr. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Nelson, B. S. in Agr. W. P. Hays ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. C. McBride, B. S. in Agr. FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, Ph. D. O. W. Letson, B. S. in Agr. Miss Regina Schulte* S. D. Gromer, A. M. E. L. Morgan, A.M. Ben H. Frame, B. S. in Agr. Owen Howells, B. S. in Agr. HORTICULTURE T. J. Talbert, A. M. H. D. Hooker, Tr., Ph. D. J. T. Rosa, Jr., Ph. D. H. G. Swartwout. B. S. in Agr. J. T. Quinn, B. S. in Agr. POULTRY HUSBANDRY H. L. Kempster, B. S. Earl W. Henderson, B.S. SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M W. A. Albrecht, Ph. D. F. L. Duley, A.M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr. Richard Bradfield, Ph. D. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Secretary S. B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsiiam, Librarian. E. E. Brown, Business Manager. In service of U. S. Department of Agriculture. STUDIES IN ANIMAL NUTRITION III. Changes in Chemical Composition on Different Planes of Nutrition. C. Robert Moulton, P. F. Trowbridge*, L. D. Haigh The changes experienced by beef cattle in form and weight and in proportions of carcass and offal when on different planes of nu- trition were presented in previous bulletinsf. The 31 representa- tive animals slaughtered at various intervals from three groups were used (with one exception) for a study of the chemical compo- sition of the various parts, organs, and cuts of beef. GENERAL TREATMENT For a general discussion of the treatment of the animals the previous bulletins must be consulted. The ration included milk for several months after birth, and timothy hay and grain were soon introduced. At weaning time the ration consisted of alfalfa hay and a grain mixture in the ratio of one to two. The grain consisted of six parts corn chop, three parts whole oats, and one part of old process linseed meal. The animals were early divided into three groups. Group I was fed all it would eat of the ration. Group II was fed for maxi- mum growth without permitting the laying on of much fat. Group III was fed for scanty or retarded growth. The Group II steers gained about a pound a day for the first two years while the Group III cattle gained but 0.69 pounds a day. The animals were slaughtered at intervals and a series of weights and measurements were taken. The wholesale cuts were divided into lean flesh, fatty tissue, and bone and tendon. Various composites and individual samples were analyzed, there being a rather large number of samples for each animal. METHODS OF PREPARATION OF SAMPLES The samples of the soft tissues and parts were passed through a power grinder equipped with four sets of plates, each plate hav- ♦Resigned September. 191S. tC. Robert Moulton, I*. F. Trowbridge, L. D. Haigh, Studies In Animal Nutrition. 1. Changes in Form and Weight on Different Planes of Nutrition, Research Bulletin 43. II. Changes in Proportions of Carcass and Offal on Different Planes of Nutri- tion, Research Bulletin 54, Missouri Agricultural Experiment Station. 4 Missouri Agr. Exp. Sta. Research Bulletin 55 in g holes of a different size than the other. Samples were ground through the coarser plate and then through the next size. The samples well mixed and quartered down if necessary and then ground through a finer plate. The large samples were then quartered again and ground through the finest plate. Very homogeneous and fine samples were easily obtained in this manner. An especially difficult sam- ple to make uniform was the respiratory system. The cartilaginous rings of the trachea would partly remain behind in the mill while the softer lungs were squeezed out past them. By means of a knife these rings were finely cut and mixed with the lungs. The hide sample was cut into thin strips with a knife, alternate strips being rejected in the larger samples. The strips were then cut into short lengths and ground through the mill already described. The grinding of the sample proceeded very solwly, but with repeated grindings the work advanced more rapidly and a final uniform and fine sample was obtained. The work of preparing and grinding the samples proceeded as rapidly as possible until the samples were in a position where there was no danger of decomposition or change. The samples were kept in jars provided with rubber gaskets, glass tops, and metal clamps so that no loss of moisture could occur. They were kept in cold storage at a temperature just above freezing, so that they re- mained fresh for analysis. The skeleton samples were ground through a Mann green bone grinder, mixed well and sampled. From this smaller samples were weighed out directly and rapidly, in triplicate, in tared procelain evaporating dishes. The size of the samples varied according to the coarseness or fineness of the bone. For finely ground samples 25 to 40 grams were considered sufficient while for coarse samples 100 grams or even more were sometimes taken. The dishes contain- ing the weighed samples were at once placed in vacuum desiccators and dried to a constant weight within 25 or 30 milligrams. They were then extracted with ether in specially constructed Sohxlet extractors. The residue was saved, the triplicates combined, and the whole ground in a steel mill until fine enough to pass through a millimeter sieve. The sample was allowed to become air dry and saved for a complete analysis later. Samples of horn and hoof were dried and reduced to a fairly fine state with a horseshoer’s rasp. A drug mill was then used to reduce the material to a finer state. Studies In Animal Nutrition — III 5 METHODS OF ANALYSIS The samples were analyzed for water, fat, nitrogen, ash and phosphorus, following in general official methods of the Association of Official Agricultural Chemists. Glycogen, dextrose, and sarco-lactic acid and similar flesh acids were not determined. The formation of the acids in flesh progressively increases from the time of slaughter up to a maximum and then a decrease follows as decomposition takes place until neutrality and finally alkalinity is reached*. The glycogenf con- tent varies considerably in different parts of the animal and de- creases quite rapidly at ordinary temperatures through hydrolysis to dextrose. Through determination of the glycogen content of a number of animals it is certain that in beef flesh the amount of glycogen will seldom exceed one-half of one percent. Water. — For this work the S. & S. extraction shells and glass tubes with hardened filter paper bottoms were filled about one-third full of ignited sea sand and then stuffed with fat-free absorbent cotton. In our later work cotton alone was used. The tubes were numbered consecutively, extracted with ether, dried in vacuo and weighed in glass stoppered weighing bottles. This was done pre- vious to the slaughtering. A counterpoised weighing bottle was found very convenient as it obviated complications arising from a broken weighing bottle, the use of a new bottle and subsequent corrections of weights. Scheibler vacuum desiccators six inches in diameter with stopcocks in the lid were filled to the depth of an inch with C. P. sulphuric acid (sp. gr. 1.84). A brass gauze or por- celain plate was placed on the shelf of the desiccator and one-half inch above this supported by corks or rubber stoppers was a second gauze. Clean paper was placed on this. It was necessary to have the ground glass surfaces and stopcocks fit well. A lubricant of three parts of hard paraffin and five parts of yellow vaseline was prepared by melting together these ingredients and allowing the mixture to cool slowly. In cold weather a little more vaseline is used and in hot weather a little more paraffin to give the mixture the proper consistency. The thoroughly mixed samples were placed in weighing bottles provided with short aluminum scoops and triplicate samples of three to five grams were weighed out. The cotton was removed from the extraction tube and placed in a flat-bottomed, shallow, ♦Trowbridge, P. F. and Orindley, IT. S., J. Araer. Chein. Soc. 28, (1906), 469. tTrowbridge, P. F. and Francis. C. K., J. Ind. and Eng. Chein. 2 (1910), 21 and 215. 6 Missouri Agr. Exp. Sta. Research Bulletin 55 glazed porcelain dish and the sand was poured carefully into the dish. The meat sample was placed on the sand and the whole was carefully and thoroughly mixed and then returned to the tube by a steel spatula. The cotton was used to wipe every trace of the sample from the dish and spatula. A large sheet of glazed paper prevented loss of sand. The last of the unused cotton was placed in the top of the tube. Later when cotton alone was used, the mixing of the sample was greatly facilitated and the danger of loss of sand was entirely removed. The sand, or cotton, was used to sep- arate the particles of the sample and so allow a more thorough dry- ing and extraction. Otherwise the samples had to be ground and reextracted a second time. The triplicate samples were placed in separate desiccators in order to avoid a loss in case a desiccator was broken or acid spilled on the cones. The desiccators held 15 to 20 tubes. The desiccators when full were exhausted to a one- centimeter vacuum by means of a Geryk duplex vacuum pump. The desiccators were rotated carefully twice a day to mix the concen- trated acid with the supernatant watery layer. After 24 to 48 hours or longer, as convenient, air was allowed to bubble slowly through a sulphuric acid tower into the desiccator until the vacuum was destroyed. The tubes were transferred to desiccators holding fresh acid and the drying was continued as before. The tubes were then transferred to glass stoppered weighing bottles and weighed in the weighing bottle. The drying was continued to constant weight as given in detail above. Fat. — The dry tubes from the moisture determinations were extracted for 24 hours in Sohxlet extractors, using ether. They were partially dried in an electric oven at a low temperature and then dried in the vacuum desiccators as given in detail above. They were weighed as above and dried again to constant weight. Loss in weight is fat. Nitrogen. — Nitrogen was determined by the Kjeldahl-Gun- ning-Arnold method. Triplicate samples were weighed out as in the fat determination and placed in S. & S. No. 595 filter papers and introduced into a 500-cc. Kjeldahl flask. For hide and hair 0.50 to 0.75 grams was used, for lean meat 1.00 to 1.25 grams, and for fat samples 2.50 to 3.50 grams. Other samples in accordance to the nitrogen content. Twenty-five cubic centimeters of C. P. con- centrated sulphuric acid was used for the meats and 35 to 50 cc. for fats. About 0.7 grams of mercury was added and the digestion was made on a digestion frame. When the sample had ceased Studies In Animal Nutrition — III 7 foaming and was not pasty, 7 to 10 grams of potassium or sodium sulphate was added and the digestion was continued for one or two hours. The flasks were then cooled and the necks washed down with water. They were again digested for an hour or more. About 300 cc. of nitrogen-free water was added to the cool flasks also a piece of paraffin the size of a pea and a few small pieces of granulated zinc. Then 85 cc. of the alkali solution (100 cc. for fats) was added carefully, the flask was connected with a condenser, the contents were mixed, the flasks boiled for 40 minutes and the distil- late caught in a wide-mouthed receiving flask containing the neces- sary amount of one-tenth normal hydrocholoric acid with some cochineal indicator. The above alkali solution was made by dis- solving 40 pounds of Greenbank alkali and 375 grams of potassium sulphide in 30 liters of distilled water. For fats and other foam- ing materials 800-cc. Kjeldahl flasks were used. Protein. — The protein was calculated by multiplying the nitro- gen by the factor 6.25. Ash. — Triplicate samples of ten to fifteen grams were weighed out as for fat and placed in numbered, tared porcelain crucibles. The samples were dried in ovens and then charred carefully. Later they were ashed over Fletcher burners, using a low heat and tak- ing plenty of time. In this way fusion and loss of chlorides was prevented. Phosphorus. — The crucibles from the ash determinations were leached with strong hydrochloric acid and a little nitric acid. The solutions were neutralized and ammonium nitrate was added. The phosphorus was precipitated to 65° C. with acid ammonium molyb- date. The yellow phospho-molybdate was filtered off, washed, dissolved in ammonia and hot water and the phosphorus was re- precipitated with magnesia mixture. The precipitate was ignited strongly in a gasoline muffle and weighed as the pyrophosphate. AIR DRY BONE SAMPLES Moisture and Ash. — Two-gram samples were weighed out in tared porcelain crucibles and dried at 100 to 110° C. The difference 'in weight between crucible plus sample and dry weight of crucible plus sample gave the moisture. The samples were then ashed by igniting over Fletcher burners until practically free from carbon and the ignition was completed in a muffle at a dull red heat. A clear white ash was readily obtained by this means in a short time. 8 Missouri Agr. Exp. Sta. Research Bulletin 55 Nitrogen. — The nitrogen was determined as given in detail above using 0.5 gram samples. Phosphorus. — The ash from the above determination was dis- solved by digestion in hot, dilute nitric acid and the solution was made up to 250 cc. Aliquots of 25 cc. were taken and the phos- phorus determined as given in detail above. COMPOSITION OF SAMPLES The percentage composition of each sample analyzed is shown in the Appendix in Tables 1 to 30 and the weights of the consti- tuents in Tables 31 to 60. The detailed weights for the separate parts included in each sample can be found in Research Bulletin 54. The weight of the entire sample is shown in the tables listed above. From this data samples can be composited and the composition of various classes of tissues, parts of the animal, or the entire animal can be calculated. The tables include the analyses of 1061 sam- ples from 30 different animals, or over 35 samples per animal. The samples listed are mutually exclusive. For example the circulatory system for Steer 500 weighed 1.562 kilograms and had 48.451 percent water, 37.638 percent fat, and so on. This sample consisted of the large arteries and blood vessels in the thorax, the pericardium, adherent fat, and the ears of the heart. The lean heart itself exclusive of the ears formed a separate sample weighing 1.284 kilograms and having 77.544 percent of water, 3.559 percent of ether soluble material, and so on. Each system listed is exclusive of those parts which follow as separate samples which parts would ordinarily be considered as part of the system. A few samples of horns, teeth or hoofs and dewclaws were lost or detsroyed before the analyses were completed. The composition of a similar sample was in such cases used to calculate the compo- sition of the sample destroyed or lost. Full explanation of this is given at the foot of each table where such instances occur. Since the plan of the experiment was slightly modified from time to time the number and content of the samples is not the same for all the animals. Consequently the samples can not all be com- pared directly. To facilitate comparison certain composited sys- tems are presented in Tables 61 to 71 in the Appendix. The Blood. — The composition of the blood is shown graphi- cally in figure 1. The water content is close to 80 percent being about 82 percent during the first two years and 78 to 80 percent Studies In Animal Nutrition — III 9 from 3 years on. There is a tendency for the percentage of water to be in inverse order to the plane of nutrition, i. e., the higher the plane the lower the percentage of water. Fat was not found in the blood by the method used for this work. The nitrogen content varies between 2.5 and 3.5 percent in- creasing with age and increased plane of nutrition, although there are a few exceptions to both rules. The ash content is close to 005 0.00 10C i2 T. 05 UJ 3 fc 00 1 O •*;=) ’ZKCl :nta< X c OMPO SITI0N 0! 1 r BL .00P> PHO$PHOR US. u — G e=Q# — ■ jU > —o ^ ASM © MITRC K5EN. O U.3.0 o o 2 - 5 1- § 90 tki UJ 80 70 o w P — k / G G ROUP ROUP IO n x £ w WA1 ELR. G ROUP XEC o ■ ■ -o — - 0 ! 10 15 20 25 30 3 ACE Ih MONTHS. 5 40 45 50 Fig. 1. — Composition of the blood of beef animals. 0.75 percent. It seems to be unaffected by age or plane of nutri- tion. The ash content of the blood of Steers 503, 504, and 505 is unusually low and probably is not normal. The phosphorus is 0.025 percent and is apparently unaffected by the plane of nutrition. It seems to be slightly less in the older animals than in the younger animals. The Nervous System. — The composition of the central nervous system — the brain and spinal cord — is shown in figure 2. The water content is between 65 and 75 percent. It decreases slightly with 10 Missouri Ac-r. Exp. Sta. Research Bulletin 55 age and does not seem to be affected by the plane of nutrition. The fat — ether soluble material — is 10 to 20 percent of this sample. It increases with age and seems not to be affected by the plane of nu- trition. The nitrogen content is low for animal tissue being about 1.7 percent. It seems to be independent of age or plane of nutri- tion. The ash varies from 1.4 to 1.9 percent, increasing with age Fig. 2. — Composition of the central nervous system of beef animals. but being independent of the plane of nutrition. The phosphorus content is 0.34 to 0.43 percent, increasing with age but being un- affected by the plane of nutrition. Steers 503 to 505 give too high values for their age. This sample is rather typical of the glands of the animal body being higher in ash and phosphorus than any class of tissue but the skeleton. Studies In Animal Nutrition — III 11 Digestive and Excretory System. — The composition of the composited digestive and excretory system is shown in figure 3. The external fatty tissue has largely been removed from this sys- tem. The water is 66 to 78 percent, the fat 3 to 19 percent, the nitrogen 2 to 2.6 percent, the ash 1.5 to 0.8 percent, and the phos- Fig. 3. — Composition of the digestive and excretory system of beef animals. phorus 0.27 to 0.15 percent. The fat content increases with age and plane of nutrition while the water, nitrogen, ash and phos- phorus decrease up to the age of 3 years. Thereafter the fat de- creases again while the other constituents increase. This may be due to the fact that the offal fat is somewhat more easily and com- pletely removed from the older and fatter animals. 12 Missouri Agr. Exp. Sta. Research Bulletin 55 The Liver. — The above sample is a conglomerate of several classes of tissue in which the glandular predominates. As an example of pure glandular tissue the liver will serve. Figure 4 shows the composition. With few exceptions the composition of the liver from 3 months to 4 years is strikingly constant. The wa- ter content is about 67 percent, the ether soluble matter 2 to 3 percent, the nitrogen 2.8 to 3.3 percent, the ash 1.35 to 1.60 per- cent and the phosphorus 0.30 to 0.35 percent. The plane of nutri- tion does not seem to affect the composition, and age has but little effect. There is a slight decrease in water and increase in fat, nitro- gen and ash with increasing age. The ash content of the 3-months- old Group I animal and the phosphorus content of Steer 527 are considered to be atypical and are probably due to errors. Again Studies In Animal Nutrition — III 13 as with the brain and spinal cord the ash and phosphorus content is quite high while the nitrogen content is about that of muscle tissue. The Spleen. — In contrast to the above glands the spleen (figure 5) has a rather constant composition and low fat content. The fat runs from 1.5 to 5 percent and is slightly greater in the Group I animals. It increases slightly with age. The water content is 75 to 78 percent. The nitrogen content is from 2.9 to 3.3 percent in the young animals and from 2.75 to 3.0 percent in the old animals. 030 PE + N RCE.r A 1TAGI E 0 OMPG o N 01 F SF r 3 n n 1. 025 T PH0 SPHC »RU3. \ g jj 020 o c z : S io © — NIT ROGE N. C ^7* y o tsL U 0- 80 70 R \T. Q WA* n 50 d I 0 5 10 15 20 25 30 35 40 45 50 ACE IN MONTHS. Fig. 5. — Composition of the spleen of beef animals. With the exception of two of the Group II animals the ash content is constant at 1.3 to 1.5 percent. The phosphorus varies from 0.260 to 0.325 percent in the young cattle and from 0.300 to 0.220 percent in the old cattle. It seems to decrease somewhat with age. On the whole the plane of nutrition has practically no effect on the composition of the spleen and there is but a slight change with age. The Heart and Neck Sweetbreads. — Not all of the other glands of the animals were analyzed as separate samples. In a number of cases the sweetbreads, spleen, pancreas and kidneys were ana- lyzed as separate samples. No figure is shown for these glands but 14 Missouri Agr. Exp. Sta. Research Bulletin 55 a study of the tables in the appendix shows that the heart and neck sweetbreads have from about 2 to over 60 percent fat and from 80 to 30 percent water. The nitrogen content runs between 1 and 3 percent following the water content. The ash and phosphorus are 1.7 and 0.34 percent respectively in the samples low in fat and about one-third those amounts in the samples high in fat. The fat content increases in the older fatter animals while the other constituents decrease. The pancreas is much like the heart and neck sweetbreads in composition. Both of these glands — the thy- mus and the pancreas — become so intermingled with fat in the older fatter animals that a good separation is impossible. The Kidneys. — The composition of the kidneys is rather con- stant. The water content is 74 to 78 percent. The fat content runs Studies In Animal Nutrition — III 15 from 2 to 12 percent, but on account of the lack of uniformity in re- moving the kidney fat from the pelvis of the kidney this variation is not considered significant. The nitrogen varies from 2.08 to 2.7 percent, the ash from 1.00 to 1.35 percent, and the phosphorus from 0.19 to 0.25 percent. The Hair and Hide. — The hair and hide (figure 6) is a rather dry tissue having from 50 to 70 percent water, 1 to 13 percent fat, 4.8 to 6.6 percent nitrogen, 1 to 1.5 percent ash, and 0.10 to 0.05 percent phosphorus. The nitrogen content is higher than in any other tissue excepting hoofs, dewclaws, and horn exclusive of the bony core. The fat increases with age and plane of nutrition while the water content does just the reverse. The nitrogen percentage increases with age and is in inverse order to the plane of nutrition. The ash content seems to be rather independent of age and nutri- tion. It was difficult at times to insure perfectly clean hides at slaughter and some of the variations in ash content may be due to dirt on the animal. The phosphorus content decreases with age and seems to vary but little between the different planes. The Offal Fat. — The composition of the offal fat is shown in figure 7. A large range in composition is shown. This tissue has from 60 to 6 percent of water, 30 to 93 percent fat, 1.7 to 0.2 percent nitrogen, 0.7 to 0.1 percent ash, and phosphorus 0.12 to 0.01 per- cent. The fat increases with age and plane of nutrition, while all the other constituents decrease. The greatest changes are between the ages of 3 and 11 months. The Skeleton. — The composition of the skeleton, or bone, is shown in figure 8. The water content is from 30 to 57 percent, the fat from 8 to 23 percent, the nitrogen from 3 to 3.5 percent, the ash from 15 to 27 percent, and the phosphorus from 2.5 to 5 percent. The fat increases with age and is generally higher in the well fed animals although the difference is not great. The water content is just the reverse. The nitrogen content averages slightly higher in the older animals than in the younger animals while the plane of nutrition seems to have no effect. The ash and phosphorus con- tent of the older animals is about double that of the 3-months-old animals. This ossification is on the whole rather gradual. The plane of nutrition is here without effect. It has been shown in earlier work of this Experiment Station (Research Bulletin 28) that it is a difficult matter to alter the com- position of the bone by the plane of nutrition. The present study shows that aside from the small difference in fat and water content 16 Missouri Agr. Exp. Sta. Research Bulletin 55 Fig. 7. — Composition of the offal fat of beef animals. Studies In Animal Nutrition — III 17 Fig. 8. — Composition of the skeleton of beef animals. 18 Missouri Agr. Exp. Sta. Research Bulletin 55 Fig. 9. — Composition of the lean and fat flesh of beef animals. Studies In Animal Nutrition — III 19 the bones are not affected in composition by the three planes of nu- trition imposed. The Lean and Fat Flesh. — The composition of the lean and fat flesh is shown in figure 9. This sample is a composite of all the skeletal musculature and the fatty tissue associated with it. The offal and thoracic fat is not included. The figure shows, mainly, the effect of increasing fatness on the composition. The fat in- creases with fatness of the animal, i. e., with increasing age and plane of nutrition, while all other constituents decrease. The nitro- gen content of the Group II and Group III animals, however, is practically constant at about 3 percent. The water runs from 77 to 36 percent, the fat from 3 to 53 percent, the nitrogen from 3.2 to 1.5 percent, the ash from 1.15 to 0.50 percent, and the phosphorus from 0.190 to 0.090 percent. The Total Animal. — The composition of the total animal ana- lyzed is shown in figure 10. The figures are for the total animal less the fill and the loss on cooling and cutting. This basis is de- signated as the analytical animal in Tables 72 and 73. The average composition of 13 beef calves at birth* is included in the figure. The water content decreases from 73 percent at birth to 39 percent in the old fat steer. The higher the plane of nutrition and the older the animal the lower is the percent of water. The fat increases from about 4 percent at birth to about 45 percent. The increase follows age and plane of nutrition. The nitrogen shows first an in- crease from 2.9 percent at birth to 3.3 percent at 3 months. It re- mains practically constant thereafter for Groups II and III but de- creases in Group I to 2 percent at 4 years. The ash content for Groups II and III increases from 4.5 percent at birth to over 5 percent at 4 years. The high value for the Group III animal at 40 months is probably an error or an abnormality. For the Group I cattle the ash increases to about 5 percent at 3 months, falls to 4 percent at 5J4 months and remains there in spite of fattening until after 3 years when it drops to nearly 3 percent. It should be noted that the Group III 3-months-old calf was so greatly retarded in development by the low plane of nutrition that its ash and phos- phorus content is actually lower than that of the calves at birth. In general the phosphorus content of the entire animal follows the ash. The values for Steer 505, Group I, and Steer 503, Group II, are so much higher than those of the other animals of their groups Research Bulletin 38, Agr. Expt. Station, University of Missouri. 20 Missouri Agr. Exp. Sta. Research Bulletin 55 that they are not averaged on the curve but are shown separately. The phosphorus content increases in percentage for Groups II and III but decreases for Group I. Studies In Animal Nutrition — III 21 THE COMPOSITION ON A PROTOPLASMIC BASIS A brief summary of the composition of various parts and sam- ples of the beef steer given above shows that those parts or or- gans that become depots of deposit for fat, exhibit an increasing percentage of fat with increasing age and plane of nutrition ; while most, if not all, other constituents decrease. Certain organs remain fairly constant in composition. It is the belief of the senior author that the composition of animal tissue should be studied also on the fat-free, or protoplasmic, basis. Its usefulness has already been demonstrated by Moulton* and Greenef. In the animal body lipoid matter can be divided largely into two classes: (1) stored and inactive lipins, largely glycerol esters of the higher fatty acids ; and, (2) those lipins that are essential to the protoplasmic structure and that take part in the physiological activities of the tissue, such as lecithins and cholesterol. The former is largely if not entirely in- ert stored matter and should not be considered as part of the proto- plasmic tissue. Unfortunately the usual method of extraction by ether removes both classes together ; but, since in the fatty tissue and even in the entire body of fat animals the former very greatly predominates, the ether extract can safely be called stored fat. For the above reasons the composition will now be considered on the fat-free basis. Tables 72 and 73 show the composition of the entire animal on the analytical basis, the empty weight basis, and the fat-free basis. The first basis has been defined just above. The second assumes that the loss on cooling and cutting is water, which it must largely be. This weight is added to the water con- tent and the composition recalculated. The result is a slightly larger water percentage and slightly smaller percentages of the other constituents. The third basis assumes that all ether soluble fat had been removed. Thirteen calves at birth and three embryos reported in Re- search Bulletin 38 of this Station are included in the tables. In or- der to complete the picture of the development of the composition of mammalian tissue a search has been made for analyses of other mammalian embryos. On account of the length of the gestation period and relative size of the animal it is thought that rabbits and other mammals cannot serve our purpose. The composition of some 21 human embryos reported by FehlingJ in 1877 and recalcu- *.T. Biol. Them. XLIII, 67. t.T. Biol. Cliem. XXXIX, 435. tArchlv. f. Gynaekologle XI, 523. 22 Missouri Agr. Exp. Sta. Research Bulletin 55 lated by the senior author to the fat-free basis have been added to the results obtained from the bovine. Figure 11 shows the water content of the animals on the fat- free basis from the beginning of gestation to maturity. The gesta- tion periods for man and the ox are practically the same. Man is less mature at birth, however, and this should be borne in mind in considering the composition of the full term human infant. The water content of the fat-free human embryo at the beginning of the sixth week of gestation, or at an intra-uterine age of 35 days, is 97.5 percent. It decreases rapidly and uniformly to about 86 percent at 6 months. It is seen that the ox embryo at this age has practi- cally the same composition as the human. At birth the ox has 76.5 percent water. The human infant being less mature has 81.5 per- cent. At the age of 3 to 5 months there is a marked change in the rate of decrease of the water in the ox. It is about 72 percent at 5 months and 70 percent at 4 years. The plane of nutrition has prac- tically no effect on the composition of the ox, on the protoplasmic basis. The percentage of nitrogen is shown in figure 11. At about 35 days (intrauterine) it is 0.4 percent. It increases rapidly and uni- formly to about 3.0 percent at birth and at 5 months is 3.5 percent. Maturity is reached at about 11 months when the percentage is 3.6. This continues to be the value excepting for a few of the old thin animals which exceed it by about 0.2 percent. The ash content at 35 days is practically nothing. It increases rapidly and uniformly to 4.3 percent at birth. At 5 months it is 5 percent and thereafter increases slowly to about 5.7 percent at 4 years. There is more variation in the ash content than in the wa- ter or nitrogen content. It is higher in the low plane animals than in the high plane animals. This is probably due to a small propor- tion of bone in the Group I cattle. The phosphorus content of the human embryos was not given. Therefore the figure shows the results for the ox only. The phos- phorus content on the protoplasmic basis is about 0.3 percent at 185 days intrauterine. It increases rapidly to about 0.74 percent at birth. By 11 months the value is about 0.90 percent and there- after it increases very slowly to 1 percent at 4 years. These figures show, then, that the evolution of the tissue of such mammals as the ox is rapid from conception to shortly after birth — about 5 to 11 months in the ox. Thereafter the composi- tion on the fat-free, or protoplasmic, basis is practically constant Studies In Animal Nutrition — III 23 Fig. 11. — Composition of the fat-free beef animal. 