— The Effect of the Age of Sire and Dam on Offspring in Dairy Cows C. L. ALLEN Reprinted without change of paging, from the Journal of Heredity (Organ of the American Genelic Association), Vol. XIII, No. 4; Wash- ington, D. C., April. 1922, feb Eee ChOF THE-AGE OF SIRE AND DAM ON THE QUALITY OF OFFSPRING IN DAIRY COWS A THESIS PRESENTED TO THE FACULTY OF THE GRADUATE SCHOOL OF CORNELL UNIVERSITY IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY BY CHARLES LORING ALLEN hl | 1920 Reprinted from THE JoURNAL OF HEREDITY (Organ of the American Genetic Association), Vol. XIII, No. 4; Washington, D. C., April, 1922. \ 7 7 t en t ot: aie ; L & 1 = * ‘ e ' > ¢ , 4 py Sans a) imag Psy (niet! ao “a eis ns P ~ ~ | es +e ‘ T ~ e - la -- . it — if >. “y i . i 4 A i i “ ’ . 4 oe mY a a 7 ij ee H, | ’ , a‘! fe y 7 a ion ist sa Meck. whith? pera aa ite ae, Buty we ; H 1 ¥ ok ae bet le ay Palin ae, be é ; ah - 7 .) € 7 Wy = Pe, 5 " i Reprinted, without change of paging, from the Journal of Heredity (Organ of the American Genetic Association), Vol. XIII, No. A; Washington, D. C., April, 1922 THE EFFECT OF THE AGE OF SIRE AND DAM ON THE QUALITY OF OFFSPRING IN DAIRY COWS C.-L. ALLEN Cornel) University, Ithaca, N. Y. A FAMOUS HOLSTEIN SIRE FiGurE 10. Ormsby Korndyke Lad (102469) one of the great sires of the Holstein Friesian Breed. His daughters are noted for their high production records for both 7 day and yearly work. He has 18 daughters with records of over 30 pounds of butter (80% fat) in 7 days. He also has 14 daughters with records of over 1000 pounds of butter (80% fat) in one year. The sire of Ormsby Korndyke Lad was only 23 months of age when he was born and his dam was just a little over 4 years of age (51 months). Photograph from Holstein Friesian World. HE effect of the age of parents on the quality of the offspring is ina large measure concerned with the idea of cumulative inheritance, par- ticularly if we do not consider the matter of the proven sire or dam. If an animal is a more valuable parent after it has attained maturity, it is more valuable for one of two rea- sons. Either the mature animal must be able, merely because of its full physical development better to im- press its characters on its offspring, or the mature animal must in the process of attaining maturity, have become endowed with something, due to train- ing or environment, which it can trans- mit to its progeny. After a century of study, this matter is far from settled and the question as to the possible effect of environment on the transmis- sion of a tendency to increase or de- crease certain quantitative functions is still important. 167 168 The Journal of Heredity JOHANNA DE KOL CONCORDIA At three and a half years of age her record was 19037 pounds of milk and 671 Her sire, Sir Clothilde Concordia, was two years nine months of age when she was born, while her dam, Cold Spring Johanna De Kol 2nd, was three years and a half old. The great number of cases where the parents of high producing cows are as young, or younger than these, leaves no doubt that the off-spring of immature parents are just as good producers or FIGuRE 11. pounds of butterfat. transmitters of production as those born when the same parents are mature. Department of Agriculture. (See text, p. 173.) In practical breeding, it would be very advantageous to know definitely whether the young born of immature parents are as valuable for production, and for reproducing their kind, as those born of more mature parents. It is the aim of this paper to show whether the parentage of a group of high producing or superior cows aver- ages older than the parentage of a group of comparatively low producing or inferior cows. The paper also attempts to show the percentage distribution of offspring for the various ages of both sire and dam; the age when cows actually make their best records; and whether the offspring of very young or very old animals are inferior. Photo by U. S. METHOD The data reported in this paper were taken from Volume 27 of the Ad- vanced Registry Year book and the Herd Books of the Holstein Friesian Association of America. The superior class of animals chosen include those that up to April 30th, 1916, had made records of 24 pounds or more of butterfat in seven days. This included all of the so-called thirty pound cows since 24 pounds of butterfat is equal to 30 pounds of eighty percent butter. For purposes of more careful study, this class was divided into three groups as follows: Group I—AIl cows producing over 27.3 pounds of butterfat in 7 days. Allen: Effect of Parents’ Age on Quality of Dairy Cows Group II—AIl cows producing be- tween 25.6 and 27.3 pounds of butter- fat in 7 days. Group IIi—AIll cows producing be- tween 24.0 and 25.6 pounds of butter- fat in 7 days. The comparatively inferior class of animals chosen includes all of the so- called mature cows, that is, cows five years of age or over, that produced less than fourteen pounds of butter- fat in seven days and were entered in the 27th Volume of the Holstein Frie- sian Advanced Registry Year Book as entries or re-entries. This class was divided into three groups that were fairly comparable in numbers to the groups of the superior class, as follows: Group I—AIl cows producing be- tween 12 and 12.5 pounds of butter- fat in 7 days. Group II—AIll between 12.5 and butterfat in 7 days. Group III—AIl cows producing be- tween 13.0 and 14.0 pounds of butter- fat in 7 days. The groups were arranged so that the very highest producers were in group 1 of the Superior Class and the very lowest producers were in group 1 of the Inferior Class. Then, if there was any difference in age, it was thought that the first groups should be further apart than the second and third groups. The 7 day record was chosen as the basis for this study because a larger cows producing 13.0 pounds of 169 number of records were available and the 7 day record is generally recog- nized as a valuable method of deter- mining the productive ability of dairy cows. Three generation pedigrees were compiled for all the cows of both the Superior and Inferior Classes. These pedigrees included the date of birth for all animals in every pedigree. This makes it possible to compute the dif- ference in age between any animal and its parents and tells us the age of the parents at the time the animal in question was born. This also provides data for determining the average differ- ence in age between generations, and allows the calculation of the percentage distribution of offspring according to the age of parents, making it possible to observe whether more good cows or poor cows are born from very young or very old dams. The difference