24 Missouri Agr. Exp. Sta. Research Bulletin 55 there being but slight changes to full maturity. Such relations as these are entirely destroyed by the presence of stored fat in the animal body and would be left undiscovered if the fresh, fat-con- taining basis were used. THE COMPOSITION OF OX MUSCLE ON THE PROTOPLASMIC BASIC The above results show the advisability of studying the com- position of such tissues as striated muscle on the fat-free basis. Un- fortunately only the composite embryo was studied by us. How- ever, Buglia and Costantino* have recently reported some analyses of ox embryo muscle. Three samples of embryo muscle at 75, 120 and 135 days were analyzed for water, fat and nitrogen. These results are included with all samples of lean muscle reported in this bulletin and are shown in Tables 74, 75, and 76. Figure 12 shows the percentage of water in ox muscle from miduterine life to maturity. At mid-term the tissue is 87.5 percent water. At birth it is 80 percent and at 5^4 months it is 77 percent. It remains practically constant then at 76.5 percent with no appar- ent effect on the plane of nutrition. Perhaps more rigidly controlled conditions in sampling and analyzing might have resulted in less variation than is shown. The water content of the muscle is 5 to 6 percent higher than in the total animal. The nitrogen content is shown in figure 12. At miduterine age it is 1.4 percent, at birth 2.9 percent, and at 11 months 3.5 per- cent which is the value maintained to the end. This value is slightly less than that for the total animal. The percentage of ash exhibits some striking changes. There are no figures preceding birth. At birth the ash in the fat-free muscle is 1.05 percent. At 3 months of age it has risen to 1.28 per- cent and falls to 1.11 percent at 6 months. From then on it de- creases slowly and gradually to 1.06 percent at 4 years. The peak at 3 months may need verification, but it is a fact that only one other animal, the Group II steer at 40 months, exhibits anywhere near as high a figure. *Z. Physiol. Chemie 81 (1921), 143 and 155. Studies In Animal Nutrition — III 25 The phosphorus percentage in the fat-free muscle exhibits some rather similar changes. At birth the tissue shows about 0.172 percent and at 3 months 0.218 percent. The value thereafter falls fairly rapidly and uniformly to about 0.200 percent at 4 years. The figure confirms in general the relations shown by the ash percent- ages. 22 21 20 19 .16 V” z u] .16 D h u *0 Z LI o U l0 U. o 15 50 •0 til o 2.0 a: u) 1S 0. 95 90 65 1 1 ( i 80; COM POST noii or o TAT FRE E. L LAN fle Sh. £>C> o / 0 * IT > o - — PERC ENT PN02 PHO^ US. o o" ~l A n * PERC Q> ENT ASH. < ( O O 9 * T s_ jq ? r\ 9 JT y/L- Q 0 0 Q 9 u 0 8 ) O o ^ > Ox o PERC ilNT NIT! ?0GE 1. G ROUP To o G G ROUP ROUP IX TTT o O PE.RC ENT WA' ER. - \ k \ 3 \ i f 1 Vo 0 ft 9 n O ) On — ° 5 5 10 15 20 25 30 3 5 4 0 45 50 AGE IN MONTHS. Fig. 12. — Composition of the fat-free lean flesh of beef animals. 26 Missouri Agr. Exp. Sta. Research Bulletin 55 AMOUNT AND COMPOSITION OF GAIN Composition of Gain From Start to Slaughter. — The compo- sition of the animal at slaughter is given above. In order to calcu- late the composition of the gains made by each steer from the time it was put in the experiment until slaughter it is necessary to know the weight and composition at the start. The weight for each ani- mal at the beginning of the experiment is shown in the Appendix of Research Bulletin 43 of this series. Since the analysis is based upon empty weight in Tables 72 and 73 it is necessary to estimate the empty weight of each calf at the start. To facilitate this the percentage of empty weight is plotted against the live weight in PERCENT EMPTY WEIGMT IN YOUNG CATTLE AS AFTECTEP BY WEIGHT 40 50 60 70 60 90 100 110 120 . LIVE WEIGHT- Kilograms. Fig. 13. — Percent empty weight in young cattle. kilograms in figure 13. Only calves of 100 kilograms empty weight or less are shown. The line shows the relation between the per- centage of empty weight and the size of the animal for a normally fed beef calf. The Group III calves lie below the line. The live weight used in this figure is the average live weight for the last five days of the animal’s life. This is usually larger than the live weight before slaughter because at that time the cattle had been without water for the morning. From this figure it is possible to estimate accurately the probable percentage of live weight in each calf at the beginning of the experiment. Table 77 gives the empty weights at the start. Studies In Animal Nutrition — III 27 In figure 14 are presented the relations between the empty- weight of young calves and the composition of the calf. The com- position at 35 kilograms is the average of 13 beef calves at birth The lines show a fairly uniform relation between empty weight and composition. Using the empty weight of the calves shown in Table 77 the composition of each calf at the start can be accurately estimated. 065 0.60 C0MF ’QSITI Am on i CTEP OF Y< BY DUNG EMP 1 CAT’ rY v FLE ^EKJH AS T. PH0S PH0R JS. X. X g*5 / £H. X g X O 35 O X u. 3c 'x O ITRO JEN. K> z : H 5 1 r AT x , — <(■ & X uJ ft- 70 X 60 WA t* UJ * 30 40 50 60 70 80 SO 100. EMPTY WEIGHT :• Kilograms. Fig. 14. — Composition of young cattle. Table 78 shows the weights and percentage of each constituent for each animal at the start and at slaughter and the composition of the gain made. The animals are not in all cases ideal checks on each other consequently the composition of the gain does not vary uniformly. Figure 15 shows the composition of the gains made by each group from the start to slaughter. The first gains of the thinnest cattle are 80 percent water the next gain is but 62 percent water. The water increases slightly to 18.5 months and then decreases slowly. The Group II cattle 28 Missouri Agr. Exp. Sta. Research Bulletin 55 Fig. 15. — Composition of total gains of beef cattle. Studies In Animal Nutrition — III 29 show much the same sort of change but in a less marked degree. For the Group I cattle the gains at first are 50 to 60 percent water and only 38 percent at 4 years. In contrast to this the gains made by all cattle become higher in percentage of fat as the age advances. The Group III calf at 3 months actually shows a loss of fat. The next gain was 10 per- cent fat and at 4 years the gain contained 18 percent fat. The Group II cattle show increasing percentages of fat up to 11 months when growth becomes so rapid that the animal becomes relatively thinner and the gains contain relatively less fat. At 4 years the gain contains 27 percent of fat. The Group I cattle show this thinning down at 8y 2 months. Thereafter the gains increase rapidly in fat, containing at 4 years as much as 46 percent of fat. The gains of the Group I cattle decrease in percentages of nitrogen excepting during the period of rapid growth at 8J4 and 11 months when there is an increase. At the start the gain contains about 3.15 percent nitrogen and at the end only 1.9 percent. The Group II cattle show a decrease in percentage of nitrogen in the gain at first followed by a slight increase. Then the value becomes constant at 2.9 percent. The Group III cattle show much the same thing excepting that the value continues to increase up to 40 months when it is almost as great a part of the gain as it was at 3 to 5 months. As for the ash gained the Group III cattle show a very low per- centage at 3 months with a very rapid recovery at 5 y 2 months. Thereafter the value is fairly constant at 5 percent. The Group I and Group II cattle at first show the opposite tendency, the gains containing a relatively smaller percentage of ash. The Group II cattle then show a gradual increase to 2 years, after which the value is rather constant at about 5 percent. The Group I cattle continue to show a decrease in the percentage of ash with some rather large individual variations. The phosphorus content of the gains made in general follows the ash. It has perhaps been noticed that the lines miss a few of the points by a large margin. The following reasons will account for this. Steers 505 and 503, representing Groups I and II respectively, were among the first animals killed and analyzed. The other ani- mals at this age — 11 months — differed in weight or composition from these first two. There must be some difference in age or treatment to account for some of the large differences. Steers 505 30 Missouri Agr. Exp. Sta. Research Bulletin 55 and 503 are not considered to be quite typical. Again Steer 527, the Group I steer at 40 months, was too fat for its age and Steers 502 and 524 were too thin for their ages and groups. The former was the Group II 45-months animal and the latter the Group III 40-months animal. Figure 15 (as does also figure 10 which gives the composition of the total animal) raises the question of the regularity of the change in composition of the cattle and of the rate of deposition of each constituent. To throw more light on these questions there is pre- sented in figure 16 the weight of water found in each animal slaughtered. At birth it is about 25 kilograms. In the Group I cattle this increases rapidly and uniformly to 250 kilograms at 21 months. The rate of increase then declines until at about 35 months the steer has almost as much water as at 47 months. The curves Studies In Animal Nutrition — III 31 for the other groups are rather similar excepting that for Group II the break in the curve is at 26 months and flattening occurs at 40 months. For Group III the rate of increase has been still less, the break occurs at about 27 months and the flattening is at the end if present at all. In marked contrast to these curves are those for the fat shown in figure 17. After the third month the Group I cattle deposited fat at a rapid and very uniform rate averaging 7.86 kilograms per month. The curve is very slightly convex to the (abscissae) hori- zontal axis. For the other groups the rate of increase in weight of fat is very much smaller and the curves are more convex to the horizontal, i. e., the rate of deposition increases more with age. 32 Missouri Agr. Exp. Sta. Research Bulletin 55 Figure 18 shows the weights of nitrogen (or protein), ash, and phosphorus for each animal slaughtered. In general the curves resemble the curves for water more than they do the curves for fat. The Group I cattle show a decided break in the building up of pro- tein at 20 months and a further break at 40 months. The curves for the other groups are very similar. Both the ash and the phos- phorus, on the other hand, fail to show the flattening of the curve after three years. INCREASE. WITH AGE AND PLANE OF NUTRITION Fig. 18 . — Quantity of nitrogen (protein), ash and phosphorus in beef cattle. Studies In Animal Nutrition — III 33 COMPOSITION OF GAIN BETWEEN EACH AGE In order to calculate the composition of the gains made be- tween the succeeding ages it is necessary to assume that each steer slaughtered had at the age at which the preceding steer was slaughtered the composition of that steer both in percentage com- position and in percentage of empty weight. Table 79 gives the percentage of empty weight referred to the live weight at the end of the feeding period (a five-day average). This is more representa- tive of conditions in the pen than when the live weight just preced- ing slaughter is used. For the live weight at the age at which the preceding animal was slaughtered the live weight at the beginning of that period which came nearest to giving the correct age was used. This fa- cilitates the calculations and is as correct as any of the assump- tions. The composition of the gains made between each succeeding age for each group is given in Table 80. A study of the table shows that on the whole each animal at the time of slaughter contained more of each constituent than it did at the time the preceding animal was slaughtered. This is true with all but one animal in each group in the latter months. In these cases the three steers — 527 in Group I, 502 in Group II, and 524 in Group III — were not of normal condition for the group, the first being too fat and the latter two too thin for the age and group. These statements are borne out by data presented above on the composition of the steers and by the proportion of lean, fat and bone shown by these animals in Research Bulletin 54. These ani- mals must be omitted entirely from a study of the composition of the gains made between successive ages or else the composition of normal animals of the respective ages and groups must be used in place of the composition shown by those abnormal steers. From figure 10 the percentage of fat a steer should have on these three planes of nutrition can be read off for any given age. By its use it is estimated that Steer 527 should have had 38.5 per- cent of fat, Steer 502 should had had 20.3 percent of fat, and Steer 524 should have had 15 percent of fat. From figure 11 the normal composition of the fat-free animal is readily determined. Calculat- ing these values to the fat content just given it is found that these animals should have had the following composition. 34 Missouri Agr. Exp. Sta. Research Bulletin 55 Fig. 19. — Composition of successive gains of beef cattle — water, fat and phosphorus. Studies In Animal Nutrition — III 35 Estimated Percentage Composition. Steer Water Fat Nitrogen Ash Phosphorus 527 43.48 38.5 2.21 3.38 0.615 502 56.03 20.3 2.87 4.46 0.797 524 60.00 15.0 3.06 4.70 0.850 Fig. 20. — Composition of successive gains of beef cattle — ash and nitrogen. From these values the composition of the gains as shown in the latter part of the table are calculated. These corrected results are shown graphically in figures 19 and 20. 36 Missouri Agr. Exp. Sta. Research Bulletin 55 Water. — The water content of the gains made by the Group I steers decreased as the animals got older and fatter. The 8^2- months-old steer had been growing rather rapidly and these gains were largely protoplasmic tissue as shown by the high water and nitrogen content of the gain. From gains containing 60 to 70 per- cent of water the change becomes rapid to gains containing 30 to 40 percent of water. After 35 months the water content drops until at 47 months it is only a little over 2 percent. The Group II steers after an initial decrease in percentage of water in the gains show a rise to 11 months. This would indicate that their development was about 3 months behind the Group I cattle. At 34 months the steers become rather fat having only 20 percent of water in the gain made during the past 8 months. The next gains contain more water. The 48-months-old steer of this Group was too fat (and too well supplied with bone at the same time). Consequently his last gain appears to contain — 113 percent of water. This is, of course, impossible. For the Group III steers there is the initial fall in percentage of water in the gains followed by a rise with the maximum in this case deferred to 18 months. The water content of the successive gains then falls to about 34 percent at 45 months and shows a rise to about 55 percent at 4 years. Fat. — In general the fat content of the successive gains is in- versely to the water content. The first gains contain about 10 per- cent of fat. This increases, after an initial fall with minima occur- ring where the water showed maxima, to 90 percent for Group I and only 30 percent for Group III. The Group II cattle lie between these, but on account of the wide abnormality of the 4-year-old Group II steer the last gain appears to contain 191 percent of fat. This is of course impossible. Nitrogen. — The nitrogen content of the gains is shown in fig- ure 20. In general it follows the water and is inversely to the per- centage of fat. The Group I cattle show gains containing over 3 percent of nitrogen at first. The percentage of nitrogen decreases until at about 4 years the gains contain no nitrogen ( — 0.45 and — 0.15 percent). The Group II and Group III cattle show gains containing over 3 percent nitrogen at first. This percentage falls to less than 2 percent for Group III and about 2.5 percent for Group II. There is then an increase to over 3 percent again. Towards the end there are some rather sudden changes which can only be Studies In Animal Nutrition — III 37 accounted for by individual differences in the steers. At the end the values are not far from 3 percent in either case. Ash. — The ash content of the gains shows some rather striking changes. For the Group I cattle the percentage drops from 5 to 3 or 3.6 percent. But the 21-months-old steer shows a gain con- taining over 6 percent ash. The value then falls to 3 percent and 1 percent at 44 y 2 months. However at 47 months the gains con- tain nearly 7 percent of ash. These changes are partly due to dif- ferent proportions of bone in the cattle. The Group II cattle fol- low Group I in general. At 8^4 months the gain contains but 2.6 percent ash. It then shows a rise to nearly 6 percent falling rapidly after 26 months. The last two steers show abnormal values, the 45-months-old steer showing the gain of the last 5 months to con- tain — 3.3 percent of ash and the 4-year-old steer showing for the last 3 months a gain containing 32.37 percent of ash. These last two values are, of course, impossible and bear witness to the abnor- mality of the 4-year-old Group II steer as well as to the different proportions of bone in the last two animals. The Group III animals indicate that the early growth of the calves has been so retarded that the bone is insufficiently developed. The gains made immediately after 3 months contain 6 percent of ash and show that the animals are recovering in this respect. There is a big drop in the ash content of the gain made just preceding 11 months. The following gain is higher in ash and there then fol- lows a decrease. Towards the end there is another increase in the percentage of ash in the gains. Phosphorus. — The percentage of phosphorus in the successive gains is shown at the top of figure 19. On the whole the values appear fairly constant. This is partly due to the scale of the figure. The percentages vary much as do the ash percentages. The 4-year- old Group II steer is again abnormal and shows the gain for the last 3 months to contain 5.5 percent of phosphorus. SUMMARY Thirty Hereford-Shorthorn beef animals ranging in age from 3 months to 4 years were used in this experiment representing three different planes of nutrition. An average of 35 samples per animal or a total of 1061 samples were analyzed for water, fat, nitrogen, ash, and phosphorus. The chief effect of age and plane of nutrition on the composi- 38 Missouri Agr. Exp. Sta. Research Bulletin 55 tion of parts and total animal is through a change in the fat con- tent, which increases in most cases with age and plane of nutrition. The skeleton shows greatly increasing ash and phosphorus content with advancing age. The total empty animal shows an increasing fat content and decreasing percentage of other constituents with age and plane of nutrition excepting where the fattening is slight and a small in- crease in nitrogen, ash and phosphorus becomes apparent. When calculated to the fat-free basis, however, the total animal shows very striking changes in composition depending on age alone. The water content decreases rapidly from conception to about the age of 6 months and then becomes constant. The other constituents show a rapid increase to a maximum. For nitrogen and phosphorus the maximum is attained at about 11 months. The ash does not attain a maximum and constant value but from 5 months to 4 years increases slowly. The composition of the composite ox muscle on the fat-free or protoplasmic basis shows somewhat similar results. The mini- mum for water and the maximum for ash occur at about 6 and 11 months respectively. The ash and phosphorus content show irreg- ularities having a marked maximum at 11 months with a decreasing percentage thereafter. The amount and composition of the gains from start to slaughter and between each succeeding age have been calculated. The beef steer may contain 4 percent fat at birth and 45 percent at 4 years. For the full fed cattle the gains become richer in fat and poorer in other constituents with advancing age until the last gains are shown to consist of 90 percent fat. With the other groups there is some variation. All groups show a thinning down during the early months and a fattening after the period of rapid growth is over. The thin cattle have a more nearly constant composition after the first few months. The irregularity of the percentage composition of the gains raises a question as to the uniformity of the treatment. It is shown that the weight of water in the fattening beef steer increases rap- idly to 21 months and then slowly to 35 months. With the poorer groups the flattening of the curve occurs at 26 months and 40 months. The deposition of fat was very uniform from 3 months on for the full fed cattle and slightly increasing with age for the poorer cattle. Studies In Animal Nutrition — III 39 APPENDIX Table 1. — Steer 500. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % 21,269 1,562 1,284 3,747 568 79.041 0.192 3.193 0.789 0.022 48.451 37.638 1.867 0.541 0.061 77.544 3.559 2.723 1.107 0.208 76.501 2.702 2.840 1.145 0.172 17.038 79.391 0.489 0.239 0.024 832 62.766 21.718 1.667 1.748 0.371 Digestive and excretory system (partial) Offal fat 19.275 12.940 538 74.635 13.501 9.971 83.556 2.203 0.422 0.834 0.184 0.118 0.020 53.199 33.943 1.837 1.043 0.224 4.634 69.786 2.902 3.243 1.579 0.323 Gall 241 91.875 0.219 0.200 1.237 0.027 1.054 74.374 5.355 2.977 1.196 0.219 625 59.139 25.091 2.140 1.201 0.256 Kidneys 1,019 1,619 77.091 4.806 2.420 1.120 0.207 Tongue, marketable (excl. bones) 69.404 11.874 2.703 0.914 0.154 Hair and hide 35,938 59.327 1.319 6.280 1.072 0.044 Head and tail, lean and fat 3.784 12.496 63.713 15.958 3.155 0.882 0.134 Shin and shank, lean and fat 70.862 6.591 3.363 0.989 0.164 Flank and plate, lean and fat 36,410 7,058 58,918 54.788 27.651 2.687 0.866 0.139 Rump, lean and fat 55.149 27.639 2.527 0.819 0.145 Chuck and neck, lean and fat 67.594 11.869 3.387 0.912 0.158 Round, lean Round, fat 39,898 4,936 29.692 74.031 27.767 3.485 61.442 3.123 1.590 1.011 0.377 0.191 0.051 Loin, lean 70.269 7.734 3.113 1.010 0.185 Loin, fat 6.830 16.464 76.508 0.598 0.245 0.038 Rib, lean 13,602 67.137 12.323 3.196 0.929 0.170 Rib, fat 1,804 2.432 20.368 71.084 1.293 0.370 0.060 Kidney, fat 7.026 90.275 0.410 0.143 0.018 Skeleton of feet 6,838 8 953 39.603 11.528 3.612 24.970 4.529 Skeleton of head 47.986 13.584 3.487 17.862 3.434 Skeleton of tail 386 39.305 24.024 2.650 15.917 2.786 Skeleton of shin 5,610 26.487 21.585 3.700 29.441 5.211 Skeleton of shank 5,750 31.458 20.187 3.454 25.183 4.429 Skeleton of flank and plate 6,350 2,988 14,450 6.438 680 41.031 18.008 3.223 18.537 3.200 Skeleton of rump 24.341 30.609 3.067 25.092 4.430 Skeleton of chuck and neck 29.775 22.517 3.060 25.926 4.575 Skeleton of round (excl. marrow) 32.552 27.793 2.589 21.074 3.786 Marrow from skeleton of round 9 460 89.251 0.147 0.213 0.031 Skeleton of loin 7,772 5.192 25.056 31.376 2.935 24.006 4.277 Skeleton of rib 27.145 22.308 3.182 27.925 4.952 Hoofs and dewclaws 2.095 50.581 0.837 7.742 2.606 0 117 Teeth 852 21.329 1.162 2.079 61.007 11.516 Table 2. — Steer 501. Analysis of Samples. Description o f sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 28,710 77.977 0.176 3.290 0.857 0.025 Circulatory system 1,836 41.962 45.889 1.907 0.476 0.049 Lean Heart 1,882 77.616 3.738 2.573 0.992 0.198 Respiratory system 3,838 76.577 3.348 2.873 1.041 0.172 Fat from tnoracic cavity 2,459 18.834 76.645 0.612 0.237 0.026 Brain and spinal cord 757 70.404 13.274 1.673 1.836 0.392 Digestive and excretory system (partial) 24 235 71.756 12.870 2.143 0.809 0.123 Offal fat 38,625 7.488 91.061 0.205 0.102 0.012 Heart and neck sweetbreads 784 30.756 61.763 0.993 0.502 0.107 liver 6,161 69.513 2.898 3.233 1.423 0.334 Gall 176 91.952 0.050 0.213 1.229 0.034 Spleen 1,178 77.892 1 953 2.773 1.386 0.239 Pancreas 836 59.873 24.564 2.203 1.153 0.263 Kidneys 1,037 77.660 4.867 2.347 1.051 0.199 Tongue, marketable (excl. bones) 2,153 65.843 16.199 2.610 0.870 0.157 Hair and hide 50,090 51.432 13 235 5.493 1.522 0.049 Head and tail, lean and fat 5,224 60.421 20.746 2.833 0 767 0 126 Shin and shank, lean and fat 17,420 58.949 22.573 2.707 0.772 0 133 Flank and plate, lean and fat 134,146 26 818 65.884 1.050 0.342 0.057 Rump, lean and fat 22.226 28 769 62.760 1.140 0.395 0.069 Chuck and neck, lean and fat 110,990 47.698 38.429 1.525 0.652 0.118 Round, lean 50,130 69.902 9.356 3.090 0.957 0.185 Round, fat 22,284 16.846 78.237 0.667 0.218 0.026 Loin, lean 45,996 62.557 17.934 2.863 0.851 0.163 Loin, fat 7 1,35 8 9.031 88.682 0.388 0.112 0.018 40 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 2. — Steer 501. Analysis of Samples — Continued. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % 20,834 58.892 22.409 2.759 0.791 0.149 Rib, fat 28,322 9.697 87.439 0.401 0.134 0.020 Kidney, fat 19,544 5.462 93.311 0.190 0.067 0.011 Skeleton of feet 7,744 10,462 304 36.054 12 326 3.531 26 132 5.015 Skeleton of head 43.603 11.776 3.287 23.776 4.199 Skeleton of tail 40 880 19.185 3.481 18.055 3 155 Skeleton of shin 6,170 7,128 7,068 3,682 15,778 6,978 32.712 14.188 3.476 28 968 5 237 Skeleton of shank 26 943 22.187 3.348 27.861 5.124 Skeleton of flank and plate 40.213 15.662 3.320 18.801 3 403 Skeleton of rump 25.333 26.213 3.172 25.973 4 688 Skeleton of chuck and neck 30 . 106 15.495 3.656 28.381 5 205 Skeleton of round (excl. marrow) 26.646 24.351 3.086 27.253 4.948 Marrow from skeleton of round 286 10.169 88.390 0.222 0.530 0.084 Skeleton of loin 8,614 25.711 22.523 3.127 28.411 5.072 Skeleton of rib 6.388 28.428 18.371 3.322 27.868 5.181 Horns 3,354 36.989 0.633 6.469 22.743 4.167 Hoofs and dewclaws 2,523 778 47.011 0.658 8.453 1.715 0.143 Teeth 22.106 0.808 2.075 59.784 11.737 Table 3. — Steer 502. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 19,728 3,446 77.433 3.492 0.720 0.023 Circulatory system 68.285 14.513 2.586 0.679 0.138 Respiratory system 3,696 76.165 3.056 2.829 1.038 0.166 Fat from thoracic cavity 1,271 21.504 73.479 0.754 0.295 0.041 Brain and spinal cord 800 69.104 14.579 1.717 1.794 0.414 Digestive and excretory system (partial) 20,933 77.020 6.035 2.495 1.264 0.136 Offal fat 11,377 10.651 86.180 0.468 0.185 0.023 Heart andneck sweetbreads 502 59.778 25.414 2.244 1.153 0.270 Liver 3,716 68.941 1.728 3.305 1.391 0.334 Gall 241 93.020 0.043 0.208 1.027 0.029 Spleen 921 77.301 2.420 2.838 1.834 0.279 Pancreas 581 53,462 29.615 2.172 1.073 0.240 Kidneys 838 73.657 6.587 2.681 1.115 0.221 Hair and hide 39,556 57.299 3.010 6.574 0.976 0.059 Head and tail, lean and fat 4,250 64.599 14.122 3.201 0.826 0.149 Shin and shank, lean and fat 12,364 68.295 8.671 3.486 0.884 0.164 Flank and plate, lean and fat 35,594 51.905 31.222 2.583 0.696 0.119 Rump, lean and fat 8,100 56.028 25.426 2.611 0.797 0.147 Chuck and neck, lean and fat 70,744 66.333 12.728 3.072 0.921 0.158 Round, lean 44,426 72.153 4.051 3.306 0.971 0.195 Round, fat 4,620 26.602 62.837 1.549 0.322 0.038 Loin, lean 35,104 69.877 8.126 3.193 0.975 0.199 Loin, fat 9,144 15.490 78.182 1.047 0.249 0.036 Rib, lean 18,256 66.358 10.116 3.068 0.893 0.164 Rib, fat 3,338 20.655 69.994 1.484 0.281 0.051 Kidney, fat 2,916 7.388 89.667 0.420 0.241 0.039 Skeleton of feet 6,982 38.632 12.205 3.788 23.868 4.240 Skeleton of head 9,577 48.757 8.378 3.185 20.196 3.460 Skeleton of tail 441 40.425 22.573 3.251 15.075 2.733 Skeleton of shin 5,490 27.399 19.319 3.532 29.466 5.252 Skeleton of shank 5,978 27.197 24.585 3.454 25.363 4.505 Skeleton of flank and plate 5,590 41.156 12.997 3.382 21.818 3.861 Skeleton of rump 2,362 25.436 29.734 3.026 23.684 4.232 Skeleton of chuck and neck 15,092 30.767 18.259 3.407 27.553 4.918 Skeleton of round (excl. marrow) 6,296 26.477 26.444 2.996 26.259 4.681 Marrow from skeleton of round 408 7.876 90.632 0.202 0.413 0.074 Skeleton of loin 7,866 26.290 26.514 3.149 25.839 4.569 Skeleton of rib Horns* 5,290 1 949 30.433 20.599 3.647 24.864 4.401 Hoofs and dewclaws 2,010 58.684 0.572 6.625 1.186 0.120 Teeth 1,038 36.211 1.025 1.632 50.004 9.483 •This sample was lost before analysis. Studies In Animal Nutrition — III 41 Table 4. — Steer 503. Analysis of Samples. Description of sample Weight in animal , grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 13,058 82.780 0.110 2.708 0.336 0.076 Circulatory system 1,782 36.244 54.914 1.249 0.329 0.060 Lean heart 1.063 78.139 3.736 2.591 0.951 0.207 Respiratory system 2,549 78.706 3.155 2.654 0.976 0.207 Brain and spinal cord 666 73.106 16.109 1.682 1.549 0.394 Digestive and excretory system (partial) .... 8,761 72.710 11.013 2.375 1.012 0.208 Offal fat 7,385 14.642 81.920 0.521 0.182 0.035 Liver 3,646 68.680 5.266 2.995 1.284 0.334 Kidneys 655 71.360 11.795 2.414 1.041 0.225 Stomach 5,765 77.658 6.925 2.211 1.085 0.207 Tongue, marketable 789 69.328 13.263 2.535 0.827 0.170 Hair and hide 23,008 67.765 2.570 4.793 0.979 0.068 Shin, snank, head and tail, lean and fat 8.614 69.707 10.223 3.196 0.847 0.168 Flank and plate, lean and fat 16,290 56.518 25.666 2.742 0.748 0.136 Chuck and neck, lean and fat 31,934 6S.526 12.574 2.954 0.870 0.171 Round and rump, lean 25.022 73.087 4.805 3.370 1.024 0.204 Round and rump, fat 3,400 22.450 70.130 1.212 0.287 0.048 Loin, lean 17,206 70.994 8.451 3.187 0.983 0.190 Loin, fat 5.746 15.748 79.917 0.777 0.190 0.035 Rib, lean 8,932 69.677 10.231 3.154 0.927 0.185 Rib, fat 840 23.294 66.936 1.481 0.318 0.061 Kidney, fat 2,126 8.676 89.467 0.340 0.125 0.027 Skeleton 41,122 38.277 15.059 3.104 23.704 4.378 Horns, hoofs and dewclaws 1.058 46.286 2.032 7.576 6.055 0.661 Teeth 253 23.299 0.525 2.252 58.876 11.280 Table 5. — Steer 504. analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen 07 / O Ash % Phosphorus % Blood 21,005 1,637 78.650 3.334 0.387 0.405 0.022 0.041 Circulatory system 28.710 63.730 1.112 Lean heart 1.428 76.650 4.310 2.774 1.015 0.203 Respiratory system 3,628 67.410 14.900 2.882 0.974 0.181 Brain and spinal cord 724 69.230 15.080 1.788 1.358 0.350 Digestive and excretory system (partial) 17,569 68.540 18.040 1.910 0.726 0.145 Offal fat 25,105 12.760 84.820 0.334 0.147 0.023 Liver 4,754 69.220 2.436 3.275 1.352 0.358 Kidneys 877 69.480 12.810 2.452 1.058 0.219 Stomachs 12,820 79.670 8.040 1.701 0.897 0.151 Tongue, marketable 1,587 60.750 23.480 2.182 0.759 0.141 Hair and hide 41,144 58.290 8.070 5.522 1.057 0.043“ Shin, shank, head and tail, lean and fat 16,070 60.640 20.560 2.951 0.803 0.14$ Flank and plate, lean and fat 49.650 41.640 45.620 1.945 0.572 0.101 Rump, lean and fat 10,846 40.720 46.810 1.847 0.574 0.106 Chuck and neck, lean and fat 59,808 58.330 24.030 2.620 0.758 0.146 Round, lean 37,238 69.510 9.210 3.208 0.983 0.194 Round, fat 9,818 16.610 78.030 0.906 0.238 0.030 Loin lean 33 676 66.920 12.220 3.051 0.946 0.181 Loin, fat 18.340 11.620 84.910 0.532 0.162 0.025 Rib, lean 18,506 63.280 17.520 2.940 0.831 0.167 Rib, fat 6,770 14.420 80.630 0.833 0.202 0.031 Kidney, fat 11,400 4.800 93.940 0.215 0.126 0.017 Skeleton of feet, head, tail, shin and shank.. 23,568 36.050 13.640 3.237 27.594 5.076 Skeleton of flank and plate 4,572 44 . 100 18.180 3.523 15.715 2.916 Skeleton of rump 2,428 25.720 26.000 3.073 27.381 5.184 Skeleton of chuck and neck 11,176 30.180 16.270 3.524 29.162 5.344 Skeleton of round 4,808 21.880 27.470 3.120 31.049 5.755 Skeleton of loin 5,850 29.840 22.100 3.184 26.452 4.812 Skeleton of rib 5.092 32.550 16.260 3.417 27.791 5.096 Horns, noofsand dewclaws Teeth* 2,532 338 69.476 0.514 4.621 2.428 0.157 1 •This sample was lost before analysis. 42 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 6. — Steer 505 . Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 13,810 82.260 0.351 2.726 0.329 0.023 Circulatory svstem 1,168 33.230 59.064 1.285 0.277 0.047 Lean heart 938 77.382 5.110 2.618 0.973 0.209 Respiratory system 2,498 76.829 5.477 2.713 0.951 0.202 Brain and spinal cord 537 73.810 14.738 1.688 1.719 0.411 Digestive and excretory system (partial) 8,258 71.893 12.815 2.476 1.053 0.226 Offal fat 12,781 12.410 85.384 0.340 0.117 0.022 Liver 3,983 68.096 5.770 3.205 1.329 0.347 Kidneys 718 75.712 7.835 2.376 1.061 0.226 Stomach 8,818 77.261 10.849 1.681 0.868 0.173 Tongue, marketable 1,115 64.061 20.160 2.352 0.759 0 156 Hair and hide 22,884 62.138 5.336 5.297 0.696 0.066 Shin, shank, nead and tail, lean and fat 9,386 64.430 15.398 3.216 0.818 0.165 Flank and plate, lean and fat 24,194 43.720 42.762 2.211 0.580 0.116 Chuck and neck, lean and fat 38,344 62.294 18.949 2.886 0.830 0.165 Round and rump, lean 25,784 69.066 9.464 3.326 0.976 0.200 Round and rump, fat 5,970 14.140 80.640 0.452 0.174 0.032 Loin, lean 19,686 68.190 9.983 3.236 0.963 0.196 Loin, fat 7,558 9.333 87.547 0.535 0.127 0.024 Rib, lean 11,300 61.762 19.191 2.964 0.841 0.176 Rib, fat 2.640 10.913 85.385 0.643 0.167 0.033 Kidney, fat 5,754 5.263 93.527 0.236 0.084 0.016 Skeleton 37,745 35.792 17.555 3.186 23.852 4.403 Horns, hoofs and dewclaws 1,206 46.098 1.104 7.724 5.340 0.611 Teeth 268 21.938 0.634 2.264 59.386 10.697 Table 7 . — Steer 507 . Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat 0/ /o Nitrogen % Ash % Phosphorus % Blood 20,316 78 . 174 3.431 0.670 0.022 Circulatory system 4,278 47.118 41.273 1.695 0.535 0.104 Respiratory system 3,768 77.467 3.927 2.729 0.954 0.169 Brain and spinal cord 744 70.806 13.738 1.498 1.749 0.418 Digestive and excretory system 27,417 70.515 14.079 2.097 0.825 0.150 Offal fat 11,037 13.096 83.962 0.412 0.140 0.029 Hair and hide 34,473 60.845 6.213 5.688 1.085 0.049 Head and tail, lean and fat 4,038 62.582 18.949 2.841 0.895 0.161 Shin and snank. lean and fat 11.860 69.883 7.860 3.343 0.922 0.167 Flank and plate, lean and fat 36,130 51.939 32.362 2.504 0.693 0.124 Rump, lean and fat 7,736 50.803 31.644 2.275 0.721 0.139 Chuck and neck, lean and fat 62,530 65.569 15.166 2.891 0.862 0.157 Round, lean 39,302 72.727 5.772 3.230 0.981 0.192 Round, fat 5,378 24.446 68.671 1.093 0.276 0.039 Loin, lean 29,724 70.696 8.096 3.128 0.972 0.185 Loin, fat 10,188 17.822 76.684 0.826 0.223 0.039 Rib, lean 15,788 67.438 12.239 2.999 0.937 0.174 Rib, fat 2,432 17.554 76.436 0.947 0.253 0.042 Kidney, fat 4,376 6.784 91.300 0.284 0.147 0.025 Skeleton of feet, head and tail 15,275 42.804 11.776 3.456 23.906 3.858 Skeleton of snin and snank 10,350 23.960 19.357 3.646 33.717 4.634 Skeleton of flank and piate 6,278 44.217 13.485 3.380 18.472 2.851 Skeleton of rump 2,536 25.041 26.339 3.229 25.836 3.940 Skeleton of chuck and neck 13,202 31.983 15.245 3.555 25.644 4.204 Skeleton of round 5,864 26.093 29.961 3.142 23.204 4.161 Skeleton of loin 6,506 26.630 26.891 2.866 26.102 3.724 Skeleton of rib 5,050 28.673 18.021 3.363 28.739 4.282 Horns 1,600 41.605 0.624 5.762 21.877 3.960 Hoofs and dewclaws 1,490 54.438 1.143 7.002 2.044 0.163 Teetn 712 26.534 1.027 1.964 56.717 10.889 Studies In Animal Nutrition — III 43 Table 8. — Steer 509. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % 18.291 78.053 3.429 0.686 0.023 2.616 69 . 945 12.261 2.590 0.922 0.167 3'283 1.248 77.032 2.879 3.004 1.011 0.176 19.808 76.095 0.720 0.297 0.043 739 66.874 17.662 1.679 1 576 0 379 Digestive and excretory svstem (partial) Offal fat 19,016 9.922 630 73.011 11.330 11.886 85.888 2.206 0.534 0.908 0.175 0.123 0.024 50.304 37.140 1.816 1.039 0.254 3,875 110 68.408 1.757 3.088 1.334 0.326 Gall 91.529 0.059 0.250 0.874 0.031 1,304 77.255 2.126 2.993 1.420 0.242 54.915 28.448 2.239 1.178 0.269 774 77.013 3.823 2.649 1.115 0.234 37,614 58.969 2.477 6.270 1.024 0.046 Head and tail, lean and fat 3,212 11,782 31,790 66.769 13.022 3.077 0.821 0 150 Shin and shank, lean and fat 67.912 9.765 3.380 0.911 0.167 Flank and Dlate, lean and fat 53.639 28.926 2.620 0.733 0.145 Rump, lean and fat 7,370 55.813 26.599 2.628 0.804 0.146 Chuck and neck, lean and fat 60.176 68.493 9.856 3.076 0 895 0.166 Round, lean 40.376 73.647 4.183 3.223 1.005 0.196 Round, fat 5.106 25.559 64.040 1.621 0.343 0.041 Loin, lean 30,836 69.632 9.010 3.103 0.951 0.180 Loin, fat 7,570 16,360 16.259 77.561 1.033 0.253 0.041 Rib, lean 67.795 10.844 2.927 0 915 0.170 Rib, fat 1,978 1,576 6,144 17.552 76.022 1.058 0.279 0.045 Kidney, fat 5.466 92.412 0.314 0.138 0.017 Skeleton of feet 41.075 10.778 3.669 23.457 4.245 Skeleton of head 8,247 47.540 8.478 3.176 21.164 3.772 Skeleton of tail 386 37.947 26.555 2.873 15.105 2 . 734 Skeleton of shin 5,046 5,498 5,124 2.860 13,682 28.351 18.203 3.839 29.339 5.168 Skeleton of shank 28.550 22.558 3.519 26.144 5.027 Skeleton of dank and plate 41.367 13.032 3.533 22.382 4 082 Skeleton of rump Skeleton of cnuck and neck 26.895 32.387 29.745 17.293 3.058 3.704 22.246 25.046 3.878 4.606 Skeleton of round (excl. marrow) 5,442 578 27.956 24.660 2.967 25.948 4.624 Marrow from skeleton of round 11.658 86.848 0.203 0.370 0.058 Skeleton of loin 6.872 27.517 25.661 3.130 25.647 4.414 Skeleton of rib 4,986 1,590 28.050 16.791 3.623 30.569 5.444 Hoofs and dewclaws 66.980 0.459 5.193 1.457 0.117 Teeth 838 28.535 0.535 1.740 56.001 10.533 Table 9. — Steer 512. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 24,176 79.949 0.055 3.073 0.790 0.023 Circulatory system 2.432 40.529 47.983 1.710 0.563 0.055 Lean heart 1,555 77.321 3.470 2.803 1.269 0.215 Respiratory system 3.881 75.487 4.670 2.803 1.079 0.164 Fat from thoracic cavity 1.171 12.232 85.522 0.263 0.144 0.015 Brain and spinal cord 666 72.058 11.117 1.687 1.871 0.385 Digestive and excretory system (partial) 20,735 73.684 10.288 2.183 0.761 0.119 Offal fat 17,454 11.212 86.273 0.369 0.144 0.019 Heart and neck sweetbreads 511 40.399 49.421 1.497 0.733 0.168 Liver 4.416 68.984 2.625 3.263 1.588 0.334 Gall 212 93.144 0.033 0.223 1.039 0.028 Spleen 1,255 77.145 2.366 2.800 1.339 0.239 Pancreas 736 57.391 26.641 2.087 1.326 0.274 Kidneys 1.074 77.235 6.831 2.080 1.053 0.203 Tongue, marketable 1,766 66.379 13.428 2.580 0.899 0.161 Hair and hide 41,268 56 193 3.612 6 . 547 1.163 0 047 Head and tail, lean and fat 4.412 61.554 19.074 2.893 0.859 0.139 Shin and shank, lean and fat 12,706 68.606 9.380 3.343 0.927 0.161 Flank and plate, lean and fat 48,946 41.914 45.337 1 910 0.570 0 094 Rump, lean and fat 10,484 44.598 41.829 2.027 0.667 0.119 Chuck and neck, lean and fat 73,512 63.188 18.191 2.826 0.919 0.150 Round, lean 43,408 73 272 4.557 3.237 1.024 0 192 Round, fat 9.940 22.030 70.658 0.765 0.311 0 040 Loin, lean 32.062 67.607 11.040 3.076 0.912 0.170 Loin, fat 15.308 12.497 83.354 0.650 0 130 0 026 Rib, lean 16,908 65 119 14.950 2.967 0.896 0.157 Rib, fat 5,398 14.938 80.367 0.840 0.212 0.035 44 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 9. — Steer 512. Analysis of Samples — Continued. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorui % Kidney, fat 4,740 4.482 93.915 0.183 0.130 0.020 Skeleton of feet 7,016 37.452 14.314 3.597 24.365 4.405 Skeleton of head 9,665 43.142 12.955 3.229 22.420 4.204 Skeleton of tail 416 36.970 24.279 3.129 18.107 3.265 Skeleton of shin 6,074 27.159 20.808 3.631 29.984 5.388 Skeleton of shank 6,156 31.979 21.541 3.540 22.768 4.094 Skeleton of flank and plate 7,788 36.792 21.061 3.033 19.564 3.586 Skeleton of rump 3,264 23.678 30.680 2.986 26.274 4.446 Skeleton of chuck and neck 16.536* 28.775 18.986 3.244 30.510 5.438 Skeleton of round (excl. marrow) 7,430 28.723 26.734 2.822 22.605 4.058 Marrow from skeleton of round 396 10.084 88.297 0.186 0.657 0.132 Skeleton of loin 8,748 25.136 24.403 2.987 26.642 5.334 Skeleton of rib 6,938 29.436 16.324 3.475 28.699 5.454 Horns 1,810 35.228 0.480 7.033 22.746 3.874 Hoofs and dewclaws 1,724 48.902 0.588 7.857 2.791 0.124 Teeth 710 19.913 0.782 2.152 63.721 12.031 Table 10. — Steer 513. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phospnorus % Blood 25,680 77.676 3.341 0.722 0 028 Circulatory system 3,485 77.600 4.500 2.673 0.909 0.157 Respiratory system 4,858 73.905 7.032 2.742 0.992 0.155 Fat from thoracic cavity 3,942 13.440 83.707 0.366 0.185 0.024 Brain and spinal cord 748 68.192 15.496 1.724 1.601 0.410 Digestive and excretory system (partial) 25,642 75.309 7.697 2.310 1.041 0.157 Offal fat 53,771 5.683 92.874 0.215 0.079 0.011 Heart and neck sweetbreads 1,334 40.327 50.545 1.405 0.824 0.198 Liver 5,920 67.926 3.126 3.225 1.397 0.333 Gall 37 91.391 0.146 0.312 1.191 0.035 Spleen 1,114 75.645 4.542 2.932 1.253 0.242 Pancreas 873 50.118 34 . 657 1.868 1 088 0 242 Kidneys 1,015 76.071 5.802 2.503 1.133 0.220 Hair and hide 45,286 57.566 10.072 5.178 0.941 0.054 Head and tail, lean and fat 5,390 56.379 26.291 2.439 0.770 0.128 Shin and shank, lean and fat 17,798 55.759 27.657 2.309 0.697 0.118 Flank and plate, lean and fat 115,774 29.221 62.259 1.242 0.385 0.065 Rump, lean and fat 19,082 31.575 58.755 1.378 0.438 0.080 Chuck and neck, lean and fat 110,940 47.879 37.191 2.152 0.629 0.116 Round, lean 50,782 65.159 14.337 2.901 0.911 0.173 Round, fat 19,108 17.763 76.089 0.921 0.199 0.024 Loin, lean 44,510 59.396 21.597 2.777 0.826 0.163 Loin, fat 49,928 8.901 88.492 0.366 0.115 0.018 Rib, lean 24,744 55.589 27.713 2.567 0.757 0.142 Rib, fat 23,608 17.797 75.455 0.408 0.138 0.021 Kidney, fat 14,490 3.912 94.928 0.156 0.074 0.010 Skeleton of feet 7,598 36.238 14.856 3.522 23.836 4.243 Skeleton of head 7,865 40.707 9.440 3.300 25.028 5.856 Skeleton of tail 297 39.282 19.765 3.353 17.252 3.031 Skeleton of shin 6,120 29.290 17.835 3.377 27.325 4.811 Skeleton of shank 6,058 26.124 19.919 3.570 29.591 5.233 Skeleton of flank and plate 7,438 40.922 15.081 3.278 20.425 4.139 Skeleton of rump 3,464 24.417 29.288 3.009 24.688 4.683 Skeleton of chuck and neck 15,526 32.008 17.781 3.508 25.202 5.058 Skeleton of round (excl. marrow) 6,564 24.756 30.638 2.968 24.757 4.366 Marrow from skeleton of round 480 8.660 89.964 0.161 0.811 0.139 Skeleton of loin 8,536 25.186 29.799 3.025 24.383 4.699 Skeleton of rib 7,096 26.559 21.539 3.363 29.216 5.528 Horns 2,144 36.851 0.766 6.450 22.622 4.168 Hoofs and dewclaws 2,180 41.930 0.890 8.956 2.960 0.239 Teeth 874 31.897 1.103 1.811 52.576 9.959 Studies In Animal Nutrition — III 45 Table 11. — Steer 515. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % 27,856 5,787 3,653 728 79.107 3.217 0.594 0.024 42.032 48.530 1.276 0.480 0.086 76.354 5.443 2.699 0.968 0.187 70.038 16.091 1.654 1.634 0.395 36.311 66.167 18.711 2.019 1.851 0.162 29,877 7.532 90.291 0.281 0.124 0.015 49^943 5,918 56.003 12.550 4.840 1.857 0.059 Head and tail, lean and fat 53.562 30.538 2.417 0.731 0.130 Shin and shank, lean and fat 16,676 57.361 26.753 2.502 0.716 0.123 Flank and plate, lean and fat 87,138 15,810 31.118 59.929 1.360 0.388 0.071 Rump, lean and fat 35.336 54.204 1.506 0.476 0.085 Chuck and neck, lean and fat 88,134 53.265 31.088 2.356 0.712 0.126 Round, lean 42,942 67.691 11.052 3.102 0.924 0.177 Round, fat 19,058 41,620 38,324 19,016 17.522 77.696 0 784 0.215 0.026 Loin, lean 64.691 15.193 2.999 0 994 0.181 Loin fat 10.576 86.623 0 412 0 115 0.017 Rib, lean 61.183 20.387 2.776 0.849 0.152 Rib, fat 16.282 9.071 88.287 0.394 0.134 0.020 Kidney, fat 9,922 4.951 94.433 0.178 0.081 0.015 Skeleton of feet, head and tail 18,179 13.900 41.368 11.161 3.032 23.490 4.066 Skeleton of shin and shank 27.713 27.063 3.045 24.670 4.032 Skeleton of flank and plate 6,368 4,074 42.196 15.964 3.410 17.779 3.113 Skeieton of rump 29.793 25 . 690 3.096 23.313 4.071 Skeleton of chuck and neck 14,528 6,344 7,784 6,464 1.804 30.500 15.517 3.333 29.568 4.253 Skeleton of round 23 . 643 29.413 3.024 28 . 767 4.890 Skeleton of loin 25.307 28.219 2.971 26.216 4.497 Skeleton of rib 26.086 20.159 3.270 31.176 4.374 Horns 40 . 640 0.591 5.617 23.674 4.223 Hoofs and dewclaws 1,893 53.752 0.529 7.363 1.823 0.072 Teeth 786 27.331 0.859 1.785 57.132 10.992 Table 12. — Steer 523. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 15,287 3,044 3,371 797 80.520 53 054 2.790 0.659 0.598 0.022 Circulatory system 35.823 1.684 0.110 Respiratory system 78 . 769 4.054 2.605 0.980 0 189 Brain and spinal cord 68.595 17.610 1.610 1.515 0 359 Digestive and excretory system 24,667 75.194 10.330 2.190 0.788 0.158 Hair and nide 33.097 62.801 1.150 5.603 1.030 0.051 Head and tail, lean and fat 3,100 68.166 11.573 3.094 0.923 0 . 154 Shin and shank, lean and fat 8,684 71.786 6 038 3.402 0.915 0.169 Flank and plate, lean and fat 26,984 59.510 22 . 920 2.693 0.739 0.139 Rump, lean and fat 5.418 54.330 29.269 2 532 0.748 0.154 Cbuck and neck, lean and fat 50,320 33,900 25,834 12.032 70 884 10 288 2 980 0 921 0.171 Round, lean 76.929 2.310 3 155 1.042 0.202 Loin, lean 69 . 754 10 406 3.027 0 921 0.172 Rib, lean 70.282 9.297 3.119 0.927 0.178 Round, fat 4,556 6,376 1,522 29.544 60 224 1.580 0 357 0.048 Loin, fat 16.497 77.934 0.863 0.281 0.394 0.045 Rib, fat 23 . 543 64 . 977 1.511 0.059 Kidney, fat 3 110 5 259 92 726 0 . 465 0.179 0.015 Offal, fat 7,915 13,120 15.418 44.251 81.284 8 120 0 495 0 204 0.030 Skeleton of feet, head and tail 3 563 22.796 4.135 Skeleton of shin and shank 8,862 3.882 29 . 464 20 364 3 381 27 . 035 5.132 Skeleton of flank and plate 44 868 10.710 3 471 19.443 3.467 Skeleton of rump 1,770 9,876 27.036 20 . 259 3.270 29 591 5.430 8keleton of cbuck and neck 33 . 866 12 894 3.681 29.394 5.391 Skeleton of round 4.630 39 . 850 26.599 1.942 20.522 3 840 Skeleton of loin 5,322 3,844 1,167 1,063 766 27.873 23 . 686 3.145 27.029 4 981 Skeleton of rib 31 218 12 517 3 592 31 713 5 769 Horns 46.232 0.625 6.616 18.740 3.410 Hoofs and dewclaws* Teeth 26.317 0.703 2.098 50.395 10.670 •This sample was lost before analysis. 46 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 13. — Steer 524. Analysis of Samples. Description of sample Weight in animal , grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 17,019 82.006 2.787 0.746 0 022 Circulatory system 2,953 62.090 22.418 2.240 0.768 0.132 Respiratory system 3,455 77.635 2.601 2.793 1.073 0.178 Brain and spinal cord 758 73.218 10.255 1.597 1.418 0.338 Digestive and excretory system (partial) 17,924 74.094 9.821 2.303 0.885 0.150 Offal fat 5,007 25.069 69.444 0.760 0.320 0.035 Heart and neck sweetbreads 541 75.123 7.448 2.603 1.873 0.463 Liver 3.019 69.840 3.316 3.047 1.408 0.327 Gall 229 94.859 0.092 0.167 0.794 0.031 Spleen 757 78.752 1.819 2.750 1.335 0.260 Pancreas 435 65.634 15.877 2.540 1.230 0.285 Kidneys 766 76.830 5.735 2.433 1.158 0.208 Hair and hide 30,092 59.259 1.813 6.300 1.577 0.058 Head and tail, lean and fat 3,364 66.261 13.126 3.127 1.026 0.179 Shin and shank, lean and fat 8,674 73.037 5.601 2.925 0.974 0.166 Flank and plate, lean and fat 19,788 63.711 15.372 3.200 0.952 0 154 Rump, lean and fat 4,030 63.611 17.147 2.957 0.972 0.175 Chuck and neck, lean and fat 46,386 72 555 6 292 3 160 0 975 0 174 Round, lean 37,714 76.864 2.716 3.257 1.043 0.192 Round, fat 2,526 36.260 49.962 2.237 0.420 0.053 Loin, lean 24,200 72.680 4.544 3.310 1.057 0.194 Loin, fat 2,444 18.811 73.319 1.237 0.350 0.053 Rib, lean and fat 13,144 70.285 7.949 3.310 0.982 0.182 Kidney, fat 766 11.440 84.187 0.440 0.179 0.031 Skeleton of feet 6,010 40.508 13.002 3.592 24.515 4.375 Skeleton of head and tail 8.318 45.930 10.389 3.053 23.867 4.263 Skeleton of shin and shank 10,262 31.581 19.317 3.365 25.920 4.619 Skeleton of flank and plate 5,926 43.358 15.272 3.211 19.164 3.345 Skeleton of rump 2,424 34.304 21.452 2.997 22.395 4.057 Skeleton of chuck and neck 12,896 40.299 15.844 3.316 21.786 3.986 Skeleton of round 5,878 31.056 28.341 2.666 22.700 4.063 Skeleton of loin 6,586 29.252 26.901 2.845 24.306 4.246 Skeleton of rib 5,310 35.735 18.535 2.969 24.951 4.429 Horns* 1,227 Hoofs and dewclaws 1,494 50.239 0.832 7.553 3.194 0.219 Teeth 806 26.967 1.106 1.907 57.863 10.988 •Tiiis sample was lost before analysis. Table 14. — Steer 525. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 13,614 80.514 2.942 0.660 0.026 Circulatory system 1,320 1,658 711 64.087 24.542 1.705 0.715 0.121 Respiratory system 78.612 2.633 2.823 1.111 0.193 Brain and spinal cord 71.054 13.931 1.737 1.504 0.370 Digestive and excretory system 23,001 77.479 9.144 1.999 0.883 0.156 Hair and hide 27,813 2,788 7,596 18,762 4,154 35,824 62.396 0.498 5.724 1.331 0.157 Head and tail, lean and fat 74.319 13.082 1.858 0.986 0.158 Shin and shank, lean and fat 72.849 5.330 3.337 0.965 0.177 Flank amd plate, lean and fat 60.827 20.204 2.873 0.828 0.149 Rump, lean and fat 60 . 691 20.925 2.760 0.864 0.162 Chuck and neck, lean and fat 71.151 8.443 3.125 0.949 0.178 Round, lean 27,524 18,710 77.034 2.403 3.120 1.064 0.205 Loin, lean 74.634 3.469 3.237 1.048 0.200 Rib, lean 11,666 70.514 8.659 3.161 0.977 0.181 Round, fat 1,962 3,758 32.685 56.826 1.599 0.433 0.056 Loin, fat 22.002 69.589 1.099 0.294 0.040 Rib, fat 664 28.087 61.613 1.611 0.605 0.092 Kidney, fat 1,258 4,961 10,782 7,014 6.721 90.181 0.470 0.161 0.020 Offal , fat 21.028 70.641 1.286 0.293 0.051 Skeleton of feet, head and tail 43.021 9.990 3.736 21.822 4.171 Skeleton of shin and shank 30.333 19.135 3.485 29.526 4.190 Skeleton of flank and plate 3,454 1,542 44.613 10.853 3.365 21.567 2.989 Skeleton of rump 30.800 23.819 3.234 24.612 4.216 Skeleton of chuck and neck 8,450 34 . 145 19.714 3.092 22.900 4.301 Skeleton of round 4,046 3,928 3,888 28.602 32.687 2.653 21.783 2.826 Skeleton of loin 28.594 25.309 3.020 24.434 3.678 Skeleton of rib 31.735 18.050 3.593 24.740 4.352 Horns 1,298 940 48.873 0.543 5.641 16.718 3.141 Hoofs and dewclaws 51.906 0.575 7.892 1.259 0.134 Teeth 690 23.125 1.063 1.903 60.713 11.737 Studies In Animal Nutrition — III 47 Table 15 . — Steer 526 . Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphoru 3 % 18,957 79.926 3.087 0.777 0.025 2,413 3,797 1,585 660 63.316 21.434 2.343 0.814 0.145 76.615 3.288 2.800 1.025 0.156 21.467 73.139 0.753 0.286 0.034 73.217 10.255 1.747 1.728 0.420 Digestive and excretory system (partial) 20,541 11.551 72.884 12.992 12.222 84.144 2.130 0.410 0.768 0.175 0.127 0.018 439 60.467 24.213 2.253 1.444 0.348 3,531 67 . 922 4.255 3.247 1.469 0.348 Gall 143 92.398 0.138 0.230 1.158 0.031 831 78.490 1.496 2.767 1.554 0.297 498 63.457 18.209 2.245 1.172 0.282 922 73.557 9.071 2.353 1.069 0.198 35,732 57.828 5.714 5.923 1.440 0.050 3.616 61.791 19.640 2.910 0.832 0.142 11,644 39,524 70.950 7.537 3.301 0 900 0.164 49.492 35.249 2.330 0.678 0.125 8,594 53.663 29.882 2.463 0.738 0.138 61,228 65.441 14.500 2 923 0 907 0 166 44,614 66.884 11.887 3.300 1.031 0.191 5.016 22.666 63.981 1.670 0.284 0.036 31,440 71.948 5.341 3.210 1.018 0.190 11.634 17,264 14.508 79.929 0.850 0.208 0.030 69.641 10.014 3.077 0.920 0.165 3,720 17.241 74.768 1.160 0.259 0.044 3,224 9.074 87.831 0.340 0.137 0.021 6.138 38.425 13.853 3.682 23.475 4.354 9.165 46.363 12.805 2.998 21.004 3.832 11,612 6.588 27.802 20.097 3.254 28 . 703 5.238 40 . 154 16 466 3.175 20.333 3 793 2!838 13,842 29.344 25 608 3.108 23.706 4.277 34.313 17.449 3.339 24.601 4.482 6,352 6.844 27.057 30.272 2.694 23.272 4.318 29.163 28.333 2.826 23.314 4.282 Skeleton of rib 5,690 1,427 1,875 31.712 20.075 3.314 26.287 4.758 Horns 41.315 0.744 6.411 19.354 3.592 Hoofs and dewclaws 54.392 0.625 7.294 2.123 0.085 Teeth 782 22.147 1.273 1.941 61.470 11.686 Table 16 . — Steer 527 . Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 27,382 4.903 78.941 3.283 0.762 0.023 Circulatory system 53.951 32.798 1.927 0.635 0.114 Respiratory system 4,326 3.988 73.330 7.186 2.717 1.024 0.166 Fat from thoracic cavity 10.340 87.646 0.335 0.128 0.015 Brain and spinal cord 701 69.957 14.059 1.583 1.544 0.368 Digestive and excretory system (partial) Offal fat 23,818 48,517 1,037 5,720 1,226 66.822 5 . 394 18.527 93 . 260 2.077 0.183 0.710 0.118 0.119 0.013 Heart and neck sweetbreads 36.622 55.347 1.187 0.734 0.308 Liver 67.872 3 472 3 293 1.575 0 184 Spleen 76.790 2.169 2.903 1.282 0.237 Pancreas 849 41.574 46.296 1.460 0.887 0.203 Kidneys 1,244 46.240 75.345 8.280 2.210 1.001 0.193 Hair and hide 54.475 11.859 5.317 1.369 0.056 Head and tail. lean and fat 5,018 53.921 30 . 635 2.410 0.691 0.123 Shin and shank, lean and fat 17,358 56.503 26.398 2.570 0.754 0.130 Flank and plate, lean and fat 118.978 27.200 65.491 1.097 0.312 0.059 Rump, lean and fat 24,020 29 026 62.962 1.233 0.315 0.059 Chuck and neck, lean and fat 112.440 46.782 39.316 2.030 0.617 0.109 Round, lean 51.396 66.109 13.931 2.993 0.864 0.175 Round, fat 21,466 50.140 16.129 79.323 0.753 0.204 0.022 Loin, lean 61 192 20.496 2.797 0.849 0.161 Loin, fat 52,724 9.519 88.271 0.357 0.127 0.017 Rib, lean 25,860 54 . 984 28.098 2.547 0.737 0.141 Rib, fat 24.278 9.463 88 . 208 0.407 0.133 0.016 Kidney, fat 18.964 5.423 93.227 0.187 0.102 0.014 Skeleton of feet 7,442 8 822 37.280 16.115 3.372 23 . 664 3.849 Skeleton of head and tail 42 136 24.822 3.165 21.749 3.076 Skeleton of shin and shank 13,136 6,082 27 . 256 21 . 276 3.149 28.369 4 550 Skeleton of flank and plate 39.334 18.437 2.996 19.531 3.404 Skeleton of rump 3.260 24 846 30 . 690 2.967 23.925 3.747 Skeleton of chuck and neck 14,870 28.058 24.570 3.082 25.037 3.944 Skeleton of round 6,446 20.998 33 . 954 2.642 27.032 4.964 Skeleton of loin 7,140 25.920 25.782 3.151 20.949 5.156 Skeleton of rib 6,546 26.920 23.074 3.022 28 . 622 5.503 Homs 1,266 35 . 662 0.775 6.950 19.811 3 . 658 Hoofs and dewclaws 2.174 44.082 0.959 8.916 2.059 0.145 Teeth 872 20.697 1.438 1.954 63.378 11.767 48 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 17. — Steer 531. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % A B h % Phosphorus % 9.457 81.882 2.841 0.703 0.040 0.126 0.204 0.363 0.121 0.046 0.427 0.321 0.052 0.291 0.288 0.213 0.075 0.155 0.188 0.176 0.177 1,971 56.721 29.318 1.968 0.676 1,915 573 78.338 2.818 2.680 1.054 72.987 13 498 1.765 1 502 Digestive and excretory system (partial) Offal fat 11,807 2,899 77.129 25.931 9.230 70.680 1.977 0.621 0.669 0.290 472 67.988 14.791 2.497 1.720 2,205 86 71.785 1.945 2.921 1 385 Gall 91.678 0.214 0 978 481 75.079 4.195 2.958 1 440 Pancreas 297 67.078 14.763 2.508 1.229 Kidneys 506 75.079 8.283 2.265 1.042 Hair and nide 16.693 63.665 0.811 5.733 1.136 Head and tail, lean and fat 1,722 5.480 10.854 67.218 14.073 2.923 0 872 Shin and shank, lean and fat 71.792 4.736 3.596 1 071 Flank and plate, lean and fat 60.374 17.191 3.055 1.005 Rump, lean and fat 2,506 61.990 18.113 2.980 0.976 Chuck and neck, lean and fat 26,902 21,496 1,490 14078 69.914 7.805 3.274 1.034 0 196 Round, lean 75.102 1.831 3.272 1.102 0.208 0.071 Round, fat 28.477 60.684 1.524 0.478 Loin, lean 73.011 4.278 3.312 1.100 0.204 Loin, fat 2.264 24.756 64.861 1.386 0.439 0.075 Rib, lean and fat 6 612 69.536 7.559 3.338 1.052 0.190 Kidney fat 726 6.066 90.265 0.596 0.240 0.031 Skeleton of feet 3,762 4.842 39.773 14.354 3.233 24.642 4.432 Skeleton of head and tail 51.145 8.071 3.041 20.849 3.629 Skeleton of shin and shank 5.998 31.730 18.346 2.937 27.416 5.113 Skeleton of flank and plate 2,546 46.771 10.571 3.220 20.395 3.464 Skeleton of rump 1.060 31.418 21.238 3.274 25.533 4.573 Skeleton of chuck and neck 6,098 37.493 16.015 3.536 23 . 526 4.187 Skeleton of round 3.640 35.030 25.913 2.679 21.363 3 806 Skeleton of loin 2.834 31.090 21.915 3.298 25.494 4 471 Skeleton of rib 2.280 32.384 16.742 3.738 25.229 4 587 Hoofs and dewclaws 760 52 . 103 0.827 7.587 1.963 0.125 Teeth 426 27.423 0.882 1.990 55.799 10.625 Table 18. — Steer 532. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 18,752 4,483 80.473 2.997 0.652 0.033 Circulatory system 47.915 40.256 1.605 0.566 0.105 Respiratory system 3,870 77.557 3.413 2.636 1.052 0.193 Brain and spinal cord 643 72.457 14.458 1.658 1.622 0.392 Digestive and excretory system (partial) Offal fat 21,741 23,697 441 72.701 9.552 13.979 88.805 1.898 0.243 0.708 0.116 0.134 0 016 Heart and neck sweetbreads 49.463 38.461 1.629 0.986 0.238 Liver 5,694 185 71.419 2.375 2.843 1.375 0.307 Gali 92 254 0.220 1.005 0.043 Spleen 884 74.698 5.033 2.888 1.256 0.262 Pancreas 630 56. 164 27.676 2.032 1.180 0.293 Kidneys 868 71.247 10.983 2.521 1.019 0.213 Hair and hide 33,988 4,260 59.564 6.839 5.225 1.032 0.071 Head and tail, lean and fat 60.370 21.950 2.687 0.807 0.146 Shin and shank, lean and fat 12,008 67.536 10.998 3.180 0.933 0.167 Flank and plate, lean and fat 44,636 43.753 41.749 2.237 0.655 0.116 Rump, lean and fat 8,058 48.528 35.883 2.345 0.716 0.133 Chuck and neck, lean and fat 66.204 61.792 18.520 2.910 0.899 0.156 Round, lean 38.064 71.904 5.223 3.304 1.065 0.200 Round, fat 6.064 21.714 70.321 1.113 0.295 0.042 Loin, lean 36,136 68.406 9.593 3.252 1.004 0.184 Loin, fat 14,954 12.520 83.405 0.674 0.191 0.033 Rib, lean 17,356 67.280 11.483 3.156 0.979 0.176 Rib, fat 6.194 15.622 78.878 0.852 0.268 0.040 Kidney fat 11.734 <5.271 95.229 0.150 0.085 0.022 Skeieton of feet 6.490 39.080 14.349 3.607 23.845 4.221 Skeleton of head and tail 7.120 46.409 12.787 2.985 21.246 3.794 Skeleton of shin and shank 10.756 28.890 21.617 4.316 23.357 4.397 Skeleton of flank and plate 5.478 44.408 18.946 3.027 14.958 2.738 Skeleton of rump 2.282 28.899 27.742 3.205 22.689 4.084 Skeleton of chuck and neck 13.014 30.283 23.624 3.367 24.027 4.351 Skeleton of round 5.624 26.867 34.032 2.514 20.794 3.713 Skeleton of loin 6.246 28.097 29.257 3.093 22.053 4.074 Skeleton of rib 5 222 34.969 22.509 2.951 22.280 4.009 Horns 228 54.753 0.511 6.327 7.572 1.439 Hoofs and dewclaws 1.406 494 51.936 0.646 7.653 1.948 0.153 Teeth 31.017 0.882 2.145 51.709 10.154 Studies In Animal Nutrition — III 49 Table 19. — Steer 538. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % 7,219 1.670 82.400 0.097 2.735 0.786 0.028 55.482 28.703 1.934 0.632 0.114 1,825 80.802 2.057 2.503 1.018 0.199 4S7 74.331 11.750 1.610 1.500 0.348 Digestive and excretory system (partial) 10,290 3,452 481 76.022 21.182 9.813 76.453 2.111 0.544 0.867 0.272 0.151 0.055 69.816 14.598 2.329 1 . 655 0.405 1,978 199 70.479 1.914 2.958 1.402 0.315 Gall 90.916 0.078 0.224 1.038 0.070 331 78.439 1.530 2.877 1.409 0.272 208 69.655 11.646 2.566 1.465 0.353 487 73.220 11.270 2.305 1.075 0 209 15.342 1,496 64.339 1.342 5.564 1.063 0 063 Head and tail, lean and fat 66.304 15.440 2.701 0.893 0.148 Shin and snank, lean and fat 4,190 11.036 71 811 5.773 3.294 1.002 0.179 Flank and plate, lean and fat 57 700 23.466 2.774 0.852 0.160 Rump, lean and fat 2,096 59 745 21.117 2.669 0.892 0 154 Chuck and neck, lean and fat 22.284 68.487 11 963 2.971 0.931 0 174 Round, lean 16.324 75.971 2.699 3.159 1.093 0.202 Round, fat 1,702 12.732 30.528 58.733 1.707 0.449 0.068 Loin, lean 73.141 5.617 3.117 1.055 0.203 Loin, fat 2,360 5,196 19.936 72.602 0.977 0 293 0.054 Rib. lean 70.881 8.150 3.075 1.017 0.191 Rib. fat 426 30 . 179 55.423 1.454 0.725 0.114 Kidney, fat 622 6.747 90 . 964 0.340 0.176 0.034 Skeleton of feet 3.168 42 . 658 15.526 3.293 18.615 3.342 Skeleton of head 4,001 49.779 7.792 2.917 21.579 4.089 Skeleton of tail 135 52 455 14.133 3.189 11 275 1 944 Skeleton of shin 2.208 28 636 23 827 2.827 25 111 3 924 Skeleton of shank 2.608 30.367 22.437 3.389 23.653 3.670 Skeleton of flank and plate 2,144 49.188 14.542 3.198 15.045 2.447 Skeleton of rump 732 34.282 23.745 3.050 22.231 3.797 Skeleton of chuck and neck 5.412 38.139 16.952 3.337 23.969 3.555 Skeleton of round 2.556 29.961 32.225 2.423 20 . 600 3.177 Skeleton of loin 2.696 34 . 968 23.196 2.940 21.259 4.163 Skeleton of rib 2,154 36.366 16.613 3.179 22.619 4.031 Horns 250 54 . 908 0.529 5.783 9.976 1.895 Hoofs and dewclaws 635 66 631* 0.465 5.359 1.015 0.067 Teeth 240 28.678f 1.111 1.961 54.175 10.200 *Hoofs and dewclaws of steer 540 and steer 538 were analyzed together, t Teeth of steer 540 and steer 538 were analyzed together. Table 20. — Steer 540. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 6,967 1,436 1.501 82.280 1.030 2.806 0.724 0.026 Circulatory system 58.131 27.726 2.039 0.676 0.127 Respiratory system 79.609 2.314 2.592 1.032 0.198 Brain and spinal cord 512 73.656 11.940 1.617 1.579 0.375 Digestive and excretory system (partial) Offal fat 9,280 2,307 408 75.239 25.244 10.756 71.058 1.995 0.548 1.703 0.282 0.129 0.047 Heart and neck sweetbreads 68.793 15.449 2.258 1.510 0.373 Liver 1,593 70.592 1.760 2.840 1.411 0 . 305 Gall 58 93.842 0.076 0.168 1.224 0.040 Spleen 331 77.793 1.372 3.097 1.408 0.271 Pancreas 180 72.201 9 . 605 2.537 1.457 0.343 Kidneys 363 72.153 10.499 2.368 1.084 0.223 Hair and hide 12,994 64 . 642 2.353 5.127 1.256 0.067 Head and tail, lean and fat. . . 1,274 07 992 13.404 2.736 0.883 0.167 8hin and shank, lean and fat 3,762 74 . 264 4.038 3.397 1 .039 0 185 Flank and plate, lean and fat 8,824 60.923 18.461 3.021 0.883 0.158 Rump, lean and fat 1,964 01 016 19 592 2.755 0.920 0.166 Chuck and neck, lean and fat 17,978 71 ins 8.347 2.961 0.986 0.178 Round, lean 13.456 75 . 698 2.255 3.208 1.082 0.210 Round, fat 810 27.830 62.313 1.356 0.388 0.060 Loin, lean 10,700 73 645 4.187 3.179 1.075 0.201 Loin, fat 2.308 10 598 73.692 0 . 958 0.288 0.058 Rib, lean and fat 5,046 69 429 9 677 3.096 1.013 0.181 Kidney, fat 682 13.401 79.809 1.156 0.185 0.035 50 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 20. — Steer 540. Analysis of Samples — Continued. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Skeleton of feet 2,784 3,682 44.008 12.128 3.288 20.841 3.573 Skeleton of head 51.688 7.055 2.754 21.417 4.056 Skeleton of tail 138 55.385 12.477 2.254 10.059 1.771 Skeleton of shin 1,952 31.687 22.158 3.498 24.179 4.561 Skeleton of shank 2,304 1,862 33.134 24.266 2.969 21.100 3.760 Skeleton of flank and plate 53 . 192 12.085 3.378 12.087 2.902 Skeleton of rump 622 35.209 20.449 3.031 22.451 4.221 Skeleton of chuck and neck 4.896 41.285 14.741 3.264 19.664 3.532 Skeleton of round 2,350 2.550 35.287 27.370 2.523 16.708 2.801 Skeleton of loin 34.808 27.381 2.786 19.208 3.575 Skeleton of rib 1.824 34.170 18.773 3.297 22.860 4.637 Horns 304 54 266 0.593 5.453 12.472 2.411 Hoofs and dewclaws 635 66.631* 0.465 5.359 1.015 0.067 Teeth 278 28 . 678f 1.111 1.961 54.175 10.200 *Hoofs and dewclaws of steer 538 and steer 540 were analyzed together. fTeeth of steer 538 and steer 540 were analyzed together. Table 21. — Steer 541. Analysis of Samples. Description of sample Weight in animal , grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus C7 /o Blood 12,470 2.646 2,387 568 82.112 0.089 2.773 0.679 0.027 Circulatory system 48.463 39.878 1.648 0.572 0.105 Respiratory system 79.217 1.976 2.738 1.040 0.213 Brain and spinal cord 72.458 13.605 1.658 1.525 0.371 Digestive and excretory system (partial) .... Offal fat 16.313 11,009 740 71.285 11.714 14.700 86.398 2.024 0.293 0.746 0.125 0.134 0.026 Heart and neck sweetbreads 65.439 19.148 2.280 1.457 0.367 Liver 3,832 202 69.540 2.502 3.158 1.441 0.349 Gall 89.007 0.174 0.286 1.207 0.060 Spleen 596 77.687 1.811 2.876 1.326 0.279 Pancreas 390 59.915 23.448 2.036 1.192 0.285 Kidneys 645 73.472 9.902 2.223 1.076 0.214 Hair and hide 26,574 2,062 6,830 24,910 4,454 40,480 27,000 3,854 23,416 60.425 3.650 5.692 1.100 0.055 Head and tail, lean and fat 66.470 14.520 2.939 1.016 0.164 Shin and shank, lean and fat 68.724 9.020 2.923 0.956 0.178 Flank and plate, lean and fat 49.802 34.241 2.427 0.717 0.127 Rump, lean and fat 50.532 32.775 2.370 0.713 0.137 Chuck and neck, lean and fat 65.263 14.772 2.942 0.913 0.165 Round, lean 73.915 3.366 3.340 1.089 0.208 Round, fat 21.202 73.399 0.987 0.282 0.046 Loin, lean 71.952 5.559 3.233 1.050 0.198 Loin, fat 8.088 14.861 80.634 0.780 0.204 0.039 Rib, lean 11,588 2,434 68.657 10.433 3.126 0.796 0.187 Rib, fat 17.784 76.039 1.080 0.312 0.059 Kidney, fat 6,056 4,506 5,400 4.494 94.350 0.202 0.086 0.020 Skeleton of feet 41.552 12.840 3.435 21.608 3.743 Skeleton of head 50.560 7.061 2.995 20.107 3.657 Skeleton of tail 206 47.886 19.594 3.175 10.884 1.894 Skeleton of shin 2,946 3,744 2,864 1,056 7,296 28.889 23.422 3.102 27.138 3.798 Skeleton of shank 32.657 20.792 2.933 27.763 3.778 Skeleton of flank and plate 51.920 15.329 2.277 11.840 1.799 Skeleton of rump 30.290 24.456 3.061 25.444 4.853 Skeleton of chuck and neck 34.849 18.389 3.368 25.110 3.622 Skeleton of round 3,250 3,916 26.958 32.458 3.253 19.430 4.211 Skeleton of loin 34.343 24.734 3.216 20.081 3.905 Skeleton of rib 3.162 31.674 21.579 3.195 25.537 3.726 Horns 468 53.053 0.655 5.345 13.848 2.689 Hoofs and dewclaws 869 58.119 0.599 6.018 1.249 0.061 Teeth 304 46.836 0.937 1.458 40.727 7.826 Studies In Animal Nutrition — III 51 Table 22 . — Steer 547 . Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 8,711 80.850 2.942 0.717 0.029 Circulatory system L624 58.440 27.812 1.966 0.682 0.124 Respiratory system 1,868 78.906 2.817 2.547 1.149 0.195 Brain and spinal cord 459 77.158 8.963 1.537 1.460 0.367 Digestive and excretory system (partial) 11,978 74.163 10.929 2.138 1.042 0.192 Offal fat 3,879 18.056 78.881 0.660 0.257 0.053 Liver 2,851 71.964 2.581 2.956 1.364 0.366 Spleen 391 77.086 2.212 3.020 1.345 0.299 Pancreas 200 71.405 10.115 2.426 1.455 0.376 Kidneys 450 73 . 255 8.553 2.560 1.144 0.240 Hair and hide 14,618 64.546 1.829 5.330 1.295 0.072 Head, tail, shin and shank, lean and fat 8.590 68.763 12.261 2.818 0.897 0.173 Flank and plate, lean and fat 14,226 55.003 27.523 2.502 0.793 0.147 Rump, lean and fat 2.256 54.409 27.572 2.464 0.818 0.157 Chuck and neck, lean and fat 23,636 67.588 12.501 2.772 0.918 0.167 Round, lean 17.092 74.440 3.584 3.223 1.079 0.210 Round, fat 2,150 29.872 60.663 1.343 0.397 0.068 Loin, lean 13,576 71.512 6.124 3.182 1.030 0.199 Loin, fat 3,972 21.083 72.247 0.987 0.300 0.057 Rib, lean 6.560 69.801 9.433 3.043 0.979 0.180 Rib, fat 1,102 25.085 66.287 1.438 0.503 0.075 Kidney, fat 1,630 7.887 90.103 0.312 0.136 0.015 Skeleton of feet, head, tail, shin and shank.. . 11,996 44.993 14.086 3.341 18.989 3.501 Skeleton of flank and plate 1,958 53.339 13.420 3.110 11.771 2.168 Skeleton of rump 760 37.716 15.923 3.294 23.682 4.444 Skeleton of chuck and neck 5,120 43.990 13.982 3.220 20.226 3.645 Skeleton of round 2,488 36.276 28.609 2.525 17.314 3.258 Skeleton of loin 3.132 39.384 20.100 2.909 20.155 3.841 Skeleton of rib 2,172 41.532 16.945 3.321 19.474 3.632 Horns, hoofs and dewclaws 737 53.209 1.331 7.310 2.465 0.220 Teeth 310 32.500 0.680 2.200 51.200 9.550 Table 23 . — Steer 548 . Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 4,603 753 81.198 3.100 0.715 0.021 Circulatory system 70 . 180 12.580 2.606 0.843 0.157 Respiratory system 1,084 76.883 3.131 2.908 1.106 0.202 Brain and spinal cord 526 74 . 634 11.062 1.637 1.434 0.350 Digestive and excretory system, partial Offal fat 5,357 845 78.972 52.573 4.268 36.778 2.386 1.461 1.066 0.720 0.179 0.122 Heart and neck sweetbreads 168 77.090 6.097 2.816 1.577 0.392 Liver 1,104 70.817 1.835 3.286 1.381 0.336 Spleen 215 77.264 1.368 3.131 1.342 0.281 Pancreas 85 75.456 6.206 2.812 1.394 0.331 Kidneys 353 78.549 3.384 2.489 1.237 0.247 Hair and hide 8,358 65.408 0.933 5.346 1.349 0.078 Head and tail, lean and fat 967 71.688 8.999 2.675 0.953 0.175 Shin and shank, lean and fat 2,462 74.845 4.448 3.267 0.998 0.175 Flank and plate, lean and fat 4,802 70.383 7.627 3.320 0.968 0.174 Rump, lean and fat 1,402 11,136 9,208 520 72.577 5.800 3.353 1.106 0.204 Chuck and neck, lean and fat 75.625 3.506 3.100 1.053 0.187 Round, lean 75.514 1.680 3.363 1.118 0.214 Round, fat 53.819 28.733 2.739 0.692 0.086 Loin, lean 5,668 384 75.325 2.020 3.292 1.128 0.209 Loin, fat 40 . 939 45.571 1.971 0.586 0.097 Rib, lean and fat 3,180 75.848 2.738 3.133 1.093 0.195 Kidney, fat 284 27.638 64.220 1.384 0.490 0.060 Skeleton of feet 2.496 2,989 94 45 . 620 13.826 3.344 18.414 3.215 Skeleton of head 56.949 6.785 2.771 17.108 3.133 Skeleton of tail 56.745 10.923 3.378 10.652 1.934 Skeleton of shin 1,584 1,712 1,514 570 38.976 19.708 3.222 19.487 3.568 Skeleton of shank 35.963 21.195 3.613 18 669 3.466 Skeleton of flank and plate 57.844 10.775 3.196 8.981 1.546 Skeleton of rump 43.892 15.924 3.252 18.272 3.323 Skeleton of chuck and neck 3.630 47 . 340 13 202 3.109 17.177 3.208 Skeleton of round 1,898 1 ,652 40.419 26.174 2.620 16.404 3.017 Skeleton of loin 43.992 18.943 3.128 17 030 3.230 Skeleton of rib 1,614 46 672 14.692 3.420 16.464 3.005 Horns, hoofs and dewclawB 435 55.591 1 . 144 6.927 2.606 0.264 Teeth 264 42.122 0.624 2.907 42.160 7.937 52 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 24 . — Steer 550 . Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 7,080 81.361 2.867 0 608 0 031 Circulatory system 1,163 55.997 29.913 1.989 0.660 0.133 Respiratory system 1,460 77.875 3.987 2.478 0.997 0.199 Brain and spinal cord 443 75.197 10.097 1.581 1.440 0.351 Digestive and excretory system (partial) .... 9,156 72.665 13.019 1.955 1.176 0.185 Offal fat 2.610 20.877 74.687 0.587 0.323 0.079 Liver 1,992 70.203 3.137 2.994 1.324 0.312 Spleen 291 76.774 2.659 2.941 1.480 0.325 Pancreas 200 66.906 15.542 2.442 1.338 0.320 Kidneys 379 75.357 6.259 2.503 1.173 0.252 Hair and hide 10,440 64.935 1.217 5.320 1.307 0.084 Head, tail, shin and shank, lean and fat 5,274 71.146 9.321 2.828 0.946 0.174 Plank and plate, lean and fat 7,896 61.653 19.406 2.857 0.888 0.164 Rump, lean and fat 1,662 59.888 20.746 2.688 0.918 0.178 Chuck and neck, lean and fat 15,814 67.239 14.099 2.685 0.950 0.180 Round, lean 11 720 75.505 2.653 3.418 1.063 0.213 Round, fat 944 32.441 57.921 1.439 0.416 0.069 Loin, lean 9.072 73.955 4.503 3.097 1.059 0.231 Loin, fat 2,134 21.819 71.450 0.918 0.326 0.065 Rib, lean and fat 4,408 69.082 11.035 2.924 0.996 0.187 Kidney, fat 756 10.430 86.878 0.455 0.229 0.057 Skeleton of feet, head, tail, shin and shank.. . 9,839 45.940 15.646 2.949 18.669 3.461 Skeleton of flank and plate 1,540 52.599 12.791 3.242 12.161 2.167 Skeleton of rump 700 40.804 16.240 3.381 19.007 3.635 Skeleton of chuck and neck 4,956 44.877 16.362 3.097 18.062 3.294 Skeleton of round 2,100 36.994 28.692 2.629 16.908 3.207 Skeleton of loin 2,464 38.480 24.756 2.896 17.775 3.340 Skeleton of rib 1,740 41.987 17.627 3.296 18.509 3.404 Horns, hoofs and dewclaws 549 53.209 1.331 7.310 2.465 0.220 Teeth 228 32.500 0.680 2.200 51.200 9.550 Table 25 . — Steer 552 . Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 5,219 79.774 3.010 0.854 0.028 Circulatory system 1,056 64.067 20.784 2.167 0.788 0.130 Respiratory system 1.096 77.900 2.614 2.724 1.206 0.205 Brain and spinal cord 466 74.074 11.478 1.564 1.369 0.335 Digestive and excretory system (partial) 5,936 75.009 9.539 2.159 1.058 0.174 Offal fat 1,784 28.924 67.756 0.924 0.380 0.069 Heart and neck sweetbreads 265 62.652 23.511 2.325 1.439 0.358 Liver 1,181 69.513 2.065 3.127 1.333 0.337 Spleen 284 77.867 1.297 3.100 1.340 0.279 Pancreas 101 73.221 9.258 2.422 1.292 0.301 Kidneys 316 72.871 12.099 2.305 1.128 0.221 Hair and hide 10,532 66.823 1.534 4.828 1.405 0.070 Head and tail, lean and fat 1,209 68.009 14.483 2.476 0.932 0.155 Shin and shank, lean and fat 2,976 74.470 4.636 3.211 0.976 0.172 Plank and plate, lean and fat 6,204 63.328 17.537 2.866 0.882 0.145 Rump, lean and fat 1,326 65.584 14.803 2.886 0.976 0.175 Chuck and neck, lean and fat 12,684 72.915 7.088 2.995 0.994 0.178 Round, lean 9,830 76.090 1.957 3.255 1.129 0.207 Round, fat 908 44.576 41.667 2.111 0.589 0.067 Loin, lean 7,034 74.559 3.588 3.131 1.101 0.194 Loin, fat 1,042 22.439 70.990 1.179 0.395 0.061 Rib, lean and fat 4,104 72.405 6.435 3.233 0.969 0.180 Kidney, fat 500 12.924 84.558 0.535 0.281 0.034 Skeleton of feet 2,861 43.452 12.823 3.478 19.568 3.480 Skeleton of head 3,052 54.140 6.234 2.916 18.811 3.453 Skeleton of tail 131 53.865 13.364 2.887 12.790 2.320 Skeleton of shin 1,550 39.817 17.664 3.553 18.794 3.332 Skeleton of shank 1,742 32.535 20.841 3.086 24.917 4.583 Skeleton of flank and plate 1,516 54.273 12.715 3.371 10.652 1.911 Skeleton of rump 640 39.791 17.248 3.300 20.750 3.671 Skeleton of chuck and neck 3,908 44.753 15.051 3.224 18.379 3.344 Skeleton of round 1.626 33.311 29.673 2.565 19.808 3.484 Skeleton of loin 2.192 41.054 20.912 2.874 18.374 3.457 Skeleton of rib 2.084 43.800 16.187 3.281 17.515 3.306 Horns, hoofs and dewclaws 550 57.684 1.828 6.410 2.692 0.261 Teeth 278 40.922 0.772 2.035 43.221 8.211 Studies In Animal Nutrition — III 53 Table 26. — Steer 554. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % 4,197 591 81.073 2.890 0.801 0.031 72.575 10.404 2.535 0.908 0.169 907 79.018 2.438 2.691 1.209 0.227 395 76.826 8.780 1.774 1.581 0.350 Digestive and excretory system (partial) .... 4,216 644 77.716 41.234 7.707 50.872 2.132 1.202 1.098 0.634 0.189 0.099 336 80.688 2.135 2.651 2.125 0.474 1,166 202 70.740 2.587 2.813 1.688 0 343 78.308 1.436 2.961 1.484 0 312 70.340 13.270 2.357 1.501 0 280 530 81.509 2.392 2.221 1 . 152 0 215 Hair and hide 7,400 883 66.018 1.615 4.976 1.442 0 096 Head and tail, lean and fat 72.777 8.344 2.812 1.253 0 191 Shin and shank, lean and fat 2,304 74.453 3.945 3.321 1.126 0.176 Flank and plate, lean and fat 4,232 70.739 8.896 2.995 0.980 0.170 Rump, lean and fat 1,052 70.442 9.091 3.057 1.166 0.178 Chuck and neck, lean and fat 10.530 7,918 520 74.853 4.416 3.102 1.245 0.189 Round, lean 75.950 2.163 3.286 1.271 0.215 Round fat 51.440 32.679 2.668 0.762 0 083 Loin lean 5,812 428 74.847 3.700 3.199 1.150 0.207 Loin fat 29.732 57.870 1 394 0 536 0 079 Rib, lean and fat Kidney, fat 2,914 240 74.793 18.564 3.459 75.417 3.261 0.727 1.138 0.344 0.198 0.057 Skeleton of feet 2,403 46.494 13.980 3.702 17.687 3.284 Skeleton of head 2.229 125 59.707 3.158 3.111 16.850 3 152 Skeleton of tail 56.788 11.330 3.423 11 074 2 065 Skeleton of shin 1.464 40 004 18.416 3.221 20.982 3.982 Skeleton of shank 1.918 40.266 19.360 3.248 17.454 3.124 Skeleton of flank and plate 1.278 60 681 9.953 3 288 7.753 1.374 Skeleton of rump 732 48.799 12.664 3.475 16.288 2.950 Skeleton of chuck and neck 3,732 51.461 10.035 3.153 16.389 2.945 Skeleton of round 1,702 1.868 1.354 338 40 . 198 24.701 2.669 15.953 2 707 Skeleton of loin 50 . 157 13.850 2.994 14.999 2 690 Skeleton of rib 50.983 12.314 3.175 15.340 2 660 Horns, hoofs and dewclaws 55.271 1.451 7.670 2.390 0.229 Teeth. 225 46.326 0.098 2.946 38.711 6.360 Table 27. — Steer 555. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 4,529 554 80.075 3 124 0.738 0.035 Circulatory system 70 . 982 12 062 2.574 0.875 0.159 Respiratory system 937 79.445 2.779 2.693 1.115 0.208 Brain and spinal cord 353 76.371 9.945 1.705 1.535 0.354 Digestive and excretory system (partial) .... Offal fat 4,534 462 78.376 58 692 6.357 31.373 2.319 1.720 1.091 0.779 0.189 0.119 Heart and neck sweetbreads 159 75 984 6.375 2.732 1.347 0.274 Liver 1,240 167 74 635 2.536 2.933 1 493 0.349 Spleen 77.376 1.363 3.129 1.308 0.276 Pancreas 106 76.261 5 510 2 622 1.458 0.293 Kidneys 439 81.997 2.743 2.220 1.181 0.223 Hair and hide 6,580 956 69.051 0 . 755 5.200 1.476 0.089 Head and tail, lean and fat 72.679 9.234 2.630 1.055 0.162 Shin and shank, lean and fat 2,688 4,068 876 77.046 2.331 3.187 1.044 0.181 Flank and plate, lean and fat 75.514 3.424 3.179 1.038 0.175 Rump, lean and fat 71.979 7.274 2.915 1.070 0.187 Chuck and neck, lean and fat 9,402 7,126 376 77 450 2 143 3.130 1 041 0.185 Round, lean 77 949 1 347 3.059 1.194 0.207 Round, fat 58 987 23.814 2.599 0.747 0.090 Loin, lean 4,618 274 77.939 1.726 3.105 1.200 0.208 Loin, fat 41 153 42.967 2.179 0.822 0.100 Rib, lean and fat 2,512 130 77 (11 1 2 . 536 3.101 1.174 0.189 Kidney, fat 32 . 970 57.414 1.209 0.549 0 091 Skeleton of feet 2,157 51 458 9.040 4.041 16 248 2 781 Skeleton of head 1,978 74 62.439 3.214 2.859 14.812 2.570 Skeleton of tail 63 . 368 6.593 3.646 9.138 1.523 Skeleton of shin 1,454 50.339 1 1 063 3.234 17.828 3.190 Skeleton of shank 1,704 49.701 13.806 2.882 14.863 2.656 54 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 27. — Steer 555. Analysis of Samples — Continued. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Skeleton of flank and plate 1,236 506 65.173 5.690 3.369 8.036 1.361 Skeleton of rump 57 . 102 5.911 3.496 16.138 2.764 Skeleton of chuck and neck 3,216 1,652 59.309 5.153 3.142 13.912 2.463 Skeleton of round 53.566 13.254 2.793 14.801 2.582 Skeleton of loin 1,264 1.256 59.306 7.164 3.098 13.007 2.313 Skeleton of rib 58.491 6.345 3.269 13.194 2.272 Horns, hoofs and dewclaws 298 48.852 0.914 7.593 2.662 0.133 Teeth 190 42.647 0.330 2.870 42.862 7.297 Table 28. — Steer 556. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 6,124 81.472 2.778 0.784 0.035 Circulatory system 993 62.355 23.955 2.146 0.856 0.148 Respiratory system 1,272 77.882 3.420 2.698 1.400 0.216 Brain and spinal cord 398 67.120 19.452 1.624 1.616 0.349 Digestive and excretory system (partial) .... 6,003 74.523 10.512 2.203 1.304 0.241 Offal fat 1,402 32.618 61.309 0.961 0.451 0.063 Heart and neck sweetbreads 328 76.405 6.578 2.590 2.052 0.452 Liver 1,760 71.112 3.500 3.060 2.295 0.366 Spleen 300 77.717 2.323 2.983 1.480 0.272 Pancreas 96 72.244 10.649 2.415 1.503 0.302 Kidneys 338 75.331 7.087 2.522 1.345 0.254 Hair and hide 10,314 66.284 2.153 5.157 1.377 0.088 Head and tail, lean and fat 914 68.746 13.192 2.739 0.897 0.165 Shin and shank, lean and fat 2,808 73.731 4.508 3.271 0.966 0.185 Flank and plate, lean and fat 6,174 67.267 13.359 3.143 0.935 0.168 Rump, lean and fat 1,244 69.336 10.388 3.015 1.036 0.191 Chuck and neck, lean and fat 12,406 72.316 7.615 3.016 1.246 0.180 Round, lean 9,472 75.179 3.195 3.289 1.251 0.210 Round, fat 686 44.562 43.541 2.194 0.577 0.069 Loin, lean 6,938 74.185 3.780 3.270 1.189 0.206 Loin, fat 820 26.114 66.035 1.399 0.542 0.064 Rib, lean and fat 3,300 71.440 7.223 3.184 1.153 0.184 Kidney, fat 420 15.362 80.619 0.587 0.355 0.037 Skeleton of feet 2,589 46.442 14.176 3.729 17.758 3.279 Skeleton of head 2,610 56.628 7.187 2.990 17.629 3.271 Skeleton of tail 92 56.900 9.965 3.538 13.365 2.393 Skeleton of shin 1,702 40.276 19.800 3.280 19.776 3.695 Skeleton of shank 1,952 37.454 22.610 3.389 19.971 3.787 Skeleton of flank and plate 1,578 57.659 11.660 3.313 10.266 1.862 Skeleton of rump 608 47.189 13.166 3.537 18.150 3.408 Skeleton of chuck and neck 3,734 47.104 12.535 3.488 18.545 3.545 Skeleton of round 1,974 38.147 25.725 2.665 16.702 3.212 Skeleton of loin 2,268 46.157 15.957 3.158 16.929 3.309 Skeleton of rib 1,646 46.368 12.349 3.310 18.157 3.720 Horns, hoofs and dewclaws 390 48.189 1.236 8.303 1.870 0.156 Teeth 218 41.753 0.042 2.938 43.927 6.910 Studies In Animal Nutrition — III 55 Table 29. — Steer 557. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % 8,952 79.941 3.136 0.842 0.027 Circulatory system 2,342 50.516 38.557 1.650 0.615 0.105 Respiratory system 1.972 77.053 5.258 2.598 1.119 0.203 Brain and spinal cord 551 72.578 12.435 1.790 1.410 0.338 Digestive and excretory system (partial) .... 9.981 73.851 11.446 2.007 1.051 0.176 Offal fat 6.757 13.945 83.471 0.435 0.229 0.029 Heart and neck sweetbreads 623 67.787 16.888 2.328 1.653 0.357 I.iver 3,003 69.463 1.882 3.084 1 344 0.332 Spleen 485 76.934 3.341 2.888 1.372 0.281 Pancreas 225 64.844 17.153 2.276 1.068 0.251 Kidneys 649 76.753 5.885 2.376 1.151 0.218 Hair and hide 14,100 63.124 5.012 4.988 1.171 0.071 Head and tail, lean and fat 1.915 60.473 24 . 133 2.352 0.848 0.144 Shin and shank, lean and fat 4,196 68.205 11.827 2.933 0.927 0.153 Flank and plate, lean and fat 15,294 50.434 34.335 2.334 0.778 0.133 Rump, lean and fat 2,582 51.177 32.845 2 317 0.797 0.142 Chuck and neck, lean and fat 22.146 63.903 17.574 2.702 0.957 0.183 Round, lean 14,960 73.824 4.610 3.073 1.011 0.194 Round, fat 2,532 27.746 63.644 1.019 0.349 0.048 Loin, lean 11.714 71 . 155 7.826 2.966 1.098 0.192 Loin, fat 4.472 15.907 79.899 0.729 0.248 0.038 Rib, lean 6.372 67.831 11 987 2.905 0.958 0.170 Rib, fat 1,616 19.571 73.914 0.959 0.360 0.059 Kidney, fat 3.228 8.267 89.768 0.367 0.159 0.024 Skeleton of feet 3,558 44.384 11.995 3.601 18.856 3.488 Skeleton of head 3,969 53.816 6.263 2.751 18.265 3.294 Skeleton of tail 193 53.009 16.791 3.163 10.458 1.954 Skeleton of shin 2,246 36.792 18.013 3.351 21.238 3.961 Skeleton of shank 2,610 36.883 19.539 3.405 20.052 3.625 Skeleton of flank and plate 2,498 56.391 12.656 3.152 9.709 1.721 Skeleton of rump 960 42.749 13.401 3.441 19.159 3.710 Skeleton of chuck and neck 5,638 42.943 12.074 3.394 20.847 3.764 Skeleton of round 2,686 32.505 29.395 2.573 19.164 3.548 Skeleton of loin 2,478 38.626 20.143 2.959 19.381 3.655 Skeleton of rib 2,572 40.350 16.831 3.239 19.671 3.611 Horns, hoofs and dewclaws 695 53.666 1.415 6.956 2.569 0.279 Teeth 261 39.979 0.283 2.874 46.115 8.632 Table 30. — Steer 558. Analysis of Samples. Description of sample Weight in animal, grams Moisture % Crude fat % Nitrogen % Ash % Phosphorus % Blood 4,666 83 . 665 2.509 0.692 0.039 Circulatory system 971 66.245 19.036 2.219 0.726 0.143 Respiratory system 1,111 79.632 1.472 2.685 1.054 0.231 Brain and spinal cord 520 75.596 10.002 1.596 1.507 0.374 Digestive and excretory system (partial) .... 6,510 77.962 6.366 2.241 0.992 0.188 Offal fat 829 47.598 43.206 1.386 0.505 0.106 Liver 1,352 71.470 2.100 3.096 1.349 0.352 Spleen 193 75.805 1.099 3.298 1.537 0.278 Pancreas 153 76.282 5.186 2.553 1.452 0.374 Kidneys 318 76.794 4.347 2.595 1.193 0.247 Hair and hide 8,138 64.713 1.078 5.287 1 . 130 0.083 Head, tail, shin and shank, lean and fat 4,324 74.380 5.509 2.948 0.943 0.163 Flank and plate, lean and fat 4,192 70.650 7.311 3.229 0.923 0.172 Rump, lean and fat 1,030 69.840 9.636 3.089 1.011 0.196 Chuck and neck, lean and fat 11,682 75.244 4.121 3.041 0.927 0.187 Round, lean 9,664 77.152 1.113 3.116 1.049 0.213 Round, fat 644 44.994 41.335 2.308 0.611 0.095 Loin, lean 5,952 74 950 2.652 3.182 1.025 0 210 Loin, fat 600 30.204 59.420 1.549 0.446 0.080 Rib, lean and fat 3,242 74.678 3.490 3.157 0.990 0.197 Kidney, fat 220 18.408 75.158 0.936 0.283 0.055 Skeleton of feet, head, tail, shin and shank . . 9,785 45.737 16 814 3.097 17.541 3.252 Skeleton of flank and plate 1,092 54.633 1 - 519 3.306 10.416 1.885 Skeleton of rump 678 42. 549 18.378 3.113 18.671 3.383 Skeleton of cnuck and neck 4,056 43 212 18 090 3.089 17.848 3.240 Skeleton of round 2,094 34.269 33.809 2.285 16.102 2.982 Skeleton of loin 2,138 39 771 25.039 2.529 17 837 3.281 Skeleton of rib 1,516 46.885 16 047 3.348 15.188 2.747 Homs, hoofs and dewclaws 443 53.209 1.331 7.310 2.465 0.220 Teetn 274 32.500 0.680 2.200 51.200 0.550 56 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 31. — Steer 500. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Pnospnorus Blood 21,269 16811.2 40.8 679.1 167.8 4.68 Circulatory system 1,562 756.8 587.9 29.2 8.5 0.95 Lean neart 1,284 995.7 45.7 35.0 14.2 2.67 Respiratory system 3,747 2866.5 101.2 106.4 42.9 6.45 Fat from thoracic cavity 568 96.8 450.9 2.8 1.4 0.14 Brain and spinal cord 832 522.2 180.7 13.9 14.5 3.09 Digestive and excretory system (partial) 19,275 14385.9 1921.9 424.6 160.8 22.75 Offal fat 12,940 1747.0 10812.2 54.6 23.8 2.59 Heart and neck sweetbreads 538 286.2 182.6 9.9 5.6 1.21 Liver 4.634 3233.9 134.5 150.3 73.2 14.97 Gall 241 221.4 0.5 0.5 3.0 0.07 Spleen 1,054 783.9 56.4 31.4 12.6 2.31 Pancreas 625 369.6 156.8 13.4 7.5 1.60 Kidneys 1,019 785.6 49.0 24.7 11.4 2.11 Tongue, marketable 1,619 1123.7 192.2 43.8 14.8 2.49 Hair and hide 35,938 21320.9 474.0 2256.9 385.3 15.81 Head and tail, lean and fat 3,784 2410.9 603.9 119.4 33.4 5.07 Shin and shank, lean and fat 12,496 8854.9 823.6 420.2 123.6 20.49 Flank and plate, lean and fat 36,410 19948.3 10067.7 978.3 315.3 50.61 Rump, lean and fat 7,058 3892.4 1950.8 178.4 57.8 10.23 Chuck and neck, lean and fat 58,918 39825.0 6993.0 1995.6 537.3 93.09 Round, lean 39,898 29536.9 1390.5 1246.0 403.4 76.21 Round, fat 4,936 1370.6 3032.8 78.5 18.6 2.52 Loin, lean 29,692 20864.3 2296.4 924.3 299.9 54.93 Loin, fat 6,830 1124.5 5225.5 40.8 16.7 2.60 Rib, lean 13,602 9132.0 1676.2 434.7 126.4 23.12 Rib, fat 1,804 367.4 1282.4 23.3 6.7 1.08 Kidney, fat 2,432 170.9 2195.5 10.0 3.5 0.44 Skeleton of feet 6,838 2708.1 788.3 247.0 1707.5 309.69 Skeleton of head 8,953 4296.2 1216.2 312.2 1599.2 307.45 Skeleton of tail 386 151.7 92.7 10.2 61.4 10.75 Skeleton of shin 5,610 1485.9 1210.9 207.6 1651.6 292.34 Ske*eton of snank 5 750 1808.8 1160.8 198.6 1448.0 254.67 Skeieton of flank and plate 6,350 2605.5 1143.5 204.7 1177.1 203.20 Skeleton of rump 2,988 727.3 914.6 91.6 749.8 132.37 Skeleton of chuck and neck 14,450 4302.5 3253.7 442.2 3746.3 661.09 Skeleton of round, (excl. marrow) 6,438 2095.7 1789.3 166.7 1356.7 243.74 Marrow from skeleton of round 680 64.3 606.9 1.0 1.5 0.21 Skeleton of loin 7,772 1947.4 2438.5 228.1 1865.8 332.41 Skeleton of rib 5,192 1409.4 1158.2 165.2 1449.9 257.11 Hoofs and dewclaws 2 095 1059 . 7 17.5 162.2 54.6 2.45 Teeth 852 181.7 9.9 17.7 519.8 98.12 Table 32. — Steer 501. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 28,710 1,836 1,882 3,838 22387.2 50.5 944.6 246.0 7.18 Circulatory system 770.4 842.5 35.0 8.7 0.90 Lean heart 1460.7 70.4 48.4 18.7 3.73 Respiratory system 2939.0 128.5 110.3 40.0 6.60 Fat from thoracic cavity 2,459 757 463.1 1884.7 15.1 5.8 0.64 Brain and spinal cord 533.0 100.5 12.7 13.9 2.97 Digestive and excretory system (partial) Offal fat 24,235 38,625 784 17390.1 2892.2 3119.0 35172.3 519.4 79.2 196.1 39.4 29.81 4.64 Heart and neck sweetbreads 241.1 484.2 7.8 3.9 0.84 Liver 6,161 176 4282.7 178.6 199.2 87.7 20.58 Gall 161.8 0.1 0.4 2.2 0.06 Spleen 1,178 836 917.6 23.0 32.7 16.3 2.82 Pancreas 500.5 205.4 18.4 9.6 2.20 Kidnevs 1,037 805.3 50.5 24.3 10.9 2.06 Tongue, marketable 2,153 50,090 5,224 17,420 134,146 22,226 110,990 50,130 22,284 45,996 71,358 1417.6 348.8 56.2 18.7 3.38 Hair and hide 25762.3 6629.4 2751.4 762.4 24.54 Head and tail, lean and fat 3156.4 1083.8 148.0 40.1 6.58 Shin and shank, lean and fat 10268.9 3932.2 471.6 134.5 23.17 Flank and plate, lean and fat 35964.5 88380.8 1408.5 458.8 76.46 Rump , lean and fat 6394 2 13949.0 253.4 87.8 15.33 Chuck and neck, lean and fat 52940.0 42652.4 1692.6 723.7 130.97 Round, lean 35041.9 4690.2 1549.0 479.7 92.74 Round, fat 3754.0 17434.3 148.6 48.6 5.79 Loin, lean 28773 . 7 8248.9 1316.9 391.4 74.97 Loin, fat 6444.3 63281.7 276.9 79.9 12.84 Rib, lean 20,834 12269.6 4668.7 574.8 164 8 31.04 Rib, fat 28,322 19,544 7,744 2746.4 24764.5 113.6 38.0 5.66 Kidnev, fat 1067.5 18236.7 37.1 13.1 2.15 Skeleton of fppt 2792.0 954.5 273.4 2023.7 390.68 Studies In Animal Nutrition — III 57 Table 32. — Steer 501. Weights of Constituents in Samples, Grams — C ont. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus 10,462 304 4561.8 1232.0 343.9 2487.5 439.30 Skeleton of tail 124.3 58.3 10.6 54 . 9 9.59 Skeleton of shin 6,170 2018 3 875.4 214.5 1787.3 323.12 Skeleton of shank 7,128 7,068 1920.5 1581.5 238.7 1985 . 9 365.24 Skeleton of flank and plate 2842.3 1107.0 234.7 1328 . 9 240.52 Skeleton of rump 3,682 932 8 965.2 116.8 956.3 172.61 Skeleton of ctruck and neck 15,778 4750.1 2444.8 576.8 4478.0 821.24 Skeleton of round, (excl. marrow) 6,978 1859.4 1699.2 215 3 190 i .7 345.27 Marrow from skeleton of round 286 29.1 252.8 0.6 1.5 0.24 Skeleton of loin 8,614 6,388 3,354 2,523 778 2214.8 1940 . 1 269.4 2447.3 436.90 Skeleton of rib 1816.0 1173.5 212.2 1780.2 330.96 Horns 1240.6 21.2 217.0 762.8 139.76 Hoofs and dewclaws 1186.1 16.6 213.3 43.3 3.61 Teeth 172.0 6.3 16.1 465.1 91.31 Table 33. — Steer 502. Weights of Constituents in Samples, Grams. Description of sample Sample Water • Crude fat Nitrogen Asn Phosphorus Blood 19,728 15276.0 688.9 142.0 4.54 Circulatory system 3,446 2353.1 500.1 89.1 23.4 4.76 Respiratory system 3,696 2815.1 113.0 104.6 38.4 6.14 Fat from thoracic cavity 1.271 273.3 933.9 9.6 3.8 0.52 Brain and spinal cord 800 552.8 116.8 13.7 14.4 3.31 Digestive and excretory system (partial) 20,933 16122.6 1263.3 522.3 264.6 28.47 Ofial fat 11,377 1211.8 9804.7 53.2 21.1 2.62 Heart and neck sweetbreads 502 300.1 127.6 11.3 5.8 1.36 Liver 3,716 2561.9 64.2 122.8 51.7 12.41 Gall 241 224.2 0.1 0.5 2.5 0.07 Spleen 921 711.9 22.3 26.1 16.9 2.57 Pancreas 581 310.6 172.1 12.6 6.2 1.39 Kidneys 838 617.3 55.2 22.5 9.3 1.85 Hair and hide 39,556 22665.2 1190.6 2600.4 386.1 23.34 Head and tail, lean and fat 4,250 2745.5 600.2 136.0 35.1 6.33 Shin and shank, lean and fat 12,364 8444.0 1072.1 431.0 109.3 20.28 Flank and plate, lean and fat 35,594 18475.1 11113.2 919.4 247.7 42.36 Rump, lean and fat 8,100 4538.3 2059.5 211.5 64.6 11.91 Chuck and neck, lean and fat 70,744 46926.6 9004.3 2173.3 651.6 111.78 Round, lean 44,426 32054.7 1799.7 1468.7 431.4 86.63 Round, fat 4,620 1229.0 2903.1 71.6 14.9 1.76 Loin, lean 35,104 24529.6 2852.5 1120.9 342.3 69.86 Loin, fat 9,144 1416.4 7149.0 95.7 22.8 3.29 Rib, lean 18,256 12114.3 1846.8 560.1 163.0 29.94 Rib, fat 3,338 689.5 2336.4 49.5 9.4 1.70 Kidney, fat 2,916 215.4 2614.7 12.3 7.0 1.14 Skeleton of feet 6.982 2897.3 852.2 264.5 1666.5 296.04 Skeleton of head 9,577 4669.5 802.4 305.0 1934.2 331.36 Skeleton of tail 441 178.3 99.6 14.3 66.5 12.05 Skeleton of shin 5,940 1627.5 1147.6 209.8 1750.3 311.97 Skeleton of shank 5.978 1625.8 1469.7 206.5 1516.2 269.31 Skeleton of flank and plate 5,590 2300.6 726.5 189.1 1219.6 215.83 Skeleton of rump 2,362 600.8 702.3 71.5 559.4 99.96 Skeleton of chuck and neck 15,092 4643.4 2755.7 514.2 4158.3 742.22 Skeleton of round (excl. marrow) 6,296 1667.0 1664.9 188.6 1653.3 294.72 Marrow from skeleton of round 408 32.1 369.8 0.8 1.7 0.30 Skeleton of loin 7,866 2068.0 2085 . 6 247.7 2032.5 359.40 Skeleton of rib 5,920 1801.6 1219.5 215.9 1472.0 260.54 Horns* 1,949 745.9 11.9 131.0 410.3 72.76 Hoofs and dewclaws 2,010 1179.6 11.5 133.2 23.8 2.41 Teeth 1,038 375.9 10.6 16.9 519.0 98.43 •This sample was lost before analysis. The average analysis of the horns of two mature steers in the. same group was used. 58 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 34 . — Steer 503 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 13,058 10809.4 14.4 353.6 43.9 9.92 Circulatory system 1,782 645.9 978.6 22.3 5.9 1.07 Lean heart 1.063 830.6 39.7 27.5 10.1 2.20 Respiratory system 2,549 2006.2 80.4 67.7 24.9 5.28 Brain and spinal cord 666 486.9 107.3 11.2 10.3 2.62 Digestive and excretory system (partial) 8,761 6370.1 964.9 208.1 88.7 18.22 Offal fat 7,385 1081.3 6049.8 38.5 13.4 2.58 Liver 3,646 2504.1 192.0 109.2 46.8 12.18 Kidneys 655 467.4 77.3 15.8 6.8 1.47 Stomach 5,765 4477.0 399.2 127.5 62.6 11.93 Tongue, marketable 789 547.0 104.7 20.0 6.5 1.34 Hair and hide 23,008 15591.4 591.3 1102.8 225.3 15.18 Shin, shank, head and tail, lean and fat 8,614 6004.6 880.6 275.3 72.9 14.46 Flank and plate, lean and fat 16,290 9206.8 4181.0 446.7 121.9 22.17 Chuck and neck, lean and fat 31,934 21883.1 4015.4 943.3 278.0 54.61 Round and rump, lean 25,022 18287.8 1202.3 843.2 256.1 51.14 Round and rump, fat 3,400 763.3 2384.4 41.2 9.8 1.62 Loin, lean 17,206 12215.2 1454.1 548.4 169.1 32.69 Loin, fat 5,746 904.9 4592.0 44.7 10.9 2.01 Rib, lean 8,932 6223.6 913.8 281.7 82.8 16.52 Rib, fat 840 195.7 562.3 12.4 2.7 0.51 Kidney, fat 2,126 184.5 1902.1 7.2 2.7 0.58 Skeleton 41,122 15740.3 6192.6 1276.4 9747.6 1800.32 Horns, hoofs and dewclaws 1,058 489.7 21.5 80.2 64.1 6.99 Teeth 253 59.0 1.3 5.7 149.0 28.54 Table 35 . — Steer 504 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 21,005 16520.4 19.5 700.3 81.3 4.62 Circulatory system 1,637 470.0 1043.3 18.2 6.6 0.67 Lean heart 1,428 1094.6 61.6 39.6 14.5 2.90 Respiratory system 3,628 2445.6 540.6 104.6 35.3 6.57 Brain and spinal cord 724 501.2 109.2 13.0 9.8 2.53 Digestive and excretory system (partial) 17,569 12041.8 3169.5 335.6 127.6 25.48 Offal fat 25,105 3203.4 21294.1 83.9 39.9 5.77 Liver 4,754 3280.7 115.8 155.7 64.3 17.02 Kidneys 877 609.3 112.3 21.5 9.3 1.92 Stomach 12.820 1Q213.7 1030.7 218.1 115.0 19.36 Tongue, marketable 1,587 964.1 372.6 34.6 12.1 2.24 Hair and hide 41,144 23982.8 3320.3 2272.0 434.9 17.69 Shin, shank, head and tail, lean and fat 16,070 9744.9 3304.0 474.2 129.0 23.46 Flank and plate, lean and fat 49,650 20674.3 22650.3 965.7 284.0 50.15 Rump, lean and fat 10,846 4416.5 5077.0 200.3 62.3 11.50 Chuck and neck, lean and fat 59,808 34886.0 14371.9 1567.0 453.3 87.32 Round, lean 37,238 25884.1 3429.6 1194.6 366.1 72.24 Round, fat 9,818 1630.8 7661.0 89.0 23.4 2.95 Loin, lean 33,676 22535.9 4115.2 1027.5 318.6 60.95 Loin, fat 18,340 2131.1 15572.5 97.6 29.7 4.59 Rib, lean 18,506 11710.6 3242.3 544.1 153.8 30.91 Rib, fat 6,770 976.2 5458.7 56.4 13.7 2.10 Kidney, fat 11,400 547.2 10709.2 24.5 14.4 1.94 Skeleton of feet, head, tail, shin and shank.. . 23,568 8496.3 3214.7 762.9 6503.4 1196.31 Skeleton of flank and plate 4,572 2916.3 831.2 161.1 718.5 133.32 Skeleton of rump 2,428 624.5 631.3 74.6 664.8 125.87 Skeleton of chuck and neck 11,176 3372.9 1818.3 393.8 3259.2 597.25 Skeleton of round 4,808 1052.0 1320.8 150.0 1492.8 276.70 Skeleton of loin 5,850 1745.6 1292.9 186.3 1547.4 281.50 Skeleton of rib 5,092 1657.5 828.0 174.0 1415.1 259.49 Horns, hoofs and dewclaws 2,532 1759.1 13.0 117.0 61.5 3.98 Teeth* 338 86.2 3.6 6.4 196.8 37.57 ♦This sample was lost before analysis. The average analysis of the teeth of four steers of the same Group was used. Studies In Animal Nutrition — III 59 Table 36 . — Steer 505 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 13.810 11360.1 48.5 376.5 45.4 3.18 Circulatory system 1.168 388.1 689.9 15.0 3.2 0.55 Lean heart 938 725.8 47.9 24.6 9.1 1.96 Respiratory system 2,498 1919.2 136.8 67.8 23.8 5.05 Brain and spinal cord 537 396.4 79.1 9.1 9.2 2.21 Digestive and excretory system (partial) Offal fat 8.258 5936.9 1058.3 204.5 87.0 18.66 12,781 1586.1 10912.9 43.5 15.0 2.81 Liver 3,983 2712.3 229.8 127.7 52.9 13.82 Kidneys 718 543.6 56.3 17.1 7.6 1.62 Stomach 8,818 6812.9 956.7 148.2 76.5 15.26 Tongue, marketable 1.115 714.3 224.8 26.2 8.5 1.74 Hair and hide 22.884 14219.7 1221.1 1212.2 159.3 15.10 Shin, shank, head and tail, lean and fat 9.386 6047.4 1445.3 301.9 76.8 15.40 Flank and plate, lean and fat 24,194 10577.6 10345.8 535.0 140.3 28.07 Chuck and neck, lean and fat 38,344 23886.0 7265.8 1106.6 318.3 63.27 Round and rump, lean 25 784 17808.0 2440.2 857.6 251.7 51.57 Round and rump, fat 5,970 844.2 4814.2 27.0 10.4 1.91 Loin, lean 19,686 13423.9 1965.3 637.0 189.6 38.58 Loin, fat 7.558 705.4 6616.8 40.4 9.6 1.83 Rib, lean 11,300 6979.1 2168.6 334.9 95.0 19.89 Rib, fat 2.640 288.1 2254.2 17.0 4.4 0.87 Kidney, fat 5,754 302.8 5381.5 13.6 4.8 0.92 Skeleton 37,745 13509.7 6626.2 1202.6 9002.9 1661.92 Horns, hoofs and dewclaws 1,206 555.9 12.2 93.2 64.4 7.37 Teeth 268 58.8 1.7 6.1 159.2 28.67 Table 37 . — Steer 507 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 20,316 4,278 15881.8 697.0 136.1 4.47 Circulatory system 2015.7 1765 . 7 72 5 22.9 4.45 6.37 3.11 41 13 Respiratory system 3,768 744 2919.0 148.0 108.8 11 2 35.9 Brain and spinal cord 526.8 102.2 13.0 Digestive and excretory system 27,417 11,307 34,473 4.038 19333.1 3860.0 574.9 226.2 Offal fat 1480.8 9493.6 46.6 15.8 3.28 16.89 Hair and hide 20975 . 1 2141.8 1960.8 367.1 Head and tail, lean and fat 2527.1 765.2 114.7 36.1 6.51 Shin and shank, lean and fat 11,860 36,130 7,736 62,530 8288 . 1 932.2 396.5 109.2 19.81 Flank and plate, lean and fat 18765.6 11692.4 904.7 250.4 44 81 Rump, lean and fat 3930.1 2448.0 176.0 55.8 10 75 Chuck and neck, lean and fat 41000.3 9483.3 1807.7 539.0 98 07 Round, lean 39,302 28583 . 2 2268.5 1269.5 385.6 75.46 Round, fat 5,378 1314.7 3693.1 58.8 14.8 2 10 Loin, lean 29,724 10,188 15,788 2,432 21013.7 2406.5 929.8 288.9 54 99 Loin, fat 1815.7 7812.6 84.2 22.7 3 97 Rib, lean 10647.1 1932.3 473.5 147 9 27.47 Rib, fat 426.9 1858.9 23.0 6.2 1.02 Kidney, fat 4,376 15,275 296.9 3995.3 12.4 6.4 1.09 Skeleton of feet, head and tail 6538.3 1798.8 527.9 3651.6 589.31 Skeleton of shin and shank 10,350 6,278 2,536 2479.9 2003.5 377.4 3489.7 479.62 Skeleton of flank and plate 2775.9 846.6 212.2 1159.7 178.99 Skeleton of rump 635 0 668.0 81.9 655.2 99 92 Skeleton of chuck and neck 13,202 5.864 4222.4 2012.6 469.3 3385.5 555.01 Skeleton of round 1530.1 1756.9 184.3 1360.7 244.00 Skeleton of loin 6,506 5,050 1732.6 1749.5 186.5 1698.2 242 . 28 Skeleton of rib 1448.0 910.1 169.8 1451.3 216.24 Horns 1,800 1,490 712 665 . 7 10.0 92.2 350.0 63.36 Hoof 8 and dewclaws 811.1 17.0 104.3 30.5 2.43 Teeth 188.9 7.3 14.0 403.8 77.53 60 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 38. — Steer 509. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus 18.291 14276.7 627.2 125.5 4.21 4.37 5.78 0.54 2.80 23.39 2.38 1.60 12.63 2,616 3,283 1829.8 320.8 67 8 24 1 2529.0 94.5 98.6 33.2 1,248 247.2 949.7 9.0 3 7 739 494.2 130.5 12.4 11 7 Digestive and excretory system (partial) .... 19.016 9,922 13883.8 1127.1 2260.2 8521.8 419.5 53.0 172.7 17.4 630 316.9 234.0 11 4 6.6 3.875 110 2650.8 68.1 119.7 51.7 Gail 100.7 0.1 0.3 1.0 0 03 1,304 562 1007.4 27.7 39 0 18.5 3.16 1.51 1.81 17.30 4.82 19.68 46 10 308.6 159.9 12.6 6.6 774 596.1 29.6 20.5 8.6 Hair and hide 37,614 3,212 22180.6 931.7 2358.4 385.2 Head and tail , lean and fat 2144.6 418.3 98.8 26 4 Shin and shank, lean and fat 11.782 8001.4 1150.5 398.2 107 3 Flank and plate, lean and fat 31.790 17051.8 9195.6 832.9 233 0 Rump, lean and fat 7.370 4113.4 1960.4 193.7 59 3 10.76 99.89 79.14 Chuck and neck, lean and fat 60.176 41216.4 5931.0 1851.0 538 6 Round, lean 40,376 29735.7 1688.9 1301.3 405 8 Round, fat 5,106 30,836 1305.0 3269.9 82.8 17.5 2.09 Loin, lean 21471.7 2778.3 956.8 293.3 55.50 Loin, fat 7,570 1230.8 5871.4 78.2 19.2 3 10 Rib, lean 16 360 11091.3 1774.1 478.9 149.7 27.81 0.89 Rib, fat 1,978 347.2 1503.7 20.9 5.5 Kidney, fat 1.576 86.1 1456.4 5.0 2.2 0 27 Skeleton of feet 6,144 2523.7 662.2 225.4 1441.2 260.81 Skeleton of head 8,247 3920.6 699.2 261.9 1745.4 311.08 Skeleton of tail 386 146.5 102.5 11.1 58.3 10 55 Skeleton of shin 5,046 1430 . 6 918.5 193.7 1480.5 260 78 Skeleton of shank 5.498 1569.7 1240.2 193.5 1437.4 276 38 Skeleton of flank and plate 5,124 2119.7 667.8 181.0 1146.9 209 16 Skeleton of rump 2,860 789.2 850.7 87.5 636.2 110.91 Skeleton of chuck and neck 13,682 5,442 4431.2 2366.0 506.8 3426.8 630 . 19 Skeleton of round (excl. marrow) 1521.4 1342.0 161.5 1412.1 251.64 Marrow from skeleton of round 578 67.4 502.0 1.2 2.1 0.34 Skeleton of loin 6,872 1891.0 1763.4 215.1 1762.5 303.33 Skeleton of rib 4,986 1,590 1398.6 837.2 180.6 1524.2 271.44 Hoofs and dewciaws 1065.0 7.3 82.6 23.2 1.86 Teeth 838 239.1 4.5 14.6 469.3 88.27 Table 39. — Steer 512. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 24,176 19328.5 13.3 742.9 191.0 5.56 Circulatory system 2'432 985.7 1167.0 41.6 13.7 1.34 Lean heart 1,555 1202.3 54.0 43.6 19.7 3.34 Respiratory system 3,881 2929.7 181.2 108.8 41.9 6.36 Fat from thoracic cavity 1.171 143.2 1001.5 3.1 1.7 0.18 Brain and spinal cord 666 479.9 74.0 11.2 12.5 2.56 Digestive and excretory system (partial) .... Offal fat 20.735 17,454 15278.4 1956.9 2133.2 15058 . 1 452.7 64.4 157.8 25.1 24.67 3.32 Heart and neck sweetbreads 511 206.4 252.5 7.7 3.8 0.86 Liver 4,416 212 3046.3 115.9 144.1 70.1 14.75 Gall 197.5 0.1 0.5 2.2 0.06 Spleen 1,255 968.2 29.7 35.1 16.8 3.00 Pancreas 736 422.4 196.1 15.4 9.8 2.02 Kidneys 1,074 1.766 829.5 73.4 22.3 11.3 2.18 Tongue, marketable 1172.3 237.1 45.6 15.9 2.84 Hair and hide 41.268 23189.7 1490.6 2701.8 480.0 19.40 Head and tail, lean and fat 4.412 2715.8 841.5 127.6 37.9 6.13 Shin and shank, lean and fat 12.706 8717.1 1191.8 424.8 117.8 20.46 Flank and plate, lean and fat 48,946 20515.2 22190.7 934.9 279.0 46.01 Rump, lean and fat 10.484 73,512 4675.7 4385.4 212.5 69.9 12.48 Chuck and neck, lean and fat 46450.8 13372.6 2077.5 675.6 110.27 Round, lean 43.408 31805.9 1978.1 1405.1 444.5 83.34 Round, fat 9,940 2189.8 7023.4 76.0 30.9 3.98 Loin, lean 32,062 15,308 16,908 5,398 21676.2 3539.6 986.2 292.4 54.51 Loin, fat 1913.0 12759.8 99.5 27.6 3.98 Rib, lean 11010.3 2527.8 501.7 151.5 26.55 Rib, fat 806.4 4338.2 45.3 11.4 1.89 Kidney, fat 4,740 212.5 4451.6 8.7 6.2 0.95 Skeleton of feet 7,016 9,665 416 2627.6 1004.3 252.4 1709.5 309.05 Skeleton of head 4169.7 1252.1 312.1 2166.9 406.32 Skeleton of tail 153.8 101.0 13.0 75.3 13.58 Studies In Animal Nutrition — III 61 Table 39. — Steer 512. Weights of Constituents in Samples, Grams — Cont. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Skeleton of shin 6,074 1649.6 1263.9 220.6 1821.2 327.27 Skeleton of shank 6.156 1968.6 1326.1 217.9 1401.6 252.03 Skeleton of flank and plate 7,788 2865.4 1640.2 236.2 1523.6 279.28 Skeleton of rump 3.264 772.9 1001.4 97.5 857.6 145.12 Skeleton of cnuck and neck 16.536 4758.2 3139.5 536.4 5045.1 899.23 Skeleton of round (excl. marrow) 7,430 2134.1 1986.3 209.7 1679.6 301 . 14 Marrow from skeleton of round 396 39.9 349.7 0.7 2.6 0.52 Skeleton of loin 8,748 2198.9 2134.8 261.3 2330.6 466.62 Skeleton of rib 6,938 2042.3 1132.6 241.1 1991.1 378.40 Horns 1.810 637.6 8.7 127.3 441.7 70.12 Hoofs and dewclaws 1.724 843.1 10.1 135.5 48.1 2.14 Teeth 710 141.4 5.6 15.3 452.4 85.42 Table 40. — Steer 513. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 25.680 19947.2 881.1 185.4 7.19 3,485 4,858 3.942 2704.4 156.8 93.2 31.7 5.47 3590.3 341.6 133 2 48.2 7.53 529.8 3299 . 7 14 4 7.3 0.95 748 510.1 115 9 12 9 12.0 3.07 Digestive and excretory system (partial) .... Offal fat 25,642 53,771 19310.7 3055 8 1973.7 49939.3 592.3 115.6 266.9 42 5 40.26 5.91 Heart and neck sweetbreads 1.334 538.0 674.3 18.7 11.0 2.64 Liver 5,920 37 4021.2 185.1 190.9 82.7 19.71 Gall 33.8 0.1 0.1 0.4 0.01 Spleen 1,114 842.7 50.6 32.7 14.0 2.70 Pancreas 873 437.5 302.6 16.5 9.5 2.11 Kidneys 1,015 45,286 5,390 17,798 115.774 772.1 58.9 25.4 11.5 2.23 Hair and hide 26069 . 3 4561.2 2344.9 426.1 24.45 Head and tail, lean and fat 3038.8 1417.1 131.5 41.5 6.90 Shin and shank, lean and fat 9924.0 4922.4 411.0 124.1 21.00 Flank and plate, lean and fat 33830.3 72079.7 1437.9 445.7 75.25 Rump, lean and fat 19,082 6025.1 11211.6 263.0 83.6 15.27 Chuck and neck, lean and fat 110,940 53117.0 41259.7 2387.4 697.8 128.69 Round, lean 50.782 19.108 33089.0 7280.6 1473.2 462.8 87.85 Round, fat 3394.2 14539 . 1 176.0 38.0 4.59 Loin, lean 44,510 49.928 26437.2 9612.8 1236.0 367.7 72.55 Loin, fat 4444.0 44182.3 182.7 57.4 8.99 Rib, lean 24,744 13754.9 6857.3 635.2 187.3 35.14 Rib, fat 23.608 4201 5 17813.4 96 3 32.6 4.96 Kidney, fat 14.490 566.9 13755.1 22 6 10.7 1.45 Skeleton of feet 7.598 2753.4 1128 8 267 6 1811.1 322.38 Skeleton of head 7,865 3201.6 742.5 259.6 1968.5 460.57 Skeleton of tail 297 116.7 58.7 10.0 51.2 9.00 Skeleton of shin 6.120 1792.6 1091.5 206.7 1672.3 294.43 Skeleton of shank 6.058 1582.6 1206.7 216.3 1792.6 317.02 Skeleton of flank and plate 7.438 3,464 15.526 3043 . 8 1121.7 243.8 1519.2 307.86 Skeleton of rump 845.8 1014.5 104.2 855.2 162.22 Skeleton of chuck and neck 4969.6 2760.7 544.7 3912.9 785.31 Skeleton of round (excl. marrow) 6,564 480 1625.0 2011.1 194.8 1625.1 286.58 Marrow from skeleton of round 41.6 431.8 0.8 3.9 0.67 Skeleton of loin 8,536 2149.9 2543.6 258.2 2081.3 401.11 Skeleton of rib 7.096 1884.6 1528.4 238.6 2073.2 392.27 Horns 2.144 790 1 16 4 138 3 485.0 89 36 Hoofs and dewclaws 2.180 914.1 19.4 195.2 64.5 5.21 Teeth 874 278.8 9 6 15.8 459.5 87.04 62 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 41. — Steer 515. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus 27,856 22036.1 896.1 165.5 6.69 Circulatory system 5,787 2432.4 2808.4 73.8 27.8 4.98 Respiratory system 3,653 2789.2 198.8 98.6 35.4 6.83 Brain and spinal cord 728 509.9 117.1 12.0 11.9 2.88 Digestive and excretory system 36,311 24025.9 6794.2 733.1 309.0 58.82 Offal fat 29,877 2250.3 26976.2 84.0 37.1 4.48 Hair and nide 49.943 27969.6 6267.9 2417.2 927.4 29.47 Head and tail, lean and fat 5.918 3169.8 1807.2 143.0 43.3 7.70 Shin and shank, lean and fat 16,676 9565.5 4461.3 417.2 119.4 20.51 Flank and plate, lean and fat 87.138 27115.6 52220.7 1185.1 338.1 61.87 Rump, lean and fat 15,810 5586.6 8569.7 238.1 75.3 13.44 Chuck and neck, lean and fat 88,134 46944.6 27399.1 2076.4 627.5 111.05 Round, lean 42,942 29067.9 4746.0 1332.1 396.8 76.01 Round, fat 19.058 3339.3 14807.3 149.4 41.0 4.96 Loin, lean 41,620 27036.8 6323.3 1248.2 413.7 75.33 Loin, fat 38,324 4053.2 33197.4 157.9 44.1 6.52 Rib, lean 19.016 11634.6 3976.8 527.9 161.5 28.90 Rib, fat 16,282 1476.9 14374.9 64.2 21.8 3.26 Kidnev, fat 9.922 491.2 9369.6 17.7 '8.0 1.49 Skeleton of feet, head and tail 18.179 7520.3 2029.0 551.2 4270.3 739.16 Skeleton of shin and shank 13,900 3852.1 3761.8 423.3 3429.1 560.45 Skeleton of flank and plate 6,368 2687.0 1016.6 217.2 1132.2 198.24 Skeleton of rump 4,074 1213.8 104.7 126.1 949.8 165.85 Skeleton of chuck and neck 14,528 4431.0 2254.3 484.2 4295.6 617.88 Skeleton of round 6.344 1499.9 1866.0 191.8 1825.0 310.22 Skeleton of loin 7,784 1969.9 2196.6 231.3 2040.7 350.05 Skeleton of rib 6,464 1686.2 1303.1 211.4 2015.2 282.74 Horns 1,804 733.2 10.7 101.3 427.1 76.18 Hoofs and dewclaws 1,893 1017.5 10.0 139.4 34.5 1.36 Teeto 786 214.8 6.8 14.0 499.1 86.40 Table 42. — Steer 523. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 15,287 3,044 12309.1 426.5 100.7 3.36 Circulatory system 1615.0 1090.5 51.3 18.2 3.35 Respiratory system 3,371 2652.3 136.7 87.8 33.2 6.37 Brain and spinal cord 797 546.7 140.4 12.9 12.1 2.86 Digestive and excretory system 24,667 18548.1 2548.1 540.2 194.4 38.97 Hair and hide 33,097 20547.0 380.6 1854.4 340.9 16.88 Head and tail, lean and fat 3,100 2113.2 358.8 95.9 28.6 4.77 Shin and shank, lean and fat 8,684 6233.9 524.2 295.4 79.5 14.68 Flank and plate, lean and fat 26,984 16058.2 6184.7 726.7 199.4 37.51 Rump, lean and fat 5,418 2943.6 1585.8 137.2 40.5 8.34 Cnuck and neck, lean and fat 50.320 35668.8 5176.9 1502.6 463.5 86.05 Round, lean 33,900 26078.9 783.1 1069.6 353.2 68.48 Loin, lean 25,834 18020.3 2688.3 782.0 237.9 44.43 Rib, lean 12.032 8456.3 1118.6 375.3 111.5 21.42 Round, fat 4,556 1346.0 2743.8 72.0 16.3 2.19 Loin, fat 6,376 1051.9 4969.1 55.0 17.9 2.87 Rib, fat 1,522 358.3 989.0 23.0 6.0 ' 0.90 Kidney, fat 3,110 163.6 2883.8 14.5 5.6 0.47 Offal fat 7,915 1220.3 6433.6 39.2 16.2 2.37 Skeleton of feet, head and tail 13.120 5805.7 1065.3 467.5 2990.8 542.51 Skeleton of shin and shank 8,862 2611.1 1804.7 299.6 2449.0 454.80 Skeleton of flank and plate 3,882 1741.8 415.8 134.7 754.8 134.59 Skeleton of rump 1,770 478.5 358.6 57.9 523.8 196.11 Skeleton of chuck, and neck 9,786 3314.1 1261.8 360.2 2876.5 527.56 Skeleton of round 4,630 1845.1 1231.5 89.9 1950.2 177.79 Skeleton of loin 5,322 1483.4 1260.6 167.4 1438.5 265.09 Skeleton of rib 3,844 1200.0 481.2 138.1 1219.1 221.76 Horns 1,167 539.5 7.3 65.5 218.7 39.87 Hoofs and dewclaws* 1.063 575.1 7.8 76.5 21.6 3 131 Teeth 766 201.6 5.9 16.1 432.0 81.78 ♦This sample was lost before analysis. The analysis of the hoofs and dewclaws of four animals of the same Group was used. Studies In Animal Nutrition — III 63 Table 43. — Steer 524. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 17,019 2.953 13956.6 474.3 127.0 3.74 1833.5 662.0 66.2 22.7 3.90 3455 758 2682.3 89.9 96.5 37.1 6.15 555.0 77.7 12.1 10.8 2.56 Digestive and excretory system (partial) 17,924 5.007 13280.6 1255.2 1760.3 3477.1 412.8 38.1 158.6 16.0 26.89 1.75 Heart and neck sweetbreads 541 406.4 40.3 14.1 10.1 2.50 3,019 2108 5 100.1 92.0 42.5 9.87 Gall 229 217 2 0.2 0.4 1.8 0.07 Spleen 596.2 13.8 20.8 10.1 1.97 435 285.5 69.1 11 1 5.4 1.24 Kidneys 766 588.5 43.9 18.6 8.9 1.59 30,092 17832.2 545.6 1895.8 474.6 17.45 Head and tail, lean and fat 3.364 2229.0 441.6 105.2 34.5 6.02 Shin and shank, lean and fat 8.674 6335.2 485.8 253.7 84.5 14.40 Flank amd plate, lean and fat 19,788 4,030 46,386 37,714 2,526 24.200 12607.1 3041.8 633.2 188.4 30.47 Rump, lean and fat 2563.5 691.0 119.2 39.2 7.05 Cnuck and neck, lean and fat 33655.4 2918.6 1465 . 8 452.3 80.71 Round, lean 28988.5 1024.3 1228.3 393.4 72.41 Round, fat 915.9 1262.0 56 . 5 10.6 1.34 Loin, lean 17588.6 1099.7 801.0 255.8 46.95 Loin, fat 2,444 13,144 459.7 1791.9 30.2 8.6 1.30 Rib, lean and fat 9238.3 1044.8 435.1 129.1 23.92 Kidney, fat 766 87.6 644.9 3.4 1.4 0.24 Skeleton of feet 6,010 2434.5 781.4 215.9 1473.4 262.94 Skeleton of head and tail 8,318 3820.5 864.2 254.0 1985.3 354.60 Skeleton of snin and shank 10,262 3240 . 8 1982.3 345.3 2659.9 474.00 Skeleton of flank and plate 5,926 2569.4 905.0 190.3 1135.7 198.22 Skeleton of rump 2,424 12,896 5,878 6,586 831 6 520.0 72.7 542.9 98.34 Skeleton of chuck and neck 5197.0 2043.2 427.6 2809.5 514.03 Skeleton of round 1825.5 1665.9 156.7 1334.3 238.82 Skeleton of loin 1926.5 1771.7 187.4 1600.8 279.64 Skeleton of rib 5,310 1.227 1897.5 984.2 157.7 1324.9 235.18 Horns* 506.9 9.1 78.7 237.5 44.07 Hoofs and dewclaws 1,494 806 750.6 12.4 112.8 47.7 3.27 Teeth 217.4 8.9 15.4 466.4 88.56 ♦This sample was lost before analysis. The analysis of (he horns of a steer of the same age was used. Table 44. — Steer 525. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 13,614 10961.2 400.5 89.9 3.58 Circulatory system 1,320 846.0 324.0 22.5 9.4 1.59 Respiratory system 1,658 1303.4 43.7 46.8 18.4 3.20 Brain and spinal cord 711 505.2 99.1 12.4 10 7 2.63 Digestive and excretory system 23,001 17820.9 2103.2 459.8 203.1 35.95 Hair and hide 27,813 17354.2 138.5 1592.0 370.2 15.85 Head and tail, lean and fat 2,788 2072.0 364.7 51.8 27.5 4 40 Shin and shank, lean and fat 7,596 5533.6 404.9 253.5 73.3 13.44 Flank and plate, lean and fat 18,762 11412.4 3790.7 539.0 155.4 27.96 Rump, lean and fat 4.154 2532.3 869.2 114.7 35.9 6.73 Chuck and neck, lean and fat 35,824 25489.1 3024.6 1119.5 340.0 63.77 Round, lean 27,524 21202.8 661.4 858.8 292.9 56.42 Loin, lean 18.710 13964.0 649.1 605.6 196.1 37.42 Rib, lean 11,666 8226.2 1010.2 368.8 114.0 21.12 Round, fat 1,962 641.3 1114.9 31.4 8.5 1.10 Loin, tat 3.758 826.8 2615.2 41.3 11.1 1.50 Rib, fat 664 186.5 409.1 10.7 4.0 0.61 Kidney, fat 1,258 84.6 1134.5 5.9 2.0 0.25 Offal tat 4,961 1043.2 3504.5 63.8 14.5 2.53 Skeleton of feet, head and tail 10,782 4638.5 1077 1 402.8 2352 . 9 4*0.32 Skeleton of shin and shank 7,014 2127.6 1342.1 244.4 2071.0 293.89 Skeleton of flank and plate 3,454 1540.9 374 9 116.2 744.9 103.24 Skeleton of rump 1,542 474.9 367.3 49.9 379.5 65.01 Skeleton of chuck and neck 8,450 2885.3 1665.8 261.3 1935.1 363.43 Skeleton of round 4,046 1157.2 1322.5 107.3 881.3 114.34 Skeleton of loin 3,928 1123.2 994.1 118.6 959 8 144.47 Skeleton of rib 3,888 1233.9 701.8 139.7 961 9 169.21 Horns 1,298 634.4 7.1 73.2 217.0 40 77 Hoofs and dewciaws 940 487.9 5.4 74.2 11.8 1.26 Teeth 690 159.6 7.3 13.1 418 9 80.99 64 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 45. — Steer 526. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus 18,957 15151.6 585.2 147.3 4 74 Circulatory system 2,413 1527.8 517.2 56.5 19.6 3.50 Respiratory system 3,797 2909.1 124.9 106.3 38.9 5.92 Fat from the thoracic cavity 1.585 340.3 1159.3 11.9 4.5 0.54 Brain and spinal cord 660 483.2 67.7 11.5 11.4 2.77 Digestive and excretory system (partial) 20,541 14971.1 2510.5 437.5 157.8 26.09 Offal fat 11,551 1500.7 9719.5 47.4 20.2 2.08 Heart and neck sweetbreads 439 265.5 106.3 9.9 6.3 1.53 Liver 3,531 2398.3 150.2 114.7 51.9 12.29 Gall 143 132.1 0.2 0.3 1.7 0.04 Spleen 831 652.3 12.4 23.0 12.9 2.47 Pancreas 498 316.0 90.7 11.2 5.8 1.40 Kidneys 922 678.2 83.6 21.7 9.9 1.83 Hair and hide 35,732 20663.1 2041.7 2116.4 514.5 17.87 Head and tail, lean and fat 3.616 2234.4 710.2 105.2 30.1 5.13 Shin and shank, lean and fat 11,644 8261.4 877.6 384.4 104.8 19.10 Flank and plate, lean and fat 39.524 19561.2 13931.8 920.9 268.0 49.41 Rump, lean and fat 8,594 4611.8 2576.6 211.7 63.3 11.9 Chuck and neck, lean and fat 61.228 40068.2 8878.1 1789.7 555.3 101.64 Round, lean 44,614 29839.6 5303.3 1472.3 460.0 85.21 Round, fat 5.016 1136.9 3460.1 83.8 14.3 1.81 Loin, lean 31,440 22620.5 1679.2 1009.2 320.1 59.74 Loin, fat 11,634 1687.9 9298.9 98.9 24.2 3.49 Rib, lean 17,264 12022.8 1728.8 531.2 158.8 28.49 Rib. fat 3,720 641.4 2781.4 43.2 9.6 1.64 Kidney, fat 3,224 292.6 2831.7 11.0 4.4 0.68 Skeleton of feet 6.138 2358.5 850.3 226.0 1440.9 267.25 Skeleton of head and tail 9.165 4249.2 1173.6 274.8 1925.0 351.20 Skeleton of shin and shank 11.612 3228.4 2333.7 377.9 3333.0 608.24 Skeleton of flank and plate 6,588 2645.4 1084.8 209.2 1339.5 249.88 Skeleton of rump 2.838 832.8 726.8 88.2 672.8 121.38 Skeleton of chuck and neck 13,842 4749.6 2415.3 462.2 3405.3 620.40 Skeleton of round 6,352 17! S. 7 1922.9 171.1 1478.2 274.28 Skeleton of loin 6 344 1995.9 1939.1 193.4 1595.6 293.06 Skeleton of rib 5.690 1804.4 1142.3 188.6 1495.7 270.73 Horns 1.427 589.6 10.6 91.5 276.2 51.26 Hoofs and dewclaws 1.875 1019.9 11.7 136.8 39.9 1.59 Teeth 782 173.2 10.0 15.2 480.7 91.39 Table 46. — Steer 527. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood . 27,382 4,903 21615 6 899 0 208 7 6 30 Circulatory system 2645.2 1608.1 94.5 31.1 5.59 Respiratory system 4.326 3172.3 310.9 117.5 44.3 7.18 Fat from thoracic cavity Brain and spinal cord 3.988 701 412.4 490.4 3495.3 98.6 13.4 11.1 5.1 10.8 0.60 2.58 Digestive and excretory system (partial) Offal fat 23,818 48,517 1.037 15915.6 2617.0 4412.8 45247.0 494.7 88.8 169.1 57.3 28.34 6.31 Heart and neck sweetbreads 379.8 574.0 12.3 7.6 3.19 Liver 5.720 3882.3 198.6 188.4 90.1 10.52 Spleen 1,226 941.5 26.6 35.6 15.7 2.91 Pancreas 849 353.0 393.1 12.4 7.5 1.72 Kidneys 1,244 937.3 103.0 27.5 12.5 2.40 Hair and hide 46.240 25189.2 5483.6 2458.6 633.0 25.89 Head and tail, lean and fat 5,018 2705.8 1537.3 120.9 34.7 6.17 Shin and shank, lean and fat 17,358 118,978 24.020 9807.8 4582.2 446.1 130.9 22.57 Flank and plate, lean and fat 32362.0 77919.9 1305.2 371.2 70.20 Rump, lean and fat 6792.1 15123.5 296.2 75.7 14.17 Chuck and neck, lean and fat 112,440 52601.7 44206.9 2282.5 693.8 122.56 Round, lean 51.396 33977.4 7160.0 1538.3 444.1 89.94 Round, fat 21.466 3462.3 17027.5 161.6 43.8 4.72 Loin, lean 50,140 30681.7 10276.7 1402.4 425.7 80.73 Loin, fat 52,724 5018.8 46540.0 188.2 67.0 8.96 Rib, lean 25.860 14218.9 7266.1 658.7 203.5 36.46 Rib, fat 24.278 2297.4 21415.1 98.8 32.3 3.88 Kidney, fat 18,964 1028.4 17679.6 35.5 19.3 2.65 Skeleton of feet 7.442 2774.4 1199.3 250.9 1761.1 286.44 Skeleton of head and tail 8'S22 3717.2 2189.8 279.2 1913.7 324.30 Skeleton of shin and shank 13.136 3580.4 2794.8 413.7 3726.6 597.69 Skeleton of flank and plate 6 082 2392.3 1121.3 182.2 1187.9 207.03 Skeleton of rump 3.260 810.0 1000.5 96.7 780.0 122.15 Skeleton of chuck and neck 14,870 4172.2 3653.6 458.3 3723.0 586.47 Skeleton of round 6.446 1253.5 2188.7 170.3 1742.5 319.98 Skeleton of loin 7.140 1850.7 1840.8 225.0 1924.2 368.14 Skeleton of rib 6,546 1762.2 1510.4 197.8 1873.6 360.23 Horns 1.266 451.5 9.8 88.0 250.8 46.31 Hoofs and dewclaws 2.174 958.3 20.9 193.8 44.8 3.15 Teeth 872 180.5 12.5 17.0 552.7 120.61 Studies In Animal Nutrition — III 65 Table 47. — Steer 531. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 9,457 7743.6 268.7 66.5 3.78 Circulatory system 1,971 1118.0 577.9 38.8 13.3 2.48 Respiratory system 1,915 1500.2 54.0 51.3 20.2 3.91 Brain and spinal cord 573 418.2 77.3 10.1 8.6 2.08 Digestive and excretory system (partial) 11,807 9106.6 1089.8 233.4 79.0 14.29 Offal fat 2.899 751.7 2049.0 18.0 8.4 1.33 Heart and neck sweetbreads 472 320.9 69.8 11.8 8.1 2.02 Liver 2,205 1582.9 42.9 64.4 30.5 7.08 Gall 86 78.8 0.2 0.8 0.04 Spleen 481 361.1 20.2 14.2 6.9 1.40 Pancreas 297 199.2 43.9 7.5 3.7 0.86 Kidneys 506 379.9 41.9 11.5 5.3 1.08 Hair and Hide 16,693 10627.6 135.4 957.0 189.6 12.52 Head and tail, lean and fat 1,722 1157.5 242.3 50.3 15.0 2.67 Shin and shank, lean and fat 5,480 3934.2 259.5 197.1 58.7 10.30 Flank and plate, lean and fat 10.854 6553.0 1865.9 331.6 109.1 19.10 Rump, lean and fat 2,506 1553.5 453.9 74.7 24.5 4.43 Chuck and neck, lean and fat 26,902 18808.4 2099.7 880.8 278.2 52.73 Round, lean 21.496 16145.9 393.6 703.4 236.9 44.71 Round, fat 1,490 424.3 904.2 22.7 7.1 1.06 Loin, lean 14,078 10278.5 602.3 466.3 154.9 28.72 Loin, fat 2.264 560.5 1468.5 31.4 9.9 1.70 Rib, lean and fat 6,612 4597.7 500.0 220.7 69.6 12.56 Kidney fat 726 44.0 655.3 4.3 1.7 0.23 Skeleton of feet 3,762 1496.3 540.0 121.6 927.0 166.73 Skeleton of head and tail 4.842 2376.4 390.8 147.3 1009.5 175.72 Skeleton of shin and shank 5,998 1903.2 1100.4 176.2 1644.4 306.68 Skeleton of flank and plate 2,546 1190.8 269.1 82.0 519.3 88.19 Skeleton of rump 1.060 333.0 225.1 34.7 270.7 48.47 Skeleton of chuck and neck 6.098 2286.3 976.6 215.6 1434.6 255.32 Skeleton of round 3,640 1275.1 943.2 97.5 777.6 138.54 Skeleton of loin 2,834 881.1 621.1 93.5 722.5 126.71 Skeleton of rib 2.280 738.4 381.7 85.2 575.2 104.58 Hoofs and dewclaws 790 411.6 6.5 59.9 15.5 0.99 Teeth 426 116.8 3.8 8.5 237.7 45.26 Table 48. — Steer 532. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 18,752 15090.3 562.0 122.3 6.19 Circulatory system 4,843 2320.4 1949.6 77.7 27.4 5.09 Respiratory system 3,870 3001.2 132.1 102.0 40.7 7.47 Brain and spinal cord 643 465.9 93.0 10.7 10.4 2.52 Digestive and excretory system partial 21,741 15805.9 3039.2 412.6 153.9 29.13 Offal fat 23,697 2263.5 21044.1 57.6 27.5 3.79 Heart and neck sweetbreads 441 218.1 169.6 7.2 4.4 1.05 Liver 5,694 4066.6 135.2 161.9 78.3 17.48 Gall 185 170.7 0.4 1.9 0.08 Spleen 884 660.3 44.5 25.5 11.1 2.32 Pancreas 630 353.8 174.4 12.8 7.4 1.85 Kidneys 868 618.4 95.3 21.9 8.8 1.85 Hair and hide 33,988 20244.6 2324.4 1775.9 350.8 24.13 Head and tail, lean and fat 4,280 2571.8 935.1 114.5 34.4 6.21 Shin and shank, lean and fat 12.008 8109.7 1320.6 379.5 112.0 20.05 Flank and plate, lean and fat 44,636 19529.6 18635.1 998.5 292.4 51.78 Rump, lean and fat 8.058 3910.4 2891.5 189.0 57.7 10.72 Chuck and neck, lean and fat 66.204 40908.8 12261.0 1926.5 595.2 103.29 Round, lean 38.064 27369.5 1988.1 1257.6 405.4 76.13 Round, fat 6,064 1316.7 4264.3 67.5 17.9 2.55 Loin, lean 36,136 24719.2 3466.5 1175.1 362.8 66.49 Loin, fat 14,954 1872.2 12472.4 100.8 28.6 4.93 Rib, lean 17,356 11677.1 1993.0 547.8 169.9 30.55 Rib, fat 6.194 967.6 4885.7 52.8 16.6 2.48 Kidney, fat 11,734 383.8 11174.1 17.6 10.0 2.58 Skeleton of feet 6,490 2536.3 031 3 234 . 1 1547.5 273.94 Skeleton of head and tail 7.120 3304.3 910.4 212.5 1512.7 270.13 Skeleton of shin and shank 10,756 3107.4 2325.1 464.2 2512.3 472.94 Skeleton of flank and plate 5,478 2132 7 1037.9 165.8 819.4 149.99 Skeleton of rump 2.282 659 . 5 633.1 73.1 517.8 93.20 Skeleton of chuck and neck 13,014 3911 0 3071 4 438.2 3126.9 506.24 Skeleton of round 5.024 1511.0 1914.0 141 4 1169.5 208.82 Skeleton of loin 6.246 17.51 9 1827.4 193.2 1377.4 254.46 Skeleton of rib 5,222 1826.1 1175.4 154.1 1163.5 209.35 Horns 228 124.8 1.2 14 4 17.3 3.28 Hoofs and dewclaws 1,406 730.2 9.1 107.0 27.4 2.15 Teeth 494 153.2 4.4 10.6 25 V 4 50.16 66 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 49 . — Steer 538 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 7,219 1,670 1,825 487 10,290 3,452 481 5948.5 7.0 197.4 56 7 2.02 926.6 479.3 32.3 10 6 1 90 Respiratory system Brain and spinal cord Digestive and excretory system (partial) . . . Offal fat 1474.6 362.0 7822.7 731.2 37.5 57.2 1009.8 2639.2 45.7 7.8 217.2 18.8 18.6 7.3 89.2 9.4 8 0 3.63 1.69 15.54 1.90 1.95 6.23 0 09 335.8 70.2 11.2 1,978 122 1394.1 37.9 58.5 27 7 Gall 110.9 0.1 0.3 1 3 331 259.6 5.1 9.5 4.7 0.90 0.73 208 144.9 24.2 5.3 3.1 487 356.6 54.9 11.2 5 2 1.02 15,342 1,496 9870.9 205.9 853.6 163.1 9 67 Head and tail, lean and fat. 991.9 231 0 40.4 13 4 2.21 7 50 Shin and shank, lean and fat 4,190 11,036 3008.9 241.9 138.0 42 0 Flank and plate, lean and fat 6367.8 2589.7 306.1 94.0 17.66 Rump, lean and fat 2,096 1252.3 442.6 55.9 18.7 3.23 Chuck and neck, lean and fat 22.284 15261.6 2665.8 662.1 207.5 38 77 Round, lean 16,324 1,702 12401.5 440.6 515.7 178.4 32.97 Round, fat 519.6 999.6 29.1 7.6 1.16 Loin, lean 12.732 9312.3 715.2 396.9 134.3 25 85 Loin, fat 2.360 5,196 470.5 1713.4 23.1 6.9 1.27 Rib, lean 3683.0 423.5 159.8 52.8 9 92 Rib, fat '426 128.6 240.4 6.2 3.1 0 49 Kidney, fat 622 42.0 565.8 2.1 1.1 0.21 Skeleton of feet 3,166 4,001 135 1350.6 491.6 104.3 589.4 105.81 Skeleton of head 1991.7 311.8 116.7 863.4 163.60 Skeleton of tail 70.8 19.1 4.3 15.2 2.63 Skeleton of shin 2,208 2,608 2.144 732 632.3 526.1 62.4 554.4 86.64 Skeleton of shank 792.0 585.2 88.4 616.9 95.71 Skeleton of flank and plate 1054.6 311.8 68.6 322.6 52.46 Skeleton of rump 250.9 173.8 22.3 162.7 27.79 Skeleton of chuck and neck 5,412 2.556 2064.1 917.4 180.6 1297.2 192.40 Skeleton of round 765.8 823.7 61.9 526.5 81.20 Skeleton of loin 2,696 942.7 625.4 79.3 573.1 112.23 Skeleton of rib 2,154 250 783.3 357.8 68.5 487.2 86.83 Horns 137.3 1.3 14.5 24.9 4.74 Hoofs and dewclaws* 635 423.1 3.0 34.0 6.5 0.43 Teeth f 240 68.83 2.7 4.7 130.0 24.48 Table 50 . — Steer 540 . Weights of Constituents in Samples, Grams. Blood 6,967 1,436 1,501 512 5732.5 70.5 195.5 50.4 1.81 Circulatory system 834.8 398.2 29.3 9.7 1.82 Respiratory system 1194.9 34.7 38.9 15.5 2.97 Brain and spinal cord 377.1 61.1 8.3 8.1 1.92 Digestive and excretory system (partial) Offal fat 9,280 2.307 6982.2 582.4 998.2 1639.3 185.1 12.6 65.2 6.5 11.97 1.08 Heart and neck sweetbreads 408 280.7 63.0 9.2 6.2 1.52 Liver 1,593 58 1124.5 28.0 45.2 22.5 4.86 Gall 54.4 0.04 0.1 0.7 0.02 Spleen 331 257.5 4.5 10.3 4.7 0.90 Pancreas 180 130.0 17.3 4.6 2.6 0.62 Kidneys 363 261.9 38.1 8.6 3.9 0.81 Hair and hide 12.994 8399.6 305.8 666.2 163.2 8.71 Head and tail, lean and fat 1.274 866.2 170.8 34.9 11.3 2.13 Shin and shank, lean and fat 3,762 2793.8 1*1.9 127.8 39.1 6.96 Flank and plate, lean and fat 8,824 1,964 5375.9 1629.0 266.6 77.9 13.94 Rump, lean and fat 1198.9 384.8 54.1 18.1 3.26 Chuck and neck, lean and fat 17,978 13,456 12783.8 1500.6 532.3 177.3 32.00 Round, lean 1018". 9 303.4 431.7 145.6 28.26 Round, fat 810 225.4 504.7 11.0 3.1 0.49 Loin, lean 10,700 2.308 7869.3 448.0 340.2 115.0 21.51 Loin, fat 452.3 1700.8 22.1 6.7 1.34 Rib, lean and fat 5,046 3503.4 488.3 156.2 51.1 9.14 Kidney fat Skeleton of feet 682 2.784 91.4 1225.2 544.3 337.6 7.9 91.5 1.3 580.2 0.24 99.47 Skeleton of head 3,682 138 1903.2 259.8 17.2 101.4 788.6 149.34 Skeleton of tail 76.4 3.1 13.9 2.44 Skeleton of shin 1,952 618.5 432.5 68.3 472.0 89.03 Skeleton of shank 2.304 763.4 558.2 68.4 486.2 86.63 Skeleton of flank and plate Skeleton of rump 1,862 622 990.4 219.0 225.0 127.2 62.9 18.9 225.1 201.9 54.04 26.25 Skeleton of chuck and neck 4.896 2021.3 721.7 159.8 962.8 172.93 Skeleton of round 2.350 829.2 643.2 59.3 392.6 65.82 Skeleton of loin 2,550 887.6 698.2 71.0 489.8 91.16 Skeleton of rib 1,824 823.3 342.4 60.1 417.0 74.58 Horns 304 165.0 1.8 16.6 37.9 7.33 Hoofs and dewclaws* 481 320.5 2.2 25.8 4.9 0.32 Teethf 278 79.7 3.1 5.5 150.6 28.36 *Hoofs and dewclaws of steer 538 and steer 540 were analyzed together. fTeeth of steer 538 and steer 540 were analyzed together. Studies In Animal Nutrition — III 67 Table 51 . — Steer 541 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 12,470 10239.4 11.1 345.8 84.7 3.37 Circulatory system 2,646 1282.3 1055.2 43.6 15.1 2.78 Respiratory system 2,387 1890.9 47.2 65.4 24.8 5.08 Brain and spinal cord 568 411.6 77.3 9.4 8.7 2.11 Digestive and excretory system (partial) 16,313 11628.7 2398.0 330.2 121.7 21.86 Offal fat 11,009 1289.6 9511.6 32.3 13.8 2.86 Heart and neck sweetbreads 740 484.3 141.7 16.9 10.8 2.72 Liver 3,832 2664.8 95.9 121.0 55.2 13.37 Gall 202 179.8 0.4 0.6 2.4 0.12 Spleen 596 463.0 10.8 17.1 7.9 1.66 Pancreas 390 233.7 91.4 7.9 4.7 1.11 Kidneys 645 473.9 63.9 14.3 6.9 1.38 Hair and hide 26,574 16057.3 970.0 1512.6 292.3 14.62 Head and tail, lean and fat 2,062 1370.6 299.4 60.6 21.0 3.38 Shin and shank, lean and fat 6.830 4693.9 616.1 199.6 65.3 12.16 Flank and plate, lean and fat 24,910 12405.7 8529.4 604.6 178.6 31.64 Rump, lean and fat 4.454 2250.7 1459.8 105.6 31.8 6.10 Chuck and neck, lean and fat 40,480 26418.5 5979.7 1190.9 369.6 66.79 Round, lean 27,000 19957.1 908.8 901.8 294.0 56.16 Round, fat 3,854 817.1 2828.8 38.0 10.9 1.77 Loin, lean 23,416 16848.3 1301.7 757.0 245.9 46.36 Loin, fat 8,088 1202.0 6521.7 63.1 16.5 3.15 Rib, lean 11,588 7956.0 1209.0 362.2 113.1 21.67 Rib, fat 2,434 432.9 1850.8 26.3 7.6 1.44 Kidney, fat 6,056 272.2 5713.8 12.2 5.2 1.21 Skeleton of feet 4,506 1872.3 578.6 154.9 973.7 168.66 Skeleton of head 5,400 2730.2 381.3 161.7 1085.8 197.48 Skeleton of tail 206 98.7 40.4 6.5 22.4 3.90 Skeleton of shin 2,946 851.4 690.0 91.4 799.5 111.89 Skeleton of shank 3,744 1222.7 778.5 109.8 1039.5 141.45 Skeleton of flank and plate 2.864 1487.0 439.0 65.2 339.1 51.52 Skeleton of rump 1,056 319.9 258.3 32.3 268.7 51.25 Skeleton of chuck and neck 7,296 2542.6 1341.7 245.7 1832.0 264.26 Skeleton of round 3,250 876.1 1054.9 105.7 631.5 136.86 Skeleton of loin 3,916 1344.9 968.6 125.9 786.4 152.92 Skeleton of rib 3,162 1001.5 682.3 101.0 807.5 117.82 Horns 468 248.3 3.1 25.0 64.8 12.59 Hoofs and dewclaws 869 505.1 5.2 52.3 10.9 0.53 Teeth 304 142.4 2.9 4.4 123.8 23.79 Table 52 . — Steer 547 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 8,711 7042.8 256.3 62.5 2.53 Circulatory system 1,624 949.1 451.7 31.9 11.1 2.01 Respiratory system 1,868 1474.0 52.6 47.6 21.5 3.64 Brain and spinal cord 459 354.2 41.1 7.1 6.7 1.68 Digestive and excretory system (partial) 11,978 8883.2 1309.1 256.1 124.8 23.00 Offal fat 3,879 700.4 3059.8 23.5 10.0 2.06 Liver 2,851 2051.7 73.6 84.3 38.9 10.43 Spleen 391 301.4 8.7 11.8 5.3 1.17 Pancreas 200 142.8 20.2 4.9 2.9 0.75 Kidneys 450 329.7 38.5 11.5 5.2 1.08 Hair and hide 14,618 9435.3 267.4 779.1 189.3 10.52 Head, tail, shin and shank, lean and fat 8,590 5906.7 1053.2 242.1 77.1 14.86 Flank and plate, lean and fat 14,226 7829.0 3915.4 355.9 112.8 20.91 Rump, lean and fat 2,256 1227.5 622.1 55.6 18.5 3.54 Chuck and neck, lean and fat 23,636 15975.1 2954.7 655.2 217.0 39.47 Round, lean 17,092 12723.3 612.6 550.9 184.4 35.89 Round, fat 2,150 642.3 1304.3 28.9 8.5 1.46 Loin, lean 13,576 9708.5 831.4 432.0 139.8 27.02 Loin, fat 3,972 837.4 2869.7 39.2 11.9 2.26 Rib, lean 6,560 4579.0 618.8 199 6 64.2 1.18 Rib, fat 1,102 276.4 730.5 15.9 5.5 0.83 Kidney, fat 1,630 128.6 1468.7 5.1 2.2 0.24 Skeleton of feet, head, tail, shin and shank.. 11,966 5383.9 1685.5 399.8 2272.2 418.93 Skeleton of flank and plate 1.958 1044.4 262.8 60.9 230.5 42.45 Skeleton of rump 760 :w, o 121.0 25.0 180.0 33.77 Skeleton of chuck and neck 5,120 2252.3 715.9 164 9 1035.6 186.62 Skeleton of round 2,488 902 6 711.8 62.8 430.8 81.06 Skeleton of loin 3,132 1233.5 629.5 91.1 631.3 120.30 Skeleton of rib 2,172 902.1 368.2 72.1 423.0 78.89 Horns, hoofs and dewciaws 737 392.2 9.8 53.9 18.2 1.62 Teeth 310 100.8 2.1 6.8 158.7 29.61 68 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 53 . — Steer 548 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 4,603 3737.5 142.7 32.9 0 97 Circulatory system 753 528.5 94.7 19.6 6.4 1.18 Respiratory system 1,084 833.4 33.9 31.5 2.0 2.19 Brain and spinal cord 526 392.6 58.2 8.6 7.5 1.84 Digestive and excretory system (partial) 5,357 4230.5 228.6 127.8 7.1 9.59 Offal fat 845 444.2 310.8 12.4 6.1 1.03 Heart and neck sweetbreads 168 129.5 10.2 4.7 2.7 0.66 Liver 1,104 781.8 20.3 36.3 15.3 3.71 Spieen 215 166.1 2.9 6.7 2.9 0.60 Pancreas 85 64.1 5.3 2.4 1.2 0.28 Kidneys 353 277.3 12.0 8.8 4.4 0.87 Hair and hide 8,358 5466.8 78.0 466.8 112.8 6.52 Head and tail, lean and fat 967 693.2 87.0 25.9 9.2 1.69 Shin and shank, lean and fat 2,462 1842.7 109.5 80.4 24.6 4.31 Flank and plate, lean and fat 4,802 3379.8 366.3 159.4 46.5 8.36 Rump, lean and fat 1,042 756.3 60.4 34.9 11.5 2.13 Chuck and neck, lean and fat 11,136 8421.6 390.4 345.2 117.3 20.82 Round, lean 9,208 6953.3 154.7 309.7 103.0 19.71 Round, fat 520 279.9 149.4 14.2 3.6 0.45 Loin, lean 5,668 4269.4 114.5 186.6 63.9 11.85 Loin, fat 384 157.2 175.0 7.6 2.3 0.37 Rib, lean and fat 3,180 2412.0 87.1 99.6 34.8 6.20 Kidney, fat 284 78.5 182.4 3.9 1.4 0.17 Skeleton of feet 2,496 1138.7 345.1 83.5 459.6 80.25 Skeleton of head 2,989 1702.2 202.8 82.8 511.4 83.65 Skeleton of tail 94 53.3 10.3 3.2 10.0 1.82 Skeleton of shin 1,584 617.4 312.2 51.0 308.7 56.52 Skeleton of shank 1,712 615.7 362.9 61.9 319.8 59 34 Skeleton of flank and plate 1,514 875.8 163.2 48.4 136.0 23.41 Skeleton of rump 570 250.2 90.8 18.5 104.2 18.94 Skeleton of chuck and neck 3,630 1718.4 479.2 112.9 623.5 116.45 Skeleton of round 1,898 767.2 496.8 49.7 311.4 57.26 Skeleton of loin 1,652 726.8 312.9 51.7 281.3 53.36 Skeleton of rib 1,614 753.3 237.1 55.2 265.7 48.50 Horns, hoofs and dewclaws 435 241.8 5.0 30.1 11.3 1.15 Teeth 264 111.2 1.7 5.5 111.3 20.95 Table 54 . — Steer 550 . Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 7,080 5760.4 203.0 43.1 2.19 Circulatory system 1,163 651.3 347.9 23.1 7.7 1.55 Respiratory system 1,460 1137.0 58.2 36.2 14.6 2.91 Brain and spinal cord 443 333.1 44.7 7.0 6.4 1.55 Digestive and excretory system (partial) .... 9,156 6653.2 1192.0 179.0 107.7 16.94 Offal fat 2,610 544.9 1949.3 15.3 8.4 2.06 Liver 1,992 1398.4 62.5 59.6 26.4 6.22 Spleen 291 233.4 7.7 8.6 4.3 0.95 Pancreas 200 133.8 31.1 4.9 2.7 0.64 Kidneys 379 285.6 23.7 9.5 4.5 0.96 Hair and nide 10,440 6779.2 127.1 555.4 136.5 8.77 Head, tail, shin, and shank, lean and fat 5,274 3752.2 491.6 149.2 49.9 9.18 Flank and plate, lean and fat 7,896 4868.1 1532.3 225.6 70.1 12.95 Rump, lean and fat 1,662 995.3 344.8 44.7 15.3 2.96 Chuck and neck, lean and fat 15,814 10633.2 2229.6 424.6 150.2 23.47 Round, lean 11,720 8849.2 310.9 400.6 124.6 24.96 Round, fat 944 306.2 546.8 13.6 3.9 0.65 Loin, lean 9,072 6709.2 408.5 281.0 96.1 20.96 Loin, fat 2,134 465.6 1524.7 19.6 7.0 1.39 Rib, lean and fat 4,408 3045.1 486.4 128.9 43.9 8.24 Kidney, fat 756 78.9 656.8 3.4 1.7 0.43 Skeleton of feet, head, tail, shin and shank. . . 9,839 4520.0 1539.4 290.2 1836.8 340.53 Skeleton of flank and plate 1,540 810.0 197.0 49.9 187.3 33.37 Skeleton of rump 700 285.6 133.7 23.7 133.1 25.45 Skeleton of chuck and neck 4,956 2224.1 810.9 153.5 895.2 163.25 Skeleton of round 2,100 776.9 602.5 55.2 355.1 67.35 Skeleton of loin 2,464 948.2 610.0 71.4 438.0 82.30 Skeleton of rib 1,740 730.6 306.7 57.4 322.1 59.23 Homs, hoofs and dewclaws 549 292.1 7.3 40.1 13.5 1.21 Teeth 228 74.1 1.6 5.0 116.7 21.77 Studies In Animal Nutrition — III 69 Table 55 . — Steer 552. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus 5,219 4163.4 157.1 44.6 1.46 1,056 1.096 466 676.6 219.5 22.9 8.3 1 37 853.8 28.7 29.9 13.2 2.25 345.2 53.5 7.3 6 4 1 56 Digestive and excretory system (partial) .... 5.936 1.784 4452.5 516 0 566.2 1208.8 128.2 16.5 62.8 6.8 10.33 1 23 265 166.0 62.3 6.2 3 8 0 95 1.181 821.0 24.4 36.9 15.7 3.98 284 221.1 3.7 8.8 3 8 0 79 101 74 0 9.4 2.5 1 3 0 30 316 230.3 38.2 7 3 3.6 0 70 10,532 7037.8 161.6 508.5 148.0 7.37 Head and tail, lean and fat 1,209 2.976 822 2 175.1 29 9 11.3 1.87 5.12 Shin and shank, lean and fat 2216.2 138.0 95.6 29.1 Flank and plate, lean and fat 6,204 3928.9 1088.0 177.8 54.7 9.00 Rump, lean and fat 1.326 869.6 196.3 38.3 12.9 2.32 Chuck and neck, lean and fat 12.684 9248.5 899.0 378.9 126.1 22.58 Round, lean 9.830 7479.7 192.4 320.0 111.0 20.35 Round, fat 908 404.8 378.3 19.2 5.4 0 61 Loin, lean 7,034 1.042 5244.5 25.2 220.2 77.4 13.65 Loin, fat 233.8 739.7 12.3 4.1 0 64 Rib. lean and fat 4,104 2971.5 264.1 132.7 39.8 7.39 Kidnev fat 500 64.6 422.8 2 7 1.4 0 17 Skeleton of feet 2,861 3.052 1243.2 366.9 99.5 559.8 99.56 Skeleton of head 1652.4 190.3 89.0 574.1 105.39 Skeleton of tail 131 70 6 17.5 3 8 16.8 3.04 Skeleton of shin 1,550 617.2 273.8 55.1 291.3 51.65 Skeleton of shank 1.742 566.8 363.1 53.8 434.1 79.84 Skeleton of flank and plate 1,516 640 822.8 192.8 51.1 161.5 28.97 Skeleton of rump 254.7 110.4 21.1 132.8 23.49 Skeleton of chuck and neck 3,908 1,626 2,192 1749.0 588.2 126.0 718.3 130.68 Skeleton of round 541.6 482.5 41.7 322.1 56.65 Skeleton of loin 899.9 458.4 63.0 402.8 75 78 Skeleton of rib 2,084 550 912.8 337.4 68.4 365 0 68.90 Horns, hoofs and dev claws 317 3 10.1 35.3 14.8 1 44 Teeth 278 113.8 2.2 5.7 120.2 22.83 Table 56. — Steer 554. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 4.197 3402 6 121.3 33.6 1.30 Circulatory system 591 428.9 61.5 5.0 5.4 1.00 Respiratory system 907 716.7 22.1 24.4 11.0 2.06 Brain and spinal cord 395 303.5 34.7 7.0 6.2 1.38 Digestive and excretory system (partial) .... Offal fat 4.216 644 3276.5 265.6 324.9 327.6 89.9 7.7 46.3 4.1 7.97 0.64 Heart and neck sweetbreads 336 271.1 7.2 8.9 7.1 1.59 Liver 1,166 824 8 30.2 32.8 19.7 4.00 Spleen 202 158.2 2.9 6.0 3.0 0.63 Pancreas 76 53.5 10.1 1.8 1.1 0.21 Kidneys 530 432.0 12.7 11.8 6.1 1.14 Hair and hide 7,400 883 4885.3 119.5 368.2 106.7 7.10 Head and tail, lean and fat 642.6 73.7 24.8 11.1 1.69 Shin and shank, lean and fat 2,304 4,232 1.052 1715.4 90.9 76.5 25.9 4.06 Flank and plate, lean and fat 2993.7 376.5 126.8 41.5 7.19 Rump, lean and fat 741.1 95.6 32.2 12.3 1.87 Chuck and neck, lean and fat 10,530 7,918 520 7882.0 465.0 326.6 131.1 19.90 Round, lean 6013 7 171.3 260.2 100.6 17.02 Round, fat 267 5 169.9 13.9 4.0 0.43 Loin, lean 5,812 428 4350.1 215.0 185.9 66.9 12.03 Loin, fat 127.3 247.7 6.0 2.3 0.34 Rib, lean and fat 2,914 240 2179.5 100.8 95.0 33.2 5.77 Kidney, fat 44.6 181.0 1.7 0.8 0.14 Skeleton of feet 2,403 2,229 1117.3 335.9 89.0 425.0 78.91 Skeleton of head 1330.9 70.4 69.3 375.6 70.26 Skeleton of tail 125 71 .0 14.2 4.3 13.8 2.58 Skeleton of shin 1.464 585.7 269.6 47.2 307.2 58.30 Skeleton of ehank 1.918 772.3 371.3 62.3 334.8 59.92 Skeleton of flank and plate 1.278 775.5 127.2 42.0 99.1 17.56 Skeleton of rump Skeleton of chuck and neck 732 3,732 1,702 357.2 1920.5 92.7 374.5 25.4 117.7 119.2 611.6 21.59 109.91 Skeleton of round 684.2 420 4 45.4 271.5 46.07 Skeleton of loin 1.868 963.9 258.7 55.9 280.2 50.25 Skeleton of rib 1.354 690.3 166.7 43.0 207.7 36.02 Horns, hoofs and dewclaws 338 186.8 4.9 25.9 8.1 0.77 Teeth 225 104.2 0.2 6.6 87.1 14.31 70 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 57. — Steer 555. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus 4,529 3626.6 141.5 33.4 1 59 Circulatory system 554 393.2 66.8 14.3 4.9 0.88 Respiratory system 937 744.4 26.0 25.2 10.5 1.95 Brain and spinal cord 353 269.6 35.1 6.0 5.4 1.25 Digestive and excretory system (partial) .... 4,534 3553.6 288.2 105.1 49.5 8.57 Offal fat 462 271.2 144.9 8.0 3.6 0.55 Heart and neck sweetbreads 159 120.8 10.1 4.3 2.1 0.44 Liver 1.240 925.5 31.5 36.4 18.5 4.33 Spleen 167 129.2 2.3 5.2 2.2 0.46 Pancreas 106 80.8 5.8 2.8 1.6 0.31 Kidneys 439 360.0 12.0 9.8 5.2 0.98 Hair and hide 6,580 4543.6 49.7 342.2 97.1 5 86 Head and tail, lean and fat 956 694.8 88.3 25.1 10.1 1.55 Shin and shank, lean and fat 2,688 2071.0 62.7 85.7 28.1 4.87 Flank and plate, lean and fat 4,068 3071.9 139.3 129.3 42.2 7.12 Rump, lean and fat 876 630.5 63.7 25.5 9.4 1.64 Chuck and neck, lean and fat 9,402 7281.9 201.5 294.3 97.9 17.39 Round, lean 7,126 5554.7 96.0 218.0 85.1 14.75 Round, fat 376 221.8 89.5 9.8 2.8 0.34 Loin, lean 4,618 3599.2 79.7 143.4 55.4 9.60 Loin, fat 274 121.0 117.7 6.0 2.3 0.27 Rib, lean and fat 2,512 1949.7 63.7 77.9 29.5 4.75 Kidney, fat 130 42.9 74.6 1.6 0.7 0.12 Skeleton of feet 2,157 1110.0 195.0 87.2 350.5 59.99 Skeleton of head 1,978 1235.0 63.6 56.6 293.0 50.83 Skeleton of tail 74 46.9 4.9 2.7 6.8 1.13 Skeleton of shin 1,454 731.9 160.9 47.0 259.2 46.38 Skeleton of shank 1,704 846.9 235.3 49.1 253.3 45.26 Skeleton of flank and plate 1,236 805.5 70.3 41.6 99.3 16.82 Skeleton of rump 506 288.9 29.9 17.7 81.7 13.99 Skeleton of chuck and neck 3,216 1907.4 165.7 101.1 447.4 79.21 Skeleton of round 1,652 884.9 219.0 46.1 244.5 42.65 Skeleton of loin 1,264 749.6 90.6 39.2 164.4 29.24 Skeleton of rib 1,256 734.7 79.7 41.1 165.7 28.54 Horns, hoofs and dewciaws 298 145.6 2.7 22.6 7.9 0.40 Teeth 190 81.0 0.6 5.5 81.4 13.86 Table 58. — Steer 556. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 6,124 4989.4 170.1 48.0 2.14 Circulatory system 993 619.2 237.9 21.3 8.5 1.47 Respiratory system 1,272 988.1 43.5 34.3 17.8 2.75 Brain and spinal cord 398 267.1 77.4 6.5 6.4 1.39 Digestive and excretory system (partial) 6,003 4473.6 631.0 132.3 78.3 14.47 Offal fat 1,402 457.3 859.6 13.5 6.3 0.88 Heart and neck sweetbreads 328 250.6 21.6 8.5 67.3 1.48 Liver 1,760 1251.6 61.6 53.9 40.4 6.44 Spleen 300 233.2 7.0 9.0 4.4 0.82 Pancreas 96 69.4 10.2 2.3 1.4 0.29 Kidneys 338 254.6 24.0 8.5 4.5 0.86 Hair and hide 10,314 6836.5 222.1 531.9 142.0 9.08 Head and tail . lean and fat 914 628.3 120.6 25.1 8.2 1.51 Shin and shank, lean and fat 2,808 2070.4 126.6 91.9 27.1 5.19 Flank and plate, lean and fat 6,174 4153.1 824.8 194.1 57.7 10.37 Rump, lean and fat 1,244 862.5 129.2 37.5 12.9 2.38 Chuck and neck, lean and fat 12,406 8971.5 944.7 374.2 154.6 22.33 Round, lean 9,472 7121.0 302.6 311.5 118.5 19.89 Round, fat 686 305.7 298.7 15.1 4.0 0.47 Loin, lean 6,938 5147.0 262.2 226.9 82.5 14.29 Loin, fat 820 214.1 541.5 11.5 4.4 0.52 Rib, lean and fat 3,300 2357.5 238.4 105.1 38.1 6.07 Kidney, fat 420 64.5 338.6 2.5 1.5 0.16 Skeleton of feet 2,589 1202.4 367.0 96.5 459.8 84.89 Skeleton of head 2,610 1478.0 187.6 78.0 460.1 85.37 Skeleton of tail 92 52.4 9.2 3.3 12.3 2.20 Skeleton of shin 1,702 685.5 337.0 55.8 336.6 62.89 Skeleton of shank 1,952 731.1 441.4 66.2 389.8 73.92 Skeleton of flank and plate 1,578 909.9 18.4 52.3 162.0 29.38 Skeleton of rump 608 286.9 80.1 21.5 110.4 20.72 Skeleton of chuck and neck 3,734 1758.9 468.1 130.2 692.5 132.37 Skeleton of round 1,974 753.0 507.8 52.6 329.7 63.40 Skeleton of loin 2,268 1046.8 361.9 71.6 384.0 75.05 Skeleton of rib 1,646 763.2 203.3 54.5 298.9 61.23 Horns, hoofs and dewciaws 390 187.9 4.8 32.4 7.3 0.60 Teeth 218 91.0 0.1 6.4 95.8 15.06 Studies In Animal Nutrition — III 71 Table 59. — Steer 557. Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phospnorus 8,952 2.342 7156.3 280.7 75.4 2.42 1183.1 903.0 38.6 14.4 2.46 1.972 1519.5 103.7 51.2 22.1 4.00 399.9 68.5 69.9 7.8 1 86 Digestive and excretory system (partial) 9,981 6,757 623 7371.1 942.3 1142.4 5640.1 200.3 29.4 104.9 15.5 17.57 1 .96 422.3 105.2 14.5 10.3 2 22 3,003 485 2086.0 56.5 92.6 40.4 9.97 373.1 16.2 14.0 6.7 1.36 225 145.9 38.6 5.1 2.4 0.56 649 498.1 38.2 15.4 7.5 1.41 14,100 1,915 4,196 8900.5 706.7 703.3 165.1 10.01 1158.1 462.2 45.0 16 2 2.76 Shin and shank, lean and fat 2861.9 496.3 123.1 38.9 6.42 Flank and plate, lean and fat 15,294 7713.4 5251.2 357.0 119.0 20.34 Rump, lean and fat 2,582 22,146 14,960 2,532 11,714 4,472 6,372 1,616 3,288 3,558 3,969 193 1321.4 848.1 59.8 20.6 3.67 Chuck and neck, lean and fat 14152.0 3891.9 598.4 211.9 40 53 Round, lean 11044.1 689.7 459.7 151.3 29.02 Round, fat 702.5 1611.5 25.8 8.8 1 22 Loin, lean 8335.1 916.7 347.4 128.6 22.49 Loin, fat 711.4 3573.1 32.6 11.1 1 70 Rib, lean 4322.2 763.8 185.1 61.9 10.83 0.95 Rib, fat 316.3 1194.5 15.5 5.8 Kidney, fat 266.9 2897.7 11.9 5.1 0.77 Skeleton of feet 1579.2 426.8 128.1 670.9 124.10 Skeleton of head 2136.0 248.6 109.2 724.9 130 . 74 Skeleton of tail 102.3 32.4 6.1 20.2 3.77 Skeleton of shin 2,246 2,010 2.498 960 826.4 404.6 75.3 477.0 88.96 Skeleton of shank 962.7 510.0 88.9 523.4 94.61 Skeleton of dank and plate 1408.7 316.2 78.7 242.5 42.99 Skeleton of rump 410.4 128.7 33.0 183.9 35.62 Skeleton of chuck and neck 5,638 2421.1 680.7 191.4 1175.4 212.21 Skeleton of round 2,686 2,478 2,572 695 873.1 789.6 69.1 514.8 95.30 Skeleton of loin 957.2 499.1 73.3 480.3 90.57 Skeleton of rib 1037.8 432.9 83.3 505.9 92.87 Horns, hoofs and dewclaws 373.0 9.8 48.3 17.9 19.39 Teeth 261 104.4 0.7 7.5 120.4 22.53 Table 60. — Steer 558 A' Weights of Constituents in Samples, Grams. Description of sample Sample Water Crude fat Nitrogen Ash Phosphorus Blood 4,666 3903.8 117.1 32.3 1 82 Circulatory system 971 643.2 184.8 21.6 7.1 1.39 Respiratory system 1,111 884.7 16.4 29.8 11.7 2.57 Brain and spinal cord 520 393.1 52.0 8.3 7.8 1.94 Digestive and excretory system (partial) .... 6,510 5075.3 414.4 145.9 64.6 12.24 Offal fat 829 394.6 358.2 11.5 4.2 0.83 Liver 1,352 966.3 28.4 41.9 18.2 4.76 Spleen 193 146.3 2.1 6.4 3.0 0.54 Pancreas 153 116.7 7.9 3.9 2.2 0.57 Kidneys 318 244.2 13.8 8.3 3.8 0.79 Hair and hide 8,138 5266.3 87.7 430.3 92.0 6.75 Head, tail, shin, and shank, lean and fat. . . . 4,324 3216.2 238.2 127.5 40.8 7.05 Flank and plate, lean and fat 4,192 2961.7 306.5 135.4 38.7 7.21 Rump, lean and fat 1,030 719.4 99.3 31.8 10.4 2.02 Chuck and neck, lean and fat 11,682 8789.8 481.4 355.3 108.3 21.85 Round, lean 9,664 7456.0 107.6 301.1 101.4 20.58 Round, fat 644 289.8 266.2 14.9 3.9 0.61 Loin, lean 5,962 4468.5 158.1 189.7 61.1 12.52 Loin, fat 600 181.2 356.5 9.3 2.7 0.48 Rib, lean and fat 3,242 2421.1 113.1 102.4 32.1 6.39 Kidney fat 220 40.5 165.4 2.1 0.6 0.12 Skeleton of feet, head, tail .shin and shank 9,785 4475.4 1645.3 303 0 1716.4 318.21 Skeleton of flank and plate 1,092 596.6 137.0 36.1 113.7 20.58 Skeleton of rump 678 288.5 124.6 21.1 126.6 22.94 Skeleton of chuck and neck 4,056 1752.7 733.8 125.3 723.9 131.41 Skeleton of round 2,094 717.6 708.0 47.9 338.4 62.44 8keleton of loin 2,138 850.3 535.3 54.1 381.4 70.15 Skeleton of rib 1,516 710.8 243.3 50.8 230.3 41.64 Homs, hoofs and dewclaws 443 235.7 5.9 32.4 10.9 0.98 Teetn 274 89.1 1.9 6.0 140.3 26.17 72 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 61. — Composition of Certain Parts and of the Total Animal. (3 Months-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorus Animal 556, Group 1 Lean and fat flesh 70.59 9.14 3.09 1.13 0.184 Bone 46.59 14.37 3.29 17.52 3.332 81.47 2.78 0.78 0 035 Circulatory system 62.36 23.96 2.15 0.86 0.148 Respiratory system 77.68 3.42 2.70 1.40 0.216 Nervous system 67.12 19.45 1.62 1.62 0.349 Digestive and excretory system 74.03 8.56 2.43 1.54 0.276 Hair and Hide 66.28 2.15 5.16 1.38 0.088 Offal fat 32.62 61.31 0.96 0.45 0.063 Total animal 65.23 9.71 3.24 4.88 0.868 Animal 554, Group II Lean and fat flesh 73.19 5.94 3.12 1.17 0.191 Bone 49.29 13.30 3.20 16.20 2.932 Blood 81.07 2.89 0.80 0.031 Circulatory system 72.58 10.40 2.53 0.91 0.169 Respiratory system 79.02 2.44 2.69 1.21 0.227 Nervous system 76.83 8.78 1.77 1.58 0.350 Digestive and excretory system 76.86 5.94 2.32 1.28 0.238 Hair and hide 66.02 1.62 4.98 1.44 0.096 Offal fat 41.23 50.87 1.20 0.63 0.099 Total animal 67.05 7.35 3.23 4.97 0.866 Animal 555, Group III Lean and fat flesh 76.42 3.26 3.08 1.10 0.189 Bone 56.63 7.97 3.21 14.34 2.510 Blood 80.08 3.12 0.74 0.035 Circulatory system 70.98 12.06 2.57 0.88 0.159 Respiratory system 79.45 2.78 2.69 1.12 0.210 Nervous system 76.37 9.95 1.71 1.54 0.354 Digestive and excretory system 77.80 5.27 2.46 1.19 0.227 Hair and hide 69.05 0.76 5.20 1.48 0.089 Offal fat 58.69 31.37 1.72 0.78 0.119 Total animal 71.11 4.38 3.25 4.36 0.739 Table 62. — Composition of Certain Parts and of the Total Animal. (5%-Months-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorous Animal 557, Group 1. Lean and fat flesh 58.12 24.82 2.48 0.86 0.155 Bone Blood 43.24 79.94 15.20 3.18 3.14 18.77 0.84 3.440 0.027 Circulatory system 50.52 38.56 1.65 0.62 0.105 Respiratory system 77.05 5.26 2.60 1.12 0.203 Nervous system 72.58 12.44 1.79 1.41 0.338 Digestive and excretory system 72.81 9.34 2.29 1.15 0.221 Hair and hide 63.12 5.01 4.99 1.17 0.071 Offal fat 13.95 83.47 0.44 0.23 0.029 Total Animal 56.77 20.99 2.75 4.04 0.731 Animal 552, Group II. Lean and fat flesh 77.03 9.45 2.99 0.99 0.175 Bone Blood 43.80 79.77 15.87 3.16 3.01 18.68 0.85 3.399 0.028 Circulatory system 64.07 20.78 2.17 0.79 0.130 Respitatory system 77.90 2.61 2.72 1.21 0.205 Nervous system 74.07 11.48 1.56 1.37 0.335 Digestive and excretory system 73.80 8.71 2.35 1.13 0.210 Hair and hide 66.82 1.53 4.83 1.41 0.070 Offal fat 28.92 67.76 0.92 0.38 0.069 Total animal 63.97 10.48 3.13 5.00 0.880 Animal 548, Group III. Lean and fat flesh 73.75 4.73 3.20 1.05 0.192 Bone Blood 46.67 81.20 15.25 3.13 3.10 16.87 0.72 3. 086 0.021 Circulatory system 70.18 12.58 2.61 0.84 0.157 Respiratory system 76.88 3.13 2.91 1.11 0.202 Nervous system 74.63 11.06 1.64 1.43 0.350 Digestive and excretory system 77.58 3.84 2.56 1.15 0.216 Hair and hide 65.41 0.93 5.35 1.35 0.078 Offal fat 52.57 36.78 1.46 0.72 0.122 Total animal 66.86 6.88 3.32 4.95 0.882 Studies In Animal Nutrition — III 73 Table 63. — Composition of Certain Parts and of the Total Animal. (8%-Months-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorous Animal 547, Group 1. Lean and fat flesh 63.12 17.92 2.72 0.89 0.156 Bone 43.50 16.29 3.18 18.86 3.486 80.85 2.94 0.72 0.029 Circulatory system 58.44 27.81 1.97 0.68 0.124 Respiratory system 78.91 2.82 2.55 1.15 0.195 Nervous system 77.16 8.96 1.54 1.46 0.367 Digestive and excretory system 73.78 9.14 2.32 1.12 0.230 Hair and hide 64.55 1.83 5.33 1.30 0.072 Offal fat 18.06 78.88 0.61 0.26 0.053 Total Animal 61.01 15.73 2.95 3.93 0.704 Animal 550, Group II. Lean and fat flesn 66.53 14.30 2.83 0.94 0.185 Bone 44.11 17.91 3.00 17.86 3.306 Blood 81.36 2.87 0 61 0 031 Circulatory system 56.00 29.91 1.99 0.66 0.133 Respiratory system 77.88 3.98 2.48 1.00 0.199 Nervous system 75.20 10.10 1.58 1.44 0.351 Digestive and excretory system 72.35 10.96 2.18 1.21 0.214 Hair and hide 64.94 1.22 5.32 1.31 0.084 Offal fat 20.88 74.69 0.59 0.32 0.079 Total animal 62.40 13.94 2.97 4.39 0.798 Animal 558, Group III. Lean and fat flesh 73.49 5.52 3.05 0.96 0.190 Bone 43.97 19.32 2.99 17.00 3.125 Blood 88.67 2.51 0.69 0 039 Circulatory system 66.25 19.04 2.22 0.73 0.143 Respiratory system 79.63 1.47 2.69 1.05 0.231 Nervous system 75.60 10.00 1.60 1.51 0.374 Digestive and excretory system 76.81 5.47 2.42 1.08 0.222 Hair and hide 64.71 1.08 5.29 1.13 0.083 Offal fat 47.60 43.21 1.39 0.51 0.106 Total animal 65.95 8.59 3.14 5.01 0.914 Table 64. — Composition of Certain Parts and of the Total Animal. (ll-Months-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorus Animal 541, Group 1. Lean and fat flesh 58.71 23.09 2.68 0.84 0.156 Bone Blood 37.42 82.11 18.81 3.13 2.77 22.39 0 68 3.646 0 027 Circulatory svstem 48.46 39.88 1.65 0.57 0.105 Respiratory system 79.22 1.98 2.74 1.04 0.213 Nervous system 72.46 13.61 1.66 1.53 0.371 Digestive and excretory system 70.99 12.33 2.24 0.92 0.186 Hair and hide 60.43 3.65 5.69 1.10 0.055 Offal fat 11.71 86.40 0.29 0.13 0.026 Total animal 56.23 21.08 2.91 3.86 0.630 Animal 538, Group II. Lean and fat flesh 67.66 14.01 2.90 0.94 0.176 Bone Blood 38.47 82.40 18.49 3.08 2.74 21.61 0.79 3.622 0.028 Circulatory system 55.48 28.70 1.93 0.63 0.114 Respiratory svstem 80.80 2.06 2.50 1.02 0.199 Nervous system 74.33 11.75 1.61 1.50 0.348 Digestive and excretory system 75.01 8.65 2.25 1.00 0.190 Hair and hide 64.34 1.34 5.56 1.06 0.063 Offal fat 21.18 76.45 0.54 0 27 0.055 Total animal 61.65 13.73 3.08 4.79 0.799' Animal 540, Group III. Lean and fat flesh 67.88 11.72 2.97 0 97 0.179' Bone Blood 40.69 82 28 17.48 3.06 2.81 19.90 0 72 3.692 0.026 0.127 Circulatory system 58.13 27.73 2.04 0 6.8 Respiratory system 79.61 2.31 2.59 1.03 0.198 Nervous system 73.66 11.94 1.62 1.58 0.375 Digestive and excretory Bystem 74.44 9.41 2.15 0.87 0.169 Hair and hide 64.64 2.35 5.13 1.26 0.067 Offal fat 25.24 71.06 0.55 0.28 0.047 Total animal 62.93 12.13 3 07 4.76 0.846 74 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 65. — Composition of Certain Parts and of the Total Animal. (ll-Montlis-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorus Animal 505, Group 1. Lean and fat flesh 53.69 29.68 2.57 0.73 0.148 Bone 35.79 17.56 3.19 23.85 4.403 Blood 82.26 2 73 0 33 0.023 0.119 Circulatory system 52.90 35.03 1.88 0.59 Respiratory system 76.83 5.48 2.71 0.95 0.202 Nervous system 73.81 14.74 1.69 1.72 0.411 Digestive and excretory system 73.04 11.03 2.29 1.03 0.223 Hair and hide 62.14 5.34 5.30 0.70 0.066 Offal fat 12.41 85.38 0.34 0.12 0.022 Total animal 53.23 25.06 2.79 4.05 0.749 Animal 503, Group II. Lean and fat flesh 63.17 18.39 2.87 0.84 0.163 Bone 38.28 15.06 3.10 23.70 4.378 Blood 82.78 2.71 0.34 0.076 Circulatory system 51.90 35.79 1.75 0.56 0.115 Respiratory system 78.71 3.16 2.65 0.98 0.207 Nervous system 73.11 16.11 1.68 1.55 0.394 Digestive and excretory system 73.23 8.86 2.45 1.08 0.230 Hair and hide 67.77 2.57 4.79 0.98 0.066 Offal fat 14.64 81.92 0.52 0.18 0.035 Total animal 59.56 16.36 2.98 4.97 0.913 Table 66. — Composition of Certain Parts and of the Total Animal. (18-Months-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorus Animal 532, Group 1. Lean and fat flesh 53.95 28.72 2.57 0.79 0.142 Bone 33.86 22.22 3.34 22.09 4.016 Blood 80.47 3.00 0.65 0.033 Circulatory system 47.92 40.26 1.61 0.57 0.105 Respiratory system 77.55 3.41 2.64 1.05 0.193 Nervous system 72.46 14.46 1.66 1.62 0.392 Digestive and excretory system 71.92 12.02 2.11 0.87 0.177 hair and hide 59.56 6.84 5.23 1.03 0.071 Offal fat 9.55 88.81 0.24 0.12 0.016 Total animal 51.70 26.74 2.75 3.81 0.680 Animal 531, Group III. Lean and fat flesh 68.05 10.03 3.17 1.03 0.189 Bone 38.05 16.48 3.19 23.84 4.268 Blood 81.88 2.84 0.70 0.040 Circulatory system 56.72 29.32 1.97 0.68 0.126 Respiratory system 78.34 2.82 2.68 1.05 0.204 Nervous system 72.99 13.50 1.77 1.50 0.363 Digestive and excretory system 75.88 8.25 2.16 0.85 0.169 Hair and hide 63.67 0.81 5.73 1.14 0.075 Offal fat 25.93 70.68 0.62 0.29 0.046 Total animal 62.64 10.75 3.26 5.37 0.950 Table 68. — Composition of Certain Parts and of the Total Animal. (3-Year-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorus Animal 515, Group 1, 34 mos. Lean and fat flesh 42.28 45.22 1.89 0.57 0.103 Bone 32.02 18.72 3.14 25.71 4.153 Blood 79.11 3.22 0.59 0.024 Circulatory system 42.03 48.53 1.28 0.48 0.086 Respiratory system 76.35 5.44 2.70 0.97 0.187 Nervous system 70.04 16.09 1.65 1.63 0.395 Digestive and excretory system 66.17 18.71 2.02 0.85 0.162 Hair and hide 56.00 12.55 4.84 1.86 0.059 Offal fat 7.53 90.29 0.28 0.12 0.015 Total animal 43.68 37.51 2.29 3.87 0.614 Animal 507, Group II, 34 mos. Lean and fat flesh 60.40 21.48 2.72 0.81 0.151 Bone 32.83 18.05 3.40 25.90 4.004 Blood 78.17 3.41 0.67 0.022 Circulatory system 47.12 41.27 1.70 0.54 0.104 Respiratory system 77.47 3.93 2.73 0.95 0.159 Nervous system 70.81 13.74 1.50 1.75 0.418 Digestive and excretory system 70.52 14.08 2.10 0.83 0.150 Hair and hide 60.85 6.21 5.69 1.07 0.049 Offal fat 13.10 83.96 0.41 0.14 0.029 Total animal 56.10 19.61 3.03 5.07 0.792 Studies In Animal Nutrition — III 75 Table 67. — Composition of Certain Parts and of the Total Animal. (2-Year-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorous Animal 504, Group 1, 21 mos. Lean and fat flesh 49.66 35.13 2.29 0.68 0.128 Bone 32.99 17.28 3.31 27.14 4.993 78.65 3.33 0.39 0 022 Circulatory system 51.05 36.05 1.88 0.69 0.116 Respiratory system 67.41 14.90 2.88 0.97 0.181 Nervous system 69.23 15.08 1.79 1.36 0.350 Digestive and excretory system 72.08 12.76 2.04 0.87 0.178 Hair and hide 58.29 8.07 5.52 1.06 0.043 Offal fat 12.76 84.82 0.33 0.15 0.023 Total animal 49.76 29.42 2.64 4.02 0.724 Animal 523, Group II, 26 mos. Lean and fat flesh 65.17 16.50 2.83 0.86 0.101 Bone 36.08 15.39 3.35 25.78 4.725 Blood 80.52 2.79 0 66 0.022 0.110 Circulatory system 53.05 35.82 1.68 0.60 Respiratory system 78.68 4.05 2.61 0.99 0.189 Nervous system 68.60 17.61 1.62 1.52 0.359 Digestive and excretory system 75.19 10.33 2.19 0.79 0.158 Hair and hide 62.80 1.15 5.60 1.03 0.051 Offal fat 15.42 81.28 0.50 0.20 0.030 Total animal 60.37 15.00 3.10 5.29 0.897 Animal 525, Group III, 26 mos. Lean and fat flesh 68.45 11.91 2.97 0.94 0.174 Bone 35.22 18.20 3.34 23.86 3.952 Blood 80.51 2.94 0.66 0 026 Circulatory svstem 64.09 24.54 1.71 0.72 0.121 Respiratory system 78.61 2.63 2.82 1.11 0.193 Nervous system 71.05 13.93 1.74 1.50 0.370 Digestive and excretory system 77.48 9.14 2.00 0.88 0.156 Hair and hide 62.40 0.50 5.72 1.33 0.057 Offal fat 21.03 70.64 1.29 0.29 0.051 Total animal 62.44 11.87 3.23 5.09 0.838 Table 69. — Composition of Certain Parts and of the Total Animal. (40-Months-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorous Animal 527, Group 1. Lean and fat flesh 37.34 51.80 1.63 0.49 0.089 Bone 30 39 23.73 3.08 25.27 4.302 Blood 78.94 3.28 0.76 0 023 Circulatory system 53.95 32.80 1.93 0.64 0.114 Respiratory system 73.33 7.19 2.72 1.02 0.166 Nervous system 69.96 14.06 1.58 1.54 0.368 Digestive and excretory system 66.12 16.84 2.27 0.89 0.145 Hair and hide 54.48 11.86 5.32 1.37 0.050 Offal fat 5.39 93.20 0.18 0.12 0.013 Thoracic fat 10.30 87.65 0.34 0.13 0.015 Total animal 38.63 45.45 2.02 3.03 0.507 Animal 528, Group II. Lean and fat flesh 59.20 22.38 2.70 0.83 0.152 Bone 34.14 19.67 3.17 24.10 4.425 Blood 79.93 3.09 0.78 0.025 Circulatory system 63.32 21.43 2.34 0.81 0.145 Respiratory system 76.62 3.29 2.80 1.03 0.156 Nervous system 73.22 10.26 1.75 1.73 0.420 Digestive and excretory system 72.16 10 98 2.30 0.92 0.161 Hair and hide 57.83 5.71 5.92 1.44 0.050 Offal fat 12.99 84.14 0 41 0.18 0.018 Thoracic fat 21.47 73.14 0 75 0 29 0.034 Total animal 55.33 20.24 3.04 4.92 0.877 Animal 524, Group III. Lean and fat flesh 70.33 8.80 3.15 0 98 0.175 Bone 37.33 18.10 3.10 23 37 4.175 Blood 82.01 2 79 0 75 0 022 Circulatory system 62 09 22.42 2.24 0.77 0 132 Respiratory system 77 64 2 60 2.79 1 07 0.178 Nervous system 73.22 10.26 1.60 1.42 0.338 Digestive and excretory system 73.86 8.57 2.41 1.00 0.180 Hair and hide 59.26 1.81 6 30 1 58 0.058 Offal fat 25.07 69.44 0.70 0 32 0.035 Thoracic fat Not enoug h to separa te. Total animal 62 43 1 10.50 3.35 5.79 1 008 76 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 70 . — Composition of Certain Parts and of the Total Animal. (45-Months-Old. Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorus Animal 513, Group !. Lean and fat flesh 38.66 49.37 1.70 0.51 0.093 Bone 31.16 20.30 3.30 25.14 4.854 Blood 77.68 3.43 0.72 0.028 0.157 Circulatory system 77.60 4.50 2.67 0.91 Respiratory system 73.91 7.03 2.74 0.99 0.155 Nervous system 68.19 15.50 1.72 1.60 0.410 Digestive and excretory system 72.23 9.03 2.44 1.10 0.194 Hair and hide 57.57 10.07 5.18 0.94 0.054 Offal fat 5.68 92.87 0.22 0.08 0.011 Thoracic fat 13.44 83.71 0.37 0.19 0.024 Total animal 39.91 42.85 2.10 3.20 0.599 Animal 502, Group II. Lean and fat flesh 61.63 18.22 2.91 0.84 0.156 Bone 33.00 19.18 3.35 24.89 4.408 Blood . . 77.43 3 49 0 72 0.023 0.138 Circulatory system 68.29 14.51 2.59 0.68 Respiratory system 76.17 3.06 2.83 1.04 0.166 Nervous system 69.10 14.60 1.72 1.79 0.414 Digestive and excretory system 75.18 6.15 2.59 1.29 0.174 Hair and hide 57.30 3.10 6.57 0.98 0.059 Offal fat 10.65 86.18 0.47 0.19 0.023 Thoracic fat 21.50 73.48 0.75 0.30 0.041 Total animal 56.60 16.97 3.28 5.09 0.887 Animal 509, Group III. Lean and fat flesh 63.17 16.96 2.89 0.85 0.160 Bone 33.59 18.43 3.42 24.78 4.466 Blood 78.05 3.43 0.69 0 023 Circulatory system 69.95 12.26 2.59 0.92 0.167 Respiratory system 77.03 2.88 3.00 1.01 0.176 Nervous system 66.87 17.66 1.68 1.58 0.379 Digestive and excretory system 71.81 10.58 2.37 1.01 0.168 Hair and hide 58.97 2.48 6.27 1.02 0.046 Offal fat 11.36 85.89 0.53 0.18 0.024 Thoracic fat 19.81 76.10 0.72 0.30 0.043 Total animal 57.72 16.27 3.23 5.01 0.887 Table 71 . — Composition of Certain Parts and of the Total Animal. (4- Year-Old Cattle) Percent Water Percent Fat Percent Nitrogen Percent Ash Percent Phos- phorus Animal 501, Group 1. Lean and fat flesh 36.25 53.12 1.46 0.49 0.087 Bone 32.09 17.72 3.36 26.34 4.808 Blood 77.98 3.29 0.86 0.025 Circulatory system 60.00 24.55 2.24 0.74 0.125 Respiratory system 76.58 3.35 2.87 1.04 0.172 Nervous system 70.40 13.27 1.67 1.84 0.392 Digestive and excretory system 70.34 12.06 2.35 0.95 0.169 Hair and hide 51.43 13.24 5.49 1.52 0.049 Offal fat 7.49 91.06 0.21 0.10 0.012 Thoracic fat 18.83 76.65 0.61 0.24 0.026 Total animal 38.75 44.34 2.00 3.33 0.587 Animal 512, Group II. Lean and fat flesh 54.96 28.29 2.48 0.77 0.133 Bone 31.56 20.31 3.23 25.62 4.698 Blood 79.95 3.07 0.79 0.023 Circulatory system 54.88 30.62 2.14 0.84 0.117 Respiratory system 75.49 4.67 2.80 1.08 0.164 Nervous system 72.06 11.12 1.69 1.87 0.385 Digestive and excretory system 72.04 9.89 2.36 0.94 0.164 Hair and hide 56.19 3.61 6.55 1.16 0.047 Offal fat 11.21 86.27 0.37 0.14 0.019 Thoracic fat 12.23 85.52 0.26 0.14 0.015 Total animal 51.88 24.09 2.93 5.10 0.906 Animal 500, Group III. Lean and fat flesh 63.11 17.23 2.96 0.89 0.156 Bone 33.05 22.09 3.19 23.55 4.208 Blood 79.04 3.19 0.79 0.022 Circulatory system 61.58 22.27 2.26 0.80 0.127 Respiratory system 76.50 2.70 2.84 1.15 0.172 Nervous system 62.77 21.72 1.67 1.75 0.371 Digestive and excretory system 73.06 9.29 2.41 1.00 0.164 Hair and hide 59.33 1.32 6.28 1.07 0.044 Offal fat 13.50 83.56 0.42 0.18 0.020 Thoracic fat 17.04 71.39 0.49 0.24 0 024 Total animal 57.25 17.21 3.20 5.08 0.884 Studies In Animal Nutrition — III 77 Table 72 . — Composition of the Total Beef Animal on Analytical, Empty, and Fat-Free Bases. Group Age Basis % Water % Fat % Nitro- gen % Ash % Phos- phorus Embryo 185 days Analytical 84.801 2.363 1.673 1.776 0.283 Embryo Embryo 185 days 232 days 86.853 1.713 1.819 0.290 Analytical 78.700 2.573 2.011 3.180 0.370 Embryo Embryo 232 days 279 days Fat-free 80.778 2.064 3.264 0.380 Analytical 74.192 3.384 2.735 4.062 0.688 Embryo Calves 279 days at birth Fat-free 76.791 2.831 4.204 0.712 Analytical 72.807 3.648 2.926 4.523 0.841 Calves at birth Live weight 73.583 3.544 2.843 4.394 0.817 76.287 2.947 4.555 0.847 I 3 months Analytical 65.226 9.712 3.243 4.875 0.868 I 3 months Empty 66.028 9.488 3.168 4.763 0.848 I 72.949 3.500 5.262 0.937 II 3 months Analytical 67.051 7.348 3.255 4.971 0.866 II 3 months Empty 67.562 7.234 3.175 4.894 0.853 II 72.830 3.422 5.276 0.919 0.739 III 3 months Analytical 71.108 4.377 3.247 4.356 III 3 months Empty 71.517 4.315 3.201 4.295 0.729 III 74.742 3.345 4.488 0.761 I 5H months Analytical 56.771 20.988 2.753 4.039 0.731 I 5 Yi months Empty 57.213 20.773 2.725 3.998 0.723 I 5 Yi months 5 l A months Fat-free empty 72.214 3.439 3.130 5.046 0.913 II Analytical 63.966 10.479 4.996 0.880 II 5M months Empty 64.388 10.356 3.093 4.937 0.870 II 5A months 5A months Fat-free empty 71.827 3.451 5.508 0.970 III Analytical 66.863 6.884 3.315 4.947 0.882 m h x A months Empty 67.800 6.689 3.221 4.807 0.857 hi h l A months 8H months Fat-free empty 72.661 3.452 5.152 0.919 i Analytical 61.009 15.728 2.952 3.931 0.704 i 8 l A months Empty 61.233 15.637 2.935 3.908 0.700 i 8 A months 8 X A months Fat-free empty 72.584 3.479 4.633 0.830 ii Analytical 62.402 13.936 2.974 4.389 0.798 ii 8A months Empty 63.055 13.695 2.922 4.312 0.784 ii 8 M months 8A months Fat -free empty 73.060 3.386 4.997 0.908 m Analytical 65.947 8.590 3.135 5.010 0.914 hi 8 X A months Empty 66.509 8.437 3.079 4.921 0.897 hi 8 X A months 11 months Fat-free empty 72.637 3.363 5.374 0.980 i Analytical 56.225 21.078 2.905 3.862 0.630 i 1 1 months Empty 57.556 20.437 2.817 3.744 0.610 i 11 months Fat-free empty 72.340 3.540 4.706 0.767 i 11 months Analytical 53.228 25.061 2.785 4.049 0.749 i 11 months Empty 54.424 24.421 2.714 3.946 0.730 i 11 months Fat-free empty 72.009 3.591 5.220 0.966 ii 11 months Analytical 61.651 13.731 3.076 4.785 0.799 11 11 months Empty 63.006 13.245 2.967 4.616 0.771 ii 11 months Fat-free empty 72.626 3.420 5.320 0.889 ii 11 months Analytical 59.557 16.361 2.983 4.969 0.913 ii 11 months Empty 60.371 16.031 2.923 4.869 0.895 ii 1 1 months Fat-free empty 71 . 897 3.481 5.799 1.066 in 11 months Analytical 62.925 12.125 3.068 4.764 0.846 hi 11 months Empty 64.800 11.512 2.913 4.532 0.803 in 11 months Fat-free empty 73.231 3.291 5.111 0.908 78 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 73. — Composition of the Total Beef Animal on Analytical, Empty, and Fat-Free Bases. Group Age Basis % Water % Fat % Nitro- gen % Ash % Phos- phorus I 18 months Analytical 51.695 26.740 2.748 3.808 0.680 I 18 months Empty 53.038 25.997 2.672 3.702 0.661 I 18 71.671 3.611 5.003 0.894 III 1,8J4 months Analytical 62.641 10.747 3.258 5.367 0.950 III l&A months Empty 65.411 9.951 3.017 4.969 0.879 III 72.639 3.350 5.518 0.976 I 21 months Analytical 49.762 29.420 2.639 4.015 0.724 I 21 months Empty 50.933 28.735 2.577 3.921 0.708 I 21 71.469 3.616 5.502 0.993 II 26 months Analytical 60.368 15.001 3.095 5.290 0.897 II 26 months Empty 61.960 14.398 2.971 5.077 0.861 II 26 72.382 3.470 5.931 1.006 III 26 months Analytical 62.444 11.871 3.231 5.088 0.838 III 26 months Empty 64.115 11.344 3.087 4.861 0.801 III 26 72.318 3.482 5.483 0.903 I 34 months Analytical 43.684 37.509 2.286 3.873 0.614 I 34 months Empty 46.601 35.566 2.167 3.672 0.582 l 34 72.324 3.364 5.699 0.904 II 34 months Analytical 56. 102 19.613 3.031 5.071 0.792 II 34 months Empty 58.014 18.759 2.899 4.850 0.758 II 34 months Fat-free empty 71.408 3.568 5.970 0.933 I 39H months Analytical 38.628 45.446 2.019 3.026 0.507 I 3934 months Empty 39.827 44.558 1.980 2.967 0.497 1 3Q14 months Fat-free empty 71.836 3.571 5.352 0.897 II 40 months Analytical 55.333 20.242 3.039 4.924 0.877 II 40 months Empty 56.556 19.688 2.956 4.789 0.853 II 40 months Fat-free empty 70.421 3.680 5.963 1.062 III 4034 months Analytical 62.430 10.499 3.353 5.794 1.008 III 4034 months Empty 63.491 10.202 3.258 5.630 0.979 III 4034 months Fat-free empty 70. 705 3.628 6.270 1.091 I 4434 months Analytical 39.912 42.850 2.103 3.201 0.599 I 4434 months Empty 41.396 41.792 2.051 3.122 0.585 I 4434 months Fat-free empty 71.117 3.524 5.363 1.004 11 4434 months Analytical 56.599 16.972 3.281 5.086 0.887 II 4434 months Empty 57.625 16.571 3.203 4.966 0.866 II 44V4 months Fat-free empty 69.071 3.840 5.952 1.038 III 45 months Analytical 57.715 16.266 3.234 5.005 0.887 III 45 months Empty 58.369 16.015 3.184 4.928 0.873 III 45 months Fat-free empty 69.498 3.791 5.867 1.040 I 47 months Analytical 38.752 44.340 1.999 3.329 0.587 1 47 months Empty 39.836 43.556 1.963 3.270 0.577 I 47 months Fat-free empty 70.576 3.478 5.793 1.022 11 48 months Analytical 51.879 24.091 2.927 5.098 0.906 II 48 months Empty 52.666 23.697 2.879 5.014 0.891 II 48 months Fat-free empty 69.022 3.773 6.572 1.167 III 48 months Analytical 57.254 17.209 3.200 5.078 0.884 III 48 months Empty 58.142 16.852 3.134 4.972 0.866 III 48 months Fat-free empty 69.926 3.769 5.980 1.041 Studies In Animal Nutrition — III 79 Table 74. — Composition of Beef Flesh on Fat-Free Basis. Group Age months Animal Sample % Water % Nitro- gen % Ash % Phos- phorus IU 48 500 Round, lean 76.704 3.236 1.048 0.198 III 48 500 Loin, lean 76.159 3.374 1.095 0.201 III 48 500 Rib, lean 76.573 3.645 1.060 0.194 III 48 500 Lean and fat composite 76.709 3.511 1.117 0.196 I 47 501 Round, lean 77.117 3.409 1.056 0.204 I 47 501 Loin, lean 76.228 3.489 1.037 0.199 1 47 501 Rib, lean 75.900 3.556 1.019 0.192 I 47 501 Lean and fat composite 76.537 3.331 1.182 0.183 II 44 H 502 Round, lean 75.199 3.446 1.012 0.203 II 44 H 502 Loin, lean 76.057 3.475 1.061 0.217 II 44 H 502 Rib, lean 73.826 3.413 0.994 0.182 II 44 502 Lean and fat composite 76.098 3.616 1.029 0.195 II 11 503 Round and rump lean 76.776 3.540 1.076 0.214 II 11 503 Loin, lean 77.548 3.481 1.074 0.208 II 11 503 Rib, lean 77.618 3.513 1.033 0.206 I 21 504 Round, lean 76.561 3.533 1.083 0.214 I 21 504 Loin, lean 76.236 3.476 1.078 0.206 I 21 504 Rib, lean 76.722 3.565 1.008 0.202 I 11 505 Round and rump, lean 76.286 3.674 1.078 0.221 I 11 505 Loin, lean 75.752 3.595 1.070 0.218 I 11 505 Rib, lean 76.430 3.668 1.041 0.218 II 34 507 Round, lean 77. 182 3.428 1.041 0.204 II 34 507 Loin, lean 76.924 3.040 1.058 0.201 II 34 507 Rib, lean 76.843 3.417 1.068 0.198 III 45 509 Round, lean 76.862 3.364 1.049 0.205 m 45 509 Loin, lean 76.527 3.410 1.045 0.198 hi 45 509 Rib, lean 76.041 3.283 1.026 0.091 hi 45 509 Lean and fat composite 76.359 3.718 1.039 0.194 ii 48 512 Round, lean 76.770 3.392 1.073 0.201 ii 48 512 Loin, lean 75.997 3.458 1.025 0.191 ii 48 512 Rib, lean 76.566 3.489 1.053 0.185 ii 48 512 Lean and fat composite 76.136 3.496 1.074 0.188 i 44H 513 Round, lean 76.064 3.387 1.063 0.202 i 44 H 513 Loin, lean 75.757 3.542 1.054 0.208 i 44 M 513 Rib, lean 76.900 3.551 1.047 0.196 i 44 H 513 Lean and fat composite 77.369 3.345 1.088 0.194 i 34 515 Round, lean 76.102 3.487 1.039 0.199 i 34 515 Loin, lean 76.599 3.536 1.172 0.213 i 34 515 Rib, lean 76.851 3.487 1.066 0.191 ii 26 523 Round, lean 78.748 3.230 1 067 0.207 ii 26 523 Loin, lean 77.856 3.379 1.028 0.192 ii 26 523 Rib, lean 77.486 3.439 1.022 0.196 80 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 75. — Composition of Beef Flesh on Fat-Free Basis. Group Age months Animal Sample % Water % Nitro- gen % Ash % Phos- phorus III 40 M 524 Round, lean 79.010 3.348 1.072 0.197 III 40H 524 Loin, lean 76.140 3.468 1.107 0.203 III 40 H 524 Rib lean and fat 76.354 3.596 1.067 0.198 III 40 H 524 Lean and fat of animal 75.735 3.509 1.102 0.201 III 26 525 Round, lean 78.931 3.197 1.090 0.210 III 26 525 Loin, lean 77.316 3.353 1.086 0.207 III 26 525 Rib, lean 77.199 3.461 1.070 0.198 II 40 526 Round, lean 75.907 3.745 1.170 0.217 II 40 526 Loin, lean 76.008 3.391 1.075 0.201 II 40 526 Rib, lean 77.391 3.419 1.022 0.183 II 40 526 Lean and fat of animal 76.605 3.595 1.093 0.186 I 39H 527 Round, lean 76.903 3.477 1.004 0.203 I 39^ 527 Loin, lean 76.967 3.518 1.068 0.203 I 39)4 527 Rib, lean 76.471 3.542 1.095 0.196 I 39H 527 Lean and fat of animal 76.535 3.390 1.073 0.191 I* 38 529 Rib, lean 76.958 3.474 1.023 0.192 III 18H 531 Round, lean 76.503 3.333 1.123 0.212 III 18 H 531 Loin, lean 76.274 3.460 1.149 0.213 III 18 H 531 Rib lean and fat 75.222 3.611 1.138 0.206 III 18H 531 Lean and fat of carcass 74.007 3.686 1.218 0.227 I 18 532 Round, lean 75.867 3.486 1.124 0.211 I 18 532 Loin, lean 75.664 3.597 1.111 0.204 I 18 532 Rib, lean 76.008 3.565 1.106 0.199 I 18 532 Lean and fat of carcass 74.809 3.747 1.157 0.209 II 11 538 Round, lean 78.078 3.247 1.123 0.208 II 11 538 Loin, lean 77.494 3.303 1.118 0.215 II 11 538 Rib, lean 77.170 2.348 1.107 0.208 II 11 538 Lean and fat of carcass, excl. kidney fat 77.379 3.379 1.136 0.205 III 11 540 Round, lean 77.444 3.282 1.107 0.215 III 11 540 Loin, lean 76.759 3.318 1.122 0.210 III 11 540 Rib lean and fat 76.867 3.428 1.122 0.200 III 11 540 Lean and fat of carcass, excl. kidney fat 76.910 3.304 1.102 0.191 I 11 541 Round, lean 76.490 3.456 1.127 0.215 I 11 541 Loin, lean 76.187 3.423 1.112 0.210 I 11 541 Rib, lean 76.654 3.490 1.090 0.209 I 11 541 Lean and fat of carcass, excl. of kidney fat 76.264 3.686 1.129 0.212 I 8 H 547 Round, lean 77.207 3.343 1.119 0.218 I VA 547 Loin, lean 76. 177 3.390 1.097 0.212 I 8A 547 Rib, lean 77.071 3.360 1.081 0.199 I 8 H 547 Lean and fat composite 76.768 3.348 1.094 0.197 III 5A 548 Round, lean 76.804 3.420 1.137 0.218 III 5A 548 Loin, lean 76.878 3.360 1.151 0.213 III 6H 548 Rib lean and fat 77.983 3.221 1.124 0.200 III 5H 548 Lean and fat composite 77.576 3.238 1.095 0.198 III 55 549 Rib, lean 77.060 3.542 1.164 0.206 ♦Maintenance one year. Then full feed. Studies In Animal Nutrition — III 81 Table 76. — Composition of Beef Flesh on Fat-Free Basis. Group Age months Animal Sample % Water % Nitro- gen % Ash % Phos- phorus n 8 H 550 Round, lean 77.563 3.511 1.092 0.219 II 8 M 550 Loin, lean 77.442 3.243 1.109 0.242 II 8 M 550 Rib lean and fat 77.651 3.287 1.120 0.210 II m 550 Lean and fat composite 77.704 3.194 1.056 0.207 n 5y 2 552 Round, lean 77.609 3.320 1.152 0.211 II 5 'A 552 Loin, lean 77.334 3.248 1.142 0.201 II 5 H 552 Rib, lean and fat 77.385 3.455 1.036 0.192 II 5^ 552 Lean and fat composite 77.421 3.237 1.135 0.187 n 3 554 Round, lean 77.629 3.359 1.299 0.220 ii 3 554 Loin, lean 77.723 3.322 1.194 0.215 ii 3 554 Rib, lean and fat 77.473 3.378 1.179 0.205 ii 3 554 Lean and fat composite 77.986 3.231 1.257 0.201 hi 3 555 Round, lean 79.013 3.101 1.210 0.210 HI 3 555 Loin, lean 79.308 3.160 1.221 0.212 hi 3 555 Rib, lean and fat 79.634 3.182 1.205 0.194 m 3 555 Lean and fat composite 79.372 3.150 1.087 0.193 i 3 556 Round, lean 77.660 3.398 1.292 0.217 i 3 556 Loin, lean 77.099 3.398 1.236 0.214 i 3 556 Rib, lean and fat 77.002 3.432 1.243 0.198 i 3 556 Lean and fat composite 77.809 3.333 1.283 0.196 i 5^ 557 Round, lean 77.392 3.222 1.060 0.203 i 5 H 557 Loin, lean 77.196 3.218 1.191 0.208 i 5 H 557 Rib, lean 77.069 3.301 1.088 0.193 i 5 H 557 Lean and fat composite 77.549 3.345 1.292 0.200 m 8H 558 Round, lean 78.020 3.151 1.061 0.215 m 8H 558 Loin, lean 76.992 3.269 1.053 0.216 hi 8H 558 Rib, lean and fat 77.379 3.271 1.026 0.204 hi sy 558 Lean and fat composite 77.830 3.203 1.006 0.204 Jersey 6 days 22A Flesh 78.477 3.743 1.062 0.204 High Plane Newborn 562B Flesh 79.108 3.118 1.110 0.175 High Plane Newborn 562C Flesh 80.539 2.724 1.031 0.180 Medium Plane Newborn 565A Flesh 80.318 2.760 0.951 0.176 Medium Plane Newborn 563A Flesh 79.401 2.921 1.047 0.163 Medium Plane Newborn 564B Flesh 79.925 2.926 0.989 0.178 Medium Plane Newborn 565B Flesh 79.874 2.909 1.091 0.187 Medium Plane Newborn 564C Flesh 79.302 2.897 1.071 0.191 Low Plane Newborn 568B Flesh 82.931 2.272 0.896 0.146 Low Plane Newborn 567B Flesh 80.332 2.746 1.119 0.180 Low Plane Newborn 566B Flesh 82.039 2.806 1.049 0.150 Low Plane Newborn 568C Flesh 79.293 2.841 1.057 0.186 High and 1 Medium Plane / Newborn Average Flesh 79.794 2.902 1.051 0.179 Low Plane Newborn Average Flesh 81.149 2.666 1.030 0.166 82 Missouri Agr. Exp. Sta. Research Bulletin 55 Table 77. — Empty Weight at Start. Animal Age atfirst feeding (days) Live Weight at first feeding (pounds) Live weight (kilograms) Empty weight (per cent) Empty weight (kilograms) 500 32 118.5 53.750 0.950 51.1 501 10 98.0 44.452 0.967 43.0 502 15 110.6 50.167 0.953 47.8 503 23 158.6 71.939 0.920 66.2 504 21 147.7 66.995 0.927 62.1 505 21 127.1 57.651 0.943 54.4 507 19 93.2 42.275 0.970 41.0 509 15 107.8 48.897 0.957 46.8 512 28 168.5 76.430 0.912 69.7 513 14 106.7 48.398 0.957 46.3 515 19 114.0 51.709 0.950 49.1 523 23 132.2 59.965 0.940 56.4 524 19 145.4 65.952 0.930 61.3 525 36 154.6 70.125 0.922 64.7 526 41 150.6 68.311 0.926 63.3 527 27 167.8 76.112 0.913 69.5 531 73 230.2 104.416 0.866 90.4 532 54 187.5 85.048 0.897 76.3 538 25 132.2 59.965 0.940 56.4 540 32 140.3 63.639 0.933 59.4 541 20 137.6 62.414 0.936 58.4 547 12 95.5 43.318 0.969 42.0 548 20 147.5 66.905 0.928 62.1 550 21 148.5 67.358 0.927 62.4 552 18 140.0 63.503 0.934 59.3 554 18 130.0 58.967 0.941 55.5 555 21 147.0 66.678 0.928 61.9 556 19 148.0 67.131 0.927 62.2 557 23 132.6 60.146 0.939 56.5 558 13 112.6 51.074 0.951 48.6 549 18 117.0 53.070 0.951 50.5 551 21 149.5 67.812 0.927 62.9 559 19 117.4 53.251 0.950 50.6 Table 78. — Amount and Composition of Gain from Start to Slaughter. (Page 83) Weight Phos- phorus 832.33 521.86 310.47 665.91 462.87 203.04 517.87 518.72 -0.85 1,250.16 471.78 778.38 864.21 496.34 367.87 737.10 521.02 216.08 1.199.78 346.08 853.70 949.39 523.54 425.85 807.60 402.41 405.19 1.759.79 488.22 1,271.57 2,002.23 453.15 1,549.08 1,225.46 470.94 754.52 2,116.15 557.40 1,558.75 1,105.98 497.18 608.80 3,035.37 648.55 2,386.82 Percent Phos- phorus IlllsISllslllSlslilllglallsllsIlgSSgllsISsIsI 00000000000000000000000000000000000000000000 Weight Ash 4.673.7 2.954.5 1.719.2 3,820.9 2.614.1 1.206.8 3.052.6 2.940.3 112.3 6.908.3 2.661.2 4.247.1 4.905.1 2,810.8 2.094.3 4.133.4 2.949.8 1.183.6 6.700.4 1.911.0 4.789.4 5.222.8 2.970.2 2.252.6 4.428.8 2.259.9 2.168.9 10,794.1 2.762.3 8.031.8 10,824.9 2.556.8 8.268.1 7.334.7 2.656.4 4,678.3 11.512.8 3.157.7 8.355.1 6.229.2 2.815.6 3.413.6 16.994.9 3,670.0 13,324.49 Percent Ash IgglSSlgllSlSSlIglillsIIlllsgsillssIlglsSlgal Weight Nitrogen 3.108.8 1.978.0 1.130.8 2.478.4 1.742.7 735.7 2.274.9 1.968.4 306.5 4.708.5 1.779.8 2.928.7 3.073.2 1.879.8 1,193.4 2.770.1 1.974.8 795.3 5.031.8 1.251.6 3.780.2 3.539.2 1.984.3 1.554.9 2.771.2 1.496.9 1.274.3 8,120.8 1.845.4 6.275.4 7.445.3 1.702.7 5.742.6 4.714.7 1.776.6 2.938.1 6.910.6 2.118.4 4.792.2 4.011.3 1.883.0 2.128.3 12,264.6 2.441.6 9.823.0 Percent Nitrogen SgSS§llSlgSllg|SSIIIillllllS§ISSllS|lllgSgglg COCOCOCOCOCOrOCOC*3DCOC^(>IC^CSlCOC^COCOeOC^COCaCO-OC^-l»-l00 Group I m II II I I III II I III II I III II I (Page 84) TABLE 78. — AMOUNT AND COMPOSITION OF GAIN FROM START TO SLAUGHTER — Continued. Weight Phos- phorus 1,688.27 777.44 910.83 3,366.90 521.02 2,845.88 2.909.44 470.94 2,438.50 2,126.98 544.13 1,582.85 3,913.72 406.55 3,507.17 3.174.44 337.02 2,837.42 3,909.04 584.05 3,324.99 3,651.97 531.72 3.120.25 3,156.16 513.69 2,642.47 4,507.90 382.44 4,125.46 3,847.63 395.31 3,452.32 3,418.30 387.04 3.031.26 4,701.00 354.32 4,346.68 4,399.23 588.97 3.810.26 3,529.88 422.47 3,107.41 Percent Phos- phorus SlIsllllllIglllKlsIIllIlgllllllsIgllslIIlllls OOOOOOOOOOOOOOOOOOOOOOOOOO'-'OOOOOOOOOOOOOOOOOO 13 9,540.5 4.384.4 5.156.1 18,658.4 2,949.8 15.708.6 17.150.6 2.656.4 14.494.2 12.911.0 3.079.7 9.831.3 24.673.2 2.288.1 22.385.1 20.316.2 1.857.3 18,458.9 23.320.7 3.307.8 20.012.9 20,498.4 3.013.1 17.485.3 18.141.7 2.911.8 15.229.9 24.073.7 2,134.4 21.939.3 22.068.9 2.217.9 19.851.0 19.290.4 2.162.2 17.128.2 26.645.1 1.965.1 24.680.0 24.764.9 3,338.6 21.426.3 20.279.1 2.382.1 17,897.0 Percent Ash llliglsSSIIlBllllllggg|glBl2slIllIlllglsglslI Weight Nitrogen 5.792.2 2.892.8 2.899.4 12.264.1 1.974.8 10.289.3 10.034.9 1.776.6 8.258.3 8.199.5 2.063.9 6.135.6 14.563.2 1.512.3 13.050.9 12.142.4 1.217.7 10.924.7 15.561.1 2.214.4 13.346.7 12.649.9 2.019.3 10.630.6 10.498.9 1.949.3 8.549.6 15.819.3 1,412.2 14.407.1 14.236.6 1.467.5 12.769.1 12.464.4 1.427.4 11.037.0 15.999.4 1,290.0 14.709.4 14.217.6 2.230.4 11.987.2 12.781.0 1.577.9 11.203.1 Percent Nitrogen sllslISgllSlSlglllllllilSSlilllllSlIlllSlISsS Weight Fat 19,105.6 7,412.8 11.692.8 136.735.0 3.229.2 133.505.8 48,636.5 2.707.2 45,929.3 30.126.9 3,429.1 26.697.8 238,975.5 2,160.4 236.815.1 78.579.9 1.640.0 76.939.9 350,228.8 3.806.0 346,422.8 84.263.0 3.291.6 80.971.4 32,874.7 3.126.3 29.748.4 322,276.3 1.944.6 320,331.7 73,645.3 2.079.3 71.566.0 62,609.6 2.012.4 60.597.2 354.940.0 1,763.0 353.177.0 117,034.5 3.833.5 113.201.0 68.726.2 2.290.5 66,436.1 Percent Fat SlselllllsllllislIIllllllllalSslisllIsgllllls IS 125,592 60,026 65,566 242,366 43,532 198,834 209,304 40,044 169,260 170,280 45,225 125,055 313,119 35.254 277,865 243,018 29,848 213,170 313,045 47,956 265,089 242,058 44,310 197,748 204,591 43,094 161,497 319,219 33,382 285,837 256,101 34,416 221,685 228,490 33,743 194,747 324,632 31,089 293,543 260,103 48,302 211,801 237,124 36,444 200,680 Percent Water 3iii§iiissiiiiiii§siisiisiiiiiiiiiiiiiiiiisii If 192.005 90.400 101,605 475,854 62,100 413,754 337,803 56.400 281,403 265,587 64.700 200,887 671,917 49,100 622,817 418.896 41.000 377.896 786.005 69,200 716,805 427,995 63.300 364,695 322,234 61.300 260,934 771,142 46.300 724,842 444,424 47.800 396,624 391,461 46.800 344,661 814.914 43.000 771.914 493,877 69.700 424,177 407,833 50.900 356,933 Condition At slaughter At start Gain At slaughter At start Gain At slaughter At start Gain At slaughter At start Gain At slaughter At start Gain At slaughter At start Gain At slaughter At start Gam At slaughter At start Gain At slaughter At start Gain At slaughter At start Gain At slaughter At start Gain At slaughter At start Gain At slaughter At start Gain At slaughter At 3tart Gain At slaughter At start Gain Animal 531 504 523 525 515 507 527 526 524 513 502 509 501 512 500 Age months Group III II I III II I III II I II III II III Studies In Animal Nutrition — III 85 Table 79. — Empty Weight at Age Previous Animal Was Slaughtered. Animal Age Live weight at end feeding Empty weight at slaughter Per cent empty weight Weight at age previous animal was slaughtered Yrs. — Mos. — Days lbs. kgs. kgs. Live kgs. Empty kg3. Group 1 556 0 3 0 247.6 112.31 98.13 87.38 34.830 557 0 5 17 443.2 201.03 172.80 85.96 109.41 95.599 547 0 8 5 450.2 204.21 171.45 83.96 142.93 122.859 541 0 10 22 724.0 328.40 288.30 87.79 250.38 210.221 505 0 10 18 690.6 313.25 274.36 87.58 255.37 214.409 532 1 5 20 1133.0 513.92 459.03 89.32 324.00 284.439 504 1 8 26 1170.0 530.70 475.85 89.67 488.74 436.545 515 2 9 19 1632.2 740.35 671.92 90.76 539.05 483.363 527 3 3 15 1869.0 847.76 786.01 92.72 777.45 705.616 513 3 8 15 1898.4 861.10 771.14 89.55 782.17 725.229 501 3 11 0 1964.8 891.21 814.91 91.44 873.71 782.403 Group 2 554 0 3 0 196.0 88.90 78.07 87.81 34.830 552 0 5 7 256.2 116.21 99.35 85.49 88.09 77.349 550 0 8 14 323.8 146.87 121.11 82.46 104.10 88.994 538 0 10 26 403.0 182.80 158.91 86.93 146.78 121.036 503 0 11 11 608.4 275.96 236.43 85.67 209.11 172.428 523 2 2 6 864.2 391.99 337.80 86.18 221.58 192.619 507 2 9 16 1014.4 460.12 418.90 91.04 440.12 379.294 526 3 4 0 1088.2 493.60 428.00 86.71 441.12 401.592 502 3 8 19 1138.6 516.46 444.42 86.05 491.69 426.346 512 3 11 21 1250.4 567.17 493.88 87.08 562.54 484.067 G^oup 3 555 0 3 0 188.4 85.46 71.08 83.17 34.830 548 0 5 9 222.0 100.70 85.99 85.39 79.02 65.717 558 0 8 12 237.8 107.86 90.00 83.44 94.12 86.369 540 0 11 2 341.8 155.04 137.73 88.83 135.90 113.392 531 1 6 12 479.6 217.54 192.01 88.26 153.99 136.793 525 2 2 8 694.6 315.06 265.59 84.30 240.49 212.259 524 3 4 13 806.2 365.68 322.23 88.12 299.60 252.559 509 3 8 22 1004.2 455.50 391.46 85.94 416.26 366.808 500 3 11 26 1061.8 481.62 407.83 84.68 438.30 376.678 (Page 86) TABLE 80. — AMOUNT AND COMPOSITION OF GAIN BETWEEN EACH SUCCEEDING AGE. If .•8§S383ffS53g33!S8!Sl5SS8gS8S3S8SgS£§§S8§S8SS Per cent. Phosphorus Is!SI!laill!lllIs§elalel§llS§IssSslllss OOOOOOOOOOOOOOOOOOOOt-hOOOOOOOO^hOOOOOOOOO b > NrfiWC0Tj<05Tj<05»0rHrl05 00O00W05Tt(c0C0OiH»O«DNW05i-iN^W»0X^*HNNNOC0t0C0 ! si Sill §§ la § IS la ill S 11 11 ISIIsIIIIy III.sII Per cent. Nitrogen IllsssisSsllsIIisSsSIsiiSlsSIlSlISsslIsl Weight Fat .O^fMOi^iONiClNCCCOOOHGOiOOONOCO^tO^HX^^eOiOWOOlrHOi^iO CO C05^oo«005NN(OOOOOCONHO(X)(NONOi Weigh Nitroge Grams 2,478 990 1,488 3,073 2,455 617 3,539 2,752 786 4.714 3,536 1,178 6,910 5,038 1,872 10,034 5.715 4,319 12,142 11,268 873 12,649 11,642 1,007 14.236 12,602 1,633 14,217 15,504 —1,287 12,755 12,602 152 14,217 13,892 324 a II SSIlSsSlSlgSlliSSllSsllllllslSIIlSIl IS T Weigh Fat Grams 5,647 1,234 4,413 10,288 5,595 4,693 16,585 9,216 7,369 21,048 16,575 4,472 37,903 23,614 14,289 48,636 25,512 23,124 78,579 54,610 23,969 84,263 75,334 8,928 73,645 83,939 —10,293 117,034 80,214 36,819 90,218 83,939 6,279 117,034 98.265 18,768 Per cent. Fat Weight Water Grams. 52,746 25,629 27,117 63,969 52,259 11,710 76,367 57,301 19,066 100.124 76,319 23,805 142,735 108,724 34,011 209,304 121,362 87,942 243,018 235.011 8,007 242,058 232,980 9,078 256,101 241.124 14,977 260,103 278,944 —18,841 249.011 241,124 7,887 260,103 271,223 —11,120 er cent. Water T T Empty Weight Grams. 78.071 34,830 43,241 99.349 77.349 22,000 121,112 88.994 32,118 158,911 121,036 37,875 236,429 172,428 64,001 337,803 192.619 145.184 418,896 379,294 39,602 427,995 401,592 26,403 444,424 426,346 18,078 493,877 484,067 9,810 444,424 426,346 18,078 493,877 484,067 9,810 Condition At slaughter At birth Gain At slaughter At 3 mo Gain At slaughter At 5^ mo Gain At slaughter At 8M mo Gain At slaughter At 8H mo Gain At slaughter At 11 mo. (538).. . Gain At slaughter At 26 mo Gain At slaughter At 34 mo Gain At slaughter At 40 mo Gain . At slaughter At 44f% mo Gain At slaughter At 40 mos Gain At slaughter At 44 Yi mo Gain Animal 554 552 550 538 503 523 507 526 502 512 502 512 months £ £ £ ^ -ill”" 'I (Page 88) Table 80. — Amount and Composition of Gain Between Each Succeedinq Age — Continued. Weight Phosphorus Grams. 517.87 284.56 233.31 737.10 479.08 258.02 807.60 688.76 118.84 1.105.98 1,017.13 88.85 1,688.27 1,098.45 589.82 2.126.98 1,865.76 261.22 3.156.16 2,023.00 1.133.16 3,418.30 3,591.05 —172.75 3,529.88 3,288.40 241.48 2.738.99 2,023.00 715.99 3,418.30 3,117.87 300.43 Per cent. Phosphorus SsISesISSlSIllIlllsIlSIglsSlIiSls O0000»-I00>-<00000»-H00000'-I00000000»-I00<-| Weight Ash ,CD^N^i005acCU5NOCq»0H^OH0iN05 00^«05HN^O05H^oO CO *0 OCOC0030a>iOTj<-«tieooo©'!t<50''*ioc&'^eo-H Grams 2,274 990 1,284 2.770 2,103 666 2.771 2,588 182 4,011 3,491 520 5,792 3,984 1,807 8,199 6,403 1.795 10,498 7.796 2,702 12,464 11,950 513 12,781 11,993 787 9,860 7,796 2,063 12,464 11,224 1,240 Per cent. Nitrogen gI3SS5i&ggiSgSgggS8S83S13381583§g Weight Fat pn W^05C0NO'H05W^05iCOC0O05OONC0^tDW00«0OO^C000«0N^ Per cent. Fat ss11s1S!1S§11b1II1II11111sI1SI11= '^CClO®-«f^OO«bg5^00«5©’^®^05«0©^«0«D®gcOCO©lCi-HOO©lC-H Weight Water Isssgs3ss-gs2sss§ssss3^||7s2ssss|a” Per cent. Water BSl!ssi!g|!l31133iS2i!llSllISI!l! Empty Weight Grams. 71,078 34,830 36,248 85,988 65,717 20,271 89,999 80,369 9,630 137,726 113,392 24,334 192,005 136,793 55,212 265,587 212,259 53,328 322,234 252,559 69,675 391,461 366,808 24,653 407,833 376,678 31,155 322,234 252,559 69,675 391,461 366,808 24,653 Condition At slaughter At birth Gain At slaughter At 3 mo Gain At slaughter At 5)4 mo Gain At slaughter At 8)4 mo Gain At slaughter At 11 mo Gain At slaughter At 18)4 mo Gain At slaughter At 26 mo Gain At slaughter At 40)4 mo Gain At slaughter At 45 mo Gain At slaughter At 26 mo Gain At slaughter At 40)4 mo Gain Animal 555 548 558 540 531 525 524 509 500 524 509 months X X X ^ ^ eoiooo^ooookooooug 1 1 J. i 1 i T i T T | UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 56 Observations on Winter Injury I — Early and Late Winter Injury F. C. Bradford II — An Aftermath of Winter Injury H. A. Cardinell (Publication Authorized September 1, 1922.) COLUMBIA, MISSOURI NOVEMBER, 1922 UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE Agricultural Experiment Station BOARD OF CONTROL THE CURATORS OF THE UNIVERSITY OF MISSOURI EXECUTIVE BOARD OF THE UNIVERSITY E. LANSING RAY P. E. BURTON H. J. BLANTON St. Louis Joplin Paris ADVISORY COUNCIL, THE MISSOURI STATE BOARD OF AGRICULTURE OFFICERS OF THE STATION J. C. JONES, PH. D., LL. D., PRESIDENT OF THE UNIVERSITY F. B. MUMFORD, M. S., DIRECTOR STATION STAFF NOVEMBER, 1922 AGRICULTURAL CHEMISTRY RURAL LIFE O. R. Johnson, A. M, S. D. GromEr, A. M. E. L. Morgan, A.M. C. R. Moulton, Ph. D. L. D. Haigh, Ph. D. W. S. Ritchie, Ph. D. E. E. Vanatta, M. S. A. R. Hall, B. S. in Agr. E. G. Sieveking, B. S. in Agr. AGRICULTURAL ENGINEERING J. C. Wooley, B. S. Mack M. Jones, B. S. ANIMAL HUSBANDRY E. A. Trowbridge, B. S. in Agr. L. A. Weaver, B. S. in Agr. A. G. Hogan, Ph. D. F. B. Mumford, M. S. D. W. Chittenden, B. S. in Agr. A. T. Edinger, B. S. in Agr. H. D. Fox, B. S. in Agr. BOTANY W. J. Robbins, Ph. D. DAIRY HUSBANDRY A. C. Ragsdale, B. S. in Agr. Wm. H. E. Reid, A. M. Samuel Brody, M. A. C. W. Turner, B. S. in Agr. D. H. Neison, B. S. in Agr. W. P. Hays ENTOMOLOGY Leonard Haseman, Ph. D. K. C. Sullivan, A. M. O. C. McBride, B. S. in Agr. FIELD CROPS W. C. Etheridge, Ph. D. C. A. Helm, A. M. L. J. Stadler, Ph. D. O W. Letson, B. S. in Agr. Miss Regina Schulte* Ben H. Frame, B. S. in Aerr. Owen Howells, B. S. in Agr. HORTICULTURE T. J. Talbert, A. M. H. D. Hooker, Tr.. Ph. D. J. T. Rosa, Jr., Ph. D. H. G. Swartwout. B. S. in Agr. J. T. Quinn, B. S. in Agr. POULTRY HUSBANDRY H L. Kempster. B. S. Earl W. Henderson, B.S. SOILS M. F. Miller, M. S. A. H. H. Krusekopf, A. M W. A. At brecht. Ph. D. F. L. Duley, A.M. Wm. DeYoung, B. S. in Agr. H. V. Jordan, B. S. in Agr Richard Bradfield, Ph. D. VETERINARY SCIENCE J. W. Connaway, D. V. S., M. D. L. S. Backus, D. V. M. O. S. Crisler, D. V. M. A. J. Durant, A. M. H. G. Newman, A. M. OTHER OFFICERS R. B. Price, M. S., Treasurer Leslie Cowan, B. S., Secretary S. B. Shirkey, A. M., Asst, to Director A. A. Jeffrey, A. B., Agricultural Editor J. F. Barham, Photographer Miss Jane Frodsham, Librarian. E. E. Brown, Business Manager. In service of U. S. Department of Agriculture. Observations on Winter Injury I. — Early and Late Winter Injury F. C. Bradford Destruction of flower buds without attendant damage to other parts of the tree is the most common manifestation of winter injury in the peach and apparently very uncommon in the apple. Conversely, injury to other tissues without attendant damage to flower buds seems relatively more common in the apple than in the peach. In short, blossom buds are ordinarily the susceptible point in the peach and the resistant part of the apple tree. The general recognition of the connection between responsiveness of peach flower buds to high temper- atures in winter and the too frequent consequent damage from ordi- nary winter cold has fostered a rather widespread assumption that injury to flower buds of any kind must be attributed to this combin- ation of weather conditions. For this reason an occurrence of injury confined to blossoms in the apple while peaches in the same orchards were wholly undamaged, constituting a case of injury supposedly due to premature starting from dormacy in a fruit supposedly least sub- ject to this weakness, merits investigation and record. MANIFESTATIONS Late in December, 1921, it was noted that a small percentage of fruit buds on Jonathan trees in the University orchard at Columbia had been injured, the floral parts being clearly discolored. In the spring of 1922 these trees blossomed very heavily. Shortly after the blossoms had fallen, the unusual persistence of the bud scales and the peculiar behavior of the axillary buds attracted attention. In many cases the terminal buds were dead and growth was proceeding from axillary buds. Most of the dead buds abscissed, leaving a smooth, flat surface, as shown in Fig. 1. In other cases the terminal growth was very feeble and fruit had set from axial blossom clusters. An in- stance of this is shown in Fig. 2. Since in Jonathan the formation of axillary flower buds without accompanying flower buds on terminals has not been observed, the association of axillary blossoms with leafy terminals invited attention. Furthermore, the setting of fruit from axillary buds in such numbers as appeared this spring was unusual ; ordinarily the terminal buds set fruit and the axillary buds do not. 4 Missouri Agr. Exp. Sta. Research Bulletin 56 Many spurs which had not blossomed this year bore fewer leaves than is usual in ncn-blossoming spurs. Figs. 3 and 4 show views from two angles of a spur of this type found on a Ben Davis tree. Careful examination of either reveals a sharply marked ring just above the lowest leaves. An enlarged view of a similar spur (Fig. 5) shows the significance of this appearance perhaps more clearly. Here the ring, just below the leaf insertions, is plainly visible. This ring, a single continuous line, is quite different from the composite belt of scale scars marking the transition from one year’s growth to the next. It occurs only on fruiting wood at the point where the vegetative axis leaves the purse. At the right of the spur, just below the ring, is a black protuber- ance; this is the dried remnant of a flower cluster which was killed in the bud. The purse, or cluster base, the swelling which bears the blossoms and fruit, so prominent in the majority of bearing spurs (cf. Fig. 15), is here reduced to extremely small dimensions. The vegetative axis, relatively small in many blossoming or bearing spurs, in these cases constitute nearly all the current season’s growth. In some the purse is reduced to even smaller size and can scarcely be dis- tinguished (Fig. 8). In others it is fairly conspicuous. Fig. 6 is a photomicrograph of a longitudional section of the bud shown in Fig. 5. Though this section does not pass through the exact center, the dead blossoms within the cluster are discernible. The smallness of the pith cylinder extending to the blossoms is rather bet- ter evidence than the possibly shrivelled blossoms could be, of the size of the bud at the time of the killing. Fig. 7, representing a Jan- uary stage in an uninjured bud of another variety, is used here to illus- trate the extent of the injury to the Jonathan bud. That part of the bud represented by the portion above the line k k’ was killed. Growth continued from the vegetative point just below this (v), resulting in the growth shown to the left of and above the injured portion. A section of another bud is shown in Fig. 9. In this case the dead tissue had abscissed and the pith cylinder extending to the point of abscission (a) is the best evidence of the former presence of blossoms at this point. Injury to the pith is shown at k and the growth from the vegetative point was less vigorous. More severe injury is shown in Fig. 10, also some regenerative tissue. Complete killing is shown in Fig. 11 and, in Fig. 12, complete killing followed by abscission. Dissection of these twigs and spurs showed, in many cases, though not invariably, some discoloration of the pith. Generally when the bud had sloughed off, as shown in Fig. 1, little injury was visible in Observations on Winter Injury Plate I. — Fig. 1. Jonathan, showing abscission of winter-killed terminal. Fig. 2. Jonathan, showing setting of fruit from axial blossom clusters. Figs. 3 and 4. Ben Davis spur, showing winter killing of blossoms. Fig. 5. Jonathan spur, showing dead blossoms. Fig. 6 Photomicrograph of section of spur shown in Fig. 5. 6 Missouri Agr. Exp. Sta. Research Bullet 56 Plate IT. — Fig. 7. Uninjured bud. In injured buds, tissues above k-k' killed and growth proceeds from v. Fig. 8. Jonathan spur, showing dead blossoms at k and extreme reduction of purse. Fig. 9 Jonathan spur, showing abscission of dead blossoms at a and injury to pith at k. Fig 10. Similar injury involving greater area. Fig. 11. Whole bud killed. Fig. 12, bud killed and abscissed. Observations on Winter Injury 7 Plate III. — Fig. 13. Kicffer pear, showing replacement of killed spurs from supernumerary buds. Fig. 14. Kieffer pear spur, injured in spring of 1921, bearing in 1922. Injury shown in blackened wood. Fig. 15. Ben Davis spur, July, 1921. Bearing from normal and from second bloom. %ai Fig. 13 8 Missouri Agr. Exp. Sta. Research Bulletin 56 Plate IV. — Series of sections of Kieffer pear. Fig. 16. Pith injury near tip of 1921 wood. Fig. 17. Injury to pith and wood in 1920 wood. Fig. 18. Section from lower level in 1920 wood; more wood and less pith injury. Fig. 19. Still lower on 1920 wood; injury confined to wood. Observations on Winter Injury 9 the remaining tissue. Some of the spurs in which the blossoms had been killed showed no further injury. In the majority of these cases, however, there were rather poorly defined small areas in which the pith was in some cases browm, in some cases orange. Other spurs, however, which were bearing normally showed somewhat the same injury. This has been observed in Oregon and in Missouri in other seasons. Parenthetically it may be recorded that spurs somewhat similar in external appearance at this time to those just described, but with the purse better developed, were showm by dissection to be infected with fire blight. Here the discoloration was black and primarily in the vessels rather than the pith. Its course from the blossom attachments was easily traced. Leaves on infected spurs at this stage are still green and practically the only external evidence of infection is the rather flaccid condition of the purse. OCCURRENCE The distribution of' this winter injury was rather general among the Jonathan trees in two parts of the University orchard. Other varieties examined, as Ben Davis, Ingram and Wealthy, showed it, but very rarely. Others, as Oldenburg, York and Gano, apparently had none. It was very common in Malus prunifolia. In Jonathan it affected certainly 10 per cent of the buds. It was found in young Jonathans in the cultivated orchard at Turner though less abundantly than in the sod orchard in Columbia. Possibly the trees in the better drained positions suffered less, but the differences were not marked. In a row on the northern boundary of the orchard injured buds were distinctly more plentiful on the north side of the trees. This might have been taken as evidence of a cold wind as a factor in the injury, were it not that elsewhere in the orchard injury was distinctly localized on other sides. Almost invariably it was most abundant on that side which faced an open space, regardless of orientation. Much more consistent was the preponderance of injury in buds on terminal shoots over that on spurs. Though spurs greatly outnum- ber terminals, probably three-fourths of the cases of injury appeared on terminal shoots. The importance of this observation will appear later. In the peach no sign of injury could be found, though many axil- lary leaf buds failed to open in the spring. Poorly developed axillary buds on vigorous shoots of many perennials, whether they are those 10 Missouri Agr. Exp. Sta. Research Bulletin 56 formed in the first flush of spring growth or those laid down just be- fore growth stops in late autumn, are inclined to remain dormant. Consequently this failure to grow is no indication of winter injury, particularly in view of the absence of any evidence of injured tissue. In the peach the pith was brown in spots, but the cells were not col- lapsed. In this fruit the pith is short-lived and discoloration often occurs in new growth before midsummer. Fruit buds were, practically without exception, uninjured. Consequently it may be said that the peach came through the winter without injury. Table 1. — Precipitation and Temperature at Columbia, September to Decem- ber, 1921, Compared to That oe the Same Months, 1911 to 1920 Inclusive. Precipitation (inches) Aug Sept. Oct. Nov. Dec. 1921 5.83 Average, 1911-1920 4.90 Maximum 7.83 Minimum 0.77 Daily Mean Temperature (°F.) 1921 Average, 1911-1920 Maximum (monthly) Minimum (monthly) Absolute Minimum Temperature (°F.) 1921 1911-1920 10.04 2.33 1.34 1.41 6.06 4.90 1.86 1.63 9.69 7.68 3.24 2.94 2.38 0.72 0.10 0.44 71.9 57.4 43.8 35.5 68.6 57.1 45.1 32.4 73.0 60.1 48.9 39.8 61.4 48.5 38.0 25.4 44 32 17 7 32 20 6 —9 CAUSE The records of the United States Weather Bureau Station at Co- lumbia, made available by Dr. George Reeder, section director for Missouri, are summarized in Table 1, to December 31, covering the period during which the injury occurred. They show the mean tem- peratures for September and December to have been rather well above the averages for the previous ten years and that for November some- what lower, but in no case did they reach the extremes recorded in the previous ten years. The absolute minima for the several months have in each case been exceeded in other recent years. Unusual cold is evidently not the primary factor in this injury. The precipitation record, however, shows an unusual rainfall in September, higher than any in the previous ten years; indeed it was exceeded but twice in 35 years. This rainfall, coupled with the con- siderable increase in mean temperature for the same month, undoubt- Observations on Winter Injury 11 edly tended to delay maturity. Raspberries made considerable new growth and fall blossoming of cherries was rather widespread. Some Japanese plums in the University orchard were partly in blossom in October. Though the subsequent winter was not at all remarkable for cold weather, in either duration or intensity, it resulted in practically complete destruction of red and purple raspberry canes and serious damage to black raspberries. Injury confined to flower buds during early winter has been re- corded but rarely. Maynard 4 * and Bartlett 1 described cases of Decem- ber killing of peach buds in Massachusetts. In the case cited by Maynard the cold was not intense (10°F.), but it followed warm weather during which the blossoms were observed to develop. Chand- ler 3 found mild injury to peach blossom buds in New York during the winter of 1914-1915. The previous August had been character- ized by heavy precipitation. The coldest temperature of the winter, — 9°F., occurred in December. The injury was in the pith of the bud and of the twig and apparently had no effect beyond retarding blos- soming. In these cases it is not altogether clear whether the buds had started to develop or had failed to mature. The only careful report of winter injury to blossom buds in apple unaccompanied by further injury is that of a case observed by Whip- ple 5 in Montana. This form is apparently identical in its manifesta- tions with that just described for Columbia. Though this form of injury seems rare, it is quite possible, as Whipple points out, that, since the injury escapes casual observation, it may occur from time to time and pass unnoticed. Whipple suggested that the damage in Montana might have been due to thawing in high winds or to freezing after warm weather in January or February. All the evidence in the occurrence at Columbia, however, con- nects immaturity with the injury. It followed weather conditions in- viting and in many cases leading directly to immaturity injuries in other fruits. Peaches, far more susceptible to injury from premature de- velopment, were entirely uninjured. The greater injury on open sides may have been due to prolonged growth as much as to greater exposure. In the varieties affected, damage was greatest in the ter- minals on shoots; these are late in differentiating flower buds, late in maturing and late in starting from dormancy. Some late blossoming varieties were affected while some early blossoming varieties escaped. Finally, the injury had occurred before the last of December. •This and subsequent superscript numerals refer to literature cited in the Bibliography. 12 Missouri Agr. Exp. Sta. Research Bulletin 56 IMPLICATIONS Attention of fruit growers in Missouri has been focused on pre- mature starting of buds during winter rather than on immaturity. In this case peach trees in the same orchard showed no injury, either in wood or in fruit bud. In addition to weather, other conditions were particularly favorable to injury from immaturity in the peach this win- ter, for, following the Easter freeze in 1921 the trees had been cut back to wood 2 or 3 inches in diameter and had made growths of three to 6 or 7 feet, with secondary and even tertiary branches. Consequently their immunity from injury while Jonathan apples were afflicted indi- cates that in the peach at Columbia immaturity is less important than premature starting. On the other hand, so far as concerns the apple, evidence is accumulating in a chain, of which the case here recorded is but one link, that even to Central Missouri, as is the case farther north, immaturity is of no mean importance. Crown rot in Grimes and crotch injury in Stayman are presumably due to injury consequent upon immaturity. Cardinell has found serious cases of winter injury — with heart rot as the consequence — in young apples, evidently tracing to immaturity and cold in the fall of 1917. Injuries due to immaturity are not patent. The type described here might well escape observation and the crop failure be attributed to lack of fruit bud formation. In- juries to other tissues are often unnoted until brought to attenion by the wood-destroying fungi which find entrance through such lesions, or until the bark comes away, long after the weather conditions re- sponsible for the injury have passed from recollection. In many cases such injuries are attributed to fungi and referred to under the general term “canker’. If the conventional notions of the effects of cultivation be accepted, peaches and apples may be bad neighbors in Central Missouri orchards, for the cultural practices which, by inducing late growth, tend to make peaches resist stimulation from warm weather in January and Feb- ruary tend to make some apples more subject to injury in November and December. When peaches are used as fillers in apple orchards, cultural practices designed to protect either fruit against winter in- jury are likely to make the other more susceptible. However, it should be recognized that the comparison made here is between peaches in cultivated soil and apples in sod. The generally accepted views of the effects of these two systems of management are, for the most part, founded on investigations in sections with a shorter and uniform growing season. For many crops there are in Missouri Observations on Winter Injury 13 two rather distinct growing seasons, separated by a season of dry, hot weather. In some cases, as with potatoes and cabbage, this may be due chiefly to the excessively high temperature of midsummer, but with others dry weather has its undoubted influence. The raspberry, companion of the potato and the cabbage in ability to grow where the summers are too cool to ripen grain, shows the same reaction to the growing season of Central Missouri. Immaturity injury in this fruit occurs every year in varying degree at Columbia before intense cold sets in ; it may be produced by temperatures certainly no lower than 12 °F. and possibly higher. In late August the canes are often more nearly mature than they are in October, following the moderate tem- perature and greater rainfall of September. Those which grow through August seem to mature better than those which stop growing at this time. Card 2 found in New York that under some circumstances the first shoots to start in the spring may be more tender in the follow- ing winter than those starting somewhat later. Prolonged tillage through the dry season may have the actual effect of inducing final maturity by so prolonging the first flush of growth that the second growth does not start. In short, for this fruit the growing season here is apparently too long; the canes mature and then resume growth. On the other hand, the peach, adjusted to warmer summers, suf- fers little or no check from heat during its growing season. The long period of warm weather enables this tree to mature properly despite high cultivation. In fact, prolonged cultivation makes it more hardy. The apple, with growing season temperature requirements higher than those of the raspberry and lower than those of the peach, un- doubtedly suffers more or less here from immaturity. The evidence at hand, however, does not warrant any conjecture as to the effects of tillage on maturity in this fruit. The smaller amount of injury in the cultivated trees at Turner than in the sod orchard at Columbia, eight miles away, is interesting, possibly suggestive, but certainly not indi- cative. The greater prevalence of injury on terminals than on spurs at Columbia points in the opposite direction. Consequently at present it cannot be stated definitely whether this immaturity injury is due to prolonged growth or to renewed growth. One thing becomes increasingly evident. Hardiness in Central Missouri is more complicated than it is farther north or farther south. In some regions it is largely a matter of maturity, in others a matter of continuing dormancy. Here it is in some fruits the first, in other fruits the second. This is the first complication. The second compli- cation comes from the fact that immaturity alone, a rather simple mat- 14 Missouri Agr. Exp. Sta. Research Bulletin 56 ter farther north, is here possibly induced by the very practices that obviate it elsewhere. Finally, since this section shares northern and southern winter weather, extreme measures for guarding against one type of injury may be efficacious in one winter and injurious in the next. Solution of the problems raised will depend on recognition of the types of injury to which each fruit is subject, determination of the probability of the occurrence of weather conditions leading to each type and formulation for each fruit of cultural practices which, over a period of years, will reduce the injuries most likely to occur. VARIABILITY OF HARDINESS After the Easter freeze of 1921 the Kieffer pears showed more damage than any other trees in the University orchard. Many spurs were killed, many branches were killed back well into 1920 wood and older wood was discolored. Other pears, such as Garber, Tyson and Surprise, were injured but little. Since the freeze the recuperative ability of Kieffer has been as remarkable as was its susceptibility. The majority of the killed spurs have been replaced by new growths arising from supernumerary buds at the base of the old spurs (Fig. 13), and the spurs whose wood was blackened in 1921 are bearing in 1922 (Fig. 14). Since this variety had proved so tender in the spring of 1921, it was examined for injury occurring in the late months of the same year. Evidence of injury to pith near the tip of 1921 wood was plenti- ful (Fig. 16), but there was no indication of injury to fruit buds and little or no indication of injury to wood. Farther back on these same branches showing pith injury in the 1921 wood, appeared injury of another kind. Just below the point where growth was resumed in 1921 both pith and wood were injured (Figs. 17 and 18). Still farther back, but yet in 1920 wood, the injury was confined to the wood. In gross appearance there was a ring of blackened tissue, which is shown by microscopic examination to be very narrow (Fig. 19). Inside the blackened ring is a narrow belt of new wood, one or two cells wide, composed of wood laid down in the spring of 1921 before the freeze. The injury was confined to parenchymatous tissue and the wood just laid down was hardy enough to withstand the freezing. Discoloration of pith is not invariably a sign of winter injury, but, under the circumstances of its occurrence in the material discussed here, it may be taken as such. The injury to the pith in the 1921 wood was undoubtedly an immaturity injury. The injury to the pith in the 1920 wood may have been due to the weather of the 1920-1921 winter or Observations on Winter Injury 15 to the Easter freeze of 1921. In any case, it is clear that the most tender tissue in the fall of 1921 was the pith while in the spring of the same year it was the wood. Extensive examination of 1920 wood in other pears and several varieties of apple showed no wood injury from the Easter freeze com- parable to that in Kieffer. In some cases Ben Davis 1920 wood showed a dark ring (Fig. 3). On microscopic examination this was found to be, not injured tissue, but a false annual ring caused by the check to growth resulting apparently from the killing of the foliage in the same freeze. The pear trees in the University orchard stand within a few feet of many Jonathan and Ben Davis trees and receive the same cultural treatment. These Jonathans show no wood injury from the Easter freeze, Ben Davis only a check to growth, while Kieffer was severely affected. In the fall of the same year, however, conditions were re- versed; Jonathan was injured rather extensively, Ben Davis much less and Kieffer least of all. This condition, coupled with the reversal of the accepted comparative hardiness of the peach and the apple, illus- trates anew the fact that hardiness is consitutional only in so far as conditions producing hardiness are constitutional. It is a condition rather than a quality. A given fruit is hardy in a locality as it reacts to the ordinary climatic conditions of that locality and comparative hardiness may be reversed in various localities according as the spring or the fall injuries are likely to prevail. THE “SECOND BLOOM” Following a frost causing widespread damage to blossoms and fruit crop there is frequently a flood of reports of crops borne on “second bloom” pushed out as a consequence of the injury to the first crop. The occurrence at times of second bloom in rather considerable quantity is unquestionable. After a destructive frost it is naturally more noticeable than it would be in a frostless season when it would come on about at the end of the first bloom. Its occurrence, however, is not necessarily a consequence of frost injury. It was noted in great abundance in the University orchard at Columbia in the spring of 1921 before any frost had occurred and in the spring of 1922 when there was no damage from frost. Fig. 15, showing a Ben Davis spur taken in the summer of 1920, is significant. Growth for that year started at g. One apple results from the first bloom and one from the second. Clearly in this case the destruction of the first bloom is not concerned with the formation of the second. 16 Missouri Agr. Exp. Sta. Research Bulletin 56 SUMMARY 1. Killing of many fruit buds in the apple occurred early in the winter of 1921-1922 in the University orchard at Columbia. 2. The attendant circumstances indicate that this injury was con- nected with immaturity. 3. Many plants with low optimum growing temperatures have, in Central Missouri, two distinct growing seasons separated by a hot, dry midsummer. The raspberry apparently belongs in this group. 4. Other plants, with higher optima, grow rather uniformily through the season. This group includes the peach. 5. Sod culture may have the effect of accentuating the duality of the growing season for the plants with low optima. Consequently its effect on maturity in these plants may be directly opposite that recognized in regions with short and relatively cool growing seasons. 6. Available evidence is not sufficient to indicate how tillage af- fects maturity in the apple in Central Missouri. 7. In the Kieffer pear the relative hardiness of the various tis- sues appears to vary with the season. 8. The Kieffer pear, under the conditions discussed in this paper, is more tender in the wood in the spring than the Jonathan apple, but more hardy in fruit buds in the fall. 9. The so-called second bloom is not necessarily the consequence of the destruction of the “first” or normal bloom. LITERATURE CITED 1. Bartlett, G., Horticulturist, 1: 549. 1847. 2. Card, F. W., Bush Fruits, p. 37., New York, 1917. 3. Chandler, W. H, Proc. Am. Soc. Hort. Sci., 12: 118. 1915. 4. Maynard, S. T., Agriculture of Massachusetts, p. 348. Bos- ton, 1884. 5. Whipple, O. B., Mont. Agr. Exp. Sta. Bui. 91. 1912. Observations on Winter Injury 17 II. — An Aftermath of Winter Injury H. A. Cardinell In the course of some continuing demonstration work in 1920 and 1921 in a young orchard at Fortescue, Holt County, Missouri, atten- tion was drawn to the failure of the pruning wounds to heal. Where- ever any wood was removed, large or small, callus formation was very slow or failed altogether (Figure 6). Not only did wounds which should have healed in one season fail to cover over, but the wood in the immediate neighborhood seemed dead. Wounds disinfected and some both disinfected and painted healed no better than those un- treated. Cuts made below old wounds revealed much dead wood in the center, surrounded by more or less live wood. Cuts through the trunk still lower on the tree, in many cases down to within three or four inches of the ground, showed the same condition (Fig. 4). For reasons which will appear later in this paper, it was possible to exam- ine 1243 trees this spring. In this group only a few cases were found of discoloration extending below the graft union and fully 50 per cent of the trees were not injured below three or four inches above the ground as shown at k in Fig. 4. Aside from this failure of wounds to heal properly the trees pre- sented no unusual appearance, except in some extreme instances. By the spring of 1922 the trees most affected, though many of these same trees were making 20 to 40 inches of growth each year, as shown in Fig. 5, had one or two dead limbs to the tree. At the base of these limbs the wood on the trunks was practically all dead. In general, however, the trees were making very good growth and on casual obser- vation the orchard would have appeared in excellent condition. It is no uncommon occurrence to find black-hearted limbs or trunks on trees that have been growing and fruiting in a perfectly sat- isfactory manner. This condition is known to be caused by winter injury, sometimes from ordinary cold in conjunction with an immature condition of the tree. It occurs when the cold is severe enough" to kill the wood but still not severe enough to kill the hardier cambium. Consequently in the following spring the cambium may resume growth and surround the dead area with a layer of new tissue. Sometimes these blackened regions are found surrounded by Healthy wood show- ing 20 or more annual rings, indicating that the injury had occurred as many years previously and apparently had not interfered materially 18 Missouri Agr. Exp. Sta. Research Bulletin 56 with the tree’s life or functions. In itself the condition is not serious, particularly in older trees. In the case here considered, however, there were two disturbing circumstances: (1) the failure of the wounds to heal, already mentioned, and (2) the fact that the discoloration was evidently advancing with the growth, spreading into new wood (Fig. 3, at B and C). This made diagnosis of the cause and prognosis of the ultimate effects somewhat uncertain. HISTORY OF THE CASE The trees involved stand in a 60-acre block, on level but well drained Missouri River bottom land. This orchard is one of the properties of George Hitz & Company of Indianapolis and is managed by Mr. C. E. Hitz. In the spring of 1918, yearling trees to the number of 2,997 were planted and in the fall of the same year 567 two-year-olds were set. Jonathans were planted as permanent trees, with fillers of five varieties : Ben Davis, Delicious, Stayman, Grimes and Ingram. All the stock used came from a nursery at St. Joseph, Missouri. While these trees were being cut back subsequent to planting, Mr. Hitz noticed that there was a slight discoloration in the wood. During the summer of the same year a pathologist from the United States Department of Agriculture, visiting the orchard on another errand, examined these trees and diagnosed the trouble as winter injury. DIAGNOSIS The complication already alluded to and the resemblance of some of the wounds to cankers of fire blight, so common on Jonathan in Northwest Missouri, rather obscured the case. Specimens of injured wood were submitted to pathologists in various sections of the country and the possibility of several other disorders eliminated. In Feb- ruary, 1922, a shipment of injured trees was sent to M. B. Waite, Pathologist in Charge, Fruit Disease Investigations, U. S. Department of Agriculture. Under date of February 25, Waite states: “I have given these samples careful study. The main trouble is winter injury. It is com- plicated by secondary trouble due to wood-rot fungi. These wood-rot fungi have produced a heart rot by entering the frozen injured centers of the trunks and main branches, and the wood-rot fungi have ex- tended the injury somewhat, and perhaps complicated and confused the primary injury. * * * There are indications on these samples that they may have been frozen a second time. I have often noticed that Observations on Winter Injury 19 trees once injured by freezing appear slightly more susceptible. Part of the two-year wood and most of the one-year wood is sound.” Of the discoloration in the one-year-old wood Waite says : “This appears to be partly due to the growth of the wood-rot fungus from the diseased part up into the healthy tissue. One of the samples shows the tip of a small trunk which has been killed completely and shows the fungus fruiting.* Mr. W. H. Diehl of this bureau has identified this fungus as Irpex tulipifera Schw ” . Weather records indicate several possibilities of winter injury in the time since these trees stood in the nursery, in the summer of 1917. However, inasmuch as injury to these trees is known to have occurred prior to planting in 1918 and the secondary injury is less certain, interest centers in the weather from October, 1917, to March, 1918. The mean temperature for the state as a whole was below nor- mal during most of 1917 and particularly in October and December. The October mean temperature was the lowest on record. Killing frosts occurred on October 6, ten days earlier than the average. That fall will be remembered by many people in this section for the great amount of soft corn. Table 1. — Precipitation at St. Joseph, Missouri. (In Inches) Aug. Sept. Oct. Nov. Dec. 1917 5.80 1.60 0.80 0.04 0.16 Average, 1888-1917 4.37 3.34 2.62 1.71 0.96 The Weather Bureau records for St. Joseph, where the trees discussed in this paper were grown, indicate that the pre- ceding autumn was rather dryer than the average. August, however, had a rainfall 1.4 inches above the average. Whether this could have had any material effect in prolonging growth and deferring maturity is problematical. Table 2. — Minimum Temperatures at St. Joseph, Missouri. (In degrees F.) Oct. Nov. Dec. Jan. Feb. Mar. 1917-18 20 17 ^13 — 19 —73 17 1909-1917 22 5 —10 —24 —16 -4 •After the manuscript was prepared many ungrrafted trees had died and one type of fruitincr bodv was noticed on all. This was identified Sept. 8, 1922, by Diehl as Polystictus versicolor, Fr. 20 Missouri Agr. Exp. Sta. Research Bulletin 56 Minimum temperatures lower than any since 1909, when the St. Joseph record begins, occurred in October and December. Just when the injury to these trees occurred, cannot be stated definitey. Selby records extensive damage in Ohio, involving even complete killing in some cases, by a temperature of 18° in October. Twenty degrees in October would seem more dangerous, particularly to nursery stock, than — 13° in December, the other month of record-making tempera- ture. The date of digging these trees is not known. If they were, ac- cording to the prevailing practice, dug in the fall, the period of injury is fixed without question. Without this evidence, however, the prob- ability of the October minimum being the chief factor in the damage is strong. Table 3.— Minimum Temperatures at Geneva, N. Y. (In degrees F.) Oct. Nov. Dec. Jan. Feb. 1917-18 26.0 9.0 — 18.0 — 10.0 — 11.0 1909-17 26.0 16.0 — 6.0 — 12.0 — 14.0 1883-1909 20.5 8.0 — 15.5 — 18.7 — 21.0 Table 3, compiled from reports of the New York Agricultural Experiment Station at Geneva, which is a considerable nursery cen- ter and located in a section where immaturity is generally known to be the chief factor in hardiness, is used here for comparison. The Octo- ber, 1917, minimum for St. Joseph is lower than at Geneva for the same year ; it is in fact a trifle lower than any in the long series of rec- ords for this place. Whether or not this particular injury was received in October, if immaturity is likely to be a factor in winter injury at Geneva, N. Y., it is likely to be a factor at St. Joseph, Mo. Table 4, giving in detail the data summarized in Tables 2 and 3, shows this clearly. In the nine years for which data are available, the October minimum for St. Jo- seph has been lower than that for Geneva in five, identical in three and higher in one. The November minimum has been lower in five years, higher in three and identical in one. If absolute cold at any time be considered the chief cause of injury, the Geneva absolute minimum of — 18°F. is offset by one of — 24°F. for St. Joseph. It is true that the higher maximum and mean temperatures of the fall months at St. Joseph may under certain conditions have some in- fluence in hastening maturity. It is also true, however, that they may Observations on Winter Injury 21 Plate \ . — Fig. 1 (Left) Jonathan, 4 years after planting in the orchard, showing growth condition and lack of external evidence that would indicate the condition shown in Fig. 7, a cross section of the same trunk. Fig. 2. (Right) Jonathan, four years after planting. Compare this view with the cross sections of the same tree shown in Fig. 3. Photographed Mar. 28, 1922. 99 Missouri Agr. Exp. Sta. Research Bulletin 56 Plate VI. — Fig 3. Jonathan, four years from time of planting, showing injured areas and heart rot rapidly advancing in later annual rings of apparently sound wood. Observations on Winter Injury 23 Plate VII. — big. 4. Jonathan, four years after planting showing longitu- dinal and cross section views of one tree through trunk and scaffold limbs. In a large percentage of the trees cut off, the injury terminated in a point a few inches above the ground. 24 Missouri Agr. Exp. Sta. Research Bulletin 56 1922. Photographed August 14, 1922. Observations on Winter Injury 25 have some influence in the opposite direction. The average rainfall at St. Joseph in August, September and October is greater than that at Geneva ; for these three months the figures are, respectively : at St. Joseph, 4.37, 3.34 and 2.62 inches; at Geneva, 3.30, 2.42 and 2.50 inches. When high rainfall is combined with high temperature, growth is prolonged and the first low temperatures are more likely to be dam- aging. Table 4.— Minimum Temperatures at St. Joseph, Mo., and at Geneva, N. Y., 1909-1918 Inclusive. Year October November December January February St. Joseph Geneva St. Joseph Geneva St. Joseph Geneva St. Joseph Geneva St. Joseph Geneva 1909-10 27 27 19 21 —10 1 —13 —8 —5 —3 3910-13 28 26 20 21 8 —2.5 —11 —1 9 4 1911-12 32 33 5 18 — 4 13 —24 —12 —6 —10 1912-13 31 31 20 20 8 12 — 4 8 —2 —10 1913-14 22 29 26 22 11 6 8 —9 —7 —14 1914-15 25 26 7 16 — 8 —6 —12 —3 13 —10 1915-16 29 29 23 21 4 4 —19 —3 —4 —8 1916-17 24 29 12 16 9 4 — 8 1 —16 —8 1917-18 20 (26 17 9 —13 —18 —19 10 —13 —11 IMPORTANCE In a large number of cases occurring in this section winter injury of apples does not command attention at the time of its occurrence. It may induce minor injuries, the consequences of which are not re- vealed until the original cause is obscured. When a crop of peach buds is killed the loss is plain, but when a small area of bark is killed it receives little attention until decay ensues and by this time possible winter injury is forgotten. This very subtlety of winter injury makes difficult any appraisal of its extent. In the case discussed here the in- jury was slight. It was noticed at the time of setting the trees, but was thought of no importance. It did not affect the growth of the trees and would have been forgotten but for the work of the wood- destroying fungi. Many cases undoubtedly occur without untoward consequence; in many others the trees will grow for some years and when they begin to go to pieces the evidence to connect the condition with a slight winter injury several years back will be scant indeed. TREATMENT Detailed account of the treatment given this orchard following diagnosis of the condition will be published elsewhere. Briefly sum- 26 Missouri Agr. Exp. Sta. Research Bulletin 56 marized, it consisted in cutting back to sound wood, frequently to within a few inches of the ground and grafting the stubs (Fig. 8). A few trees cut back without grafting, after growth had started and when the carbohydrate reserve was low, died. Those treated earlier, with grafts inserted in the crowns, have made a practically perfect stand and are growing vigorously. This procedure has involved the sacrifice of the wood grown in the four years these trees have stood in the orchard ; but, with proper attention to the grafts, it will ensure perfectly sound trees, with every promise of long life and productive- ness. CONCLUSIONS Though winter conditions rarely kill apple trees outright in this section, they may have hardly less serious consequences. Evidence of winter injury should, therefore, put the grower on his guard. If new evidence appears every few years it may signify the need of re- vision of his cultural practices. Injury to wood at any time justifies great care in pruning. If the cuts can be made far enough back to re- move all injured wood, there is little danger of infection. If the re- moval of all injured wood is not practicable there are two courses re- maining: (1) careful disinfection and painting of all wounds, (2) omission of pruning altogether till the cuts can be made in sound tissue growing subsequent to the freeze. The practicability of these methods will be discussed elsewhere. However, one guiding principle may be stated: injured wood should not be exposed. Sealed within living wood, it is harmless ; exposed it is a source of constant danger. UNIVERSITY OF MISSOURI COLLEGE OF AGRICULTURE AGRICULTURAL EXPERIMENT STATION RESEARCH BULLETIN 56 Observations on Winter Injury I — Early and Late Winter Injury II — An Aftermath of Winter Injury COLUMBIA. MISSOURI NOVEMBER, 1922 46-M Ik].22 UETIN COLOmIia. MO. 3 0112 019681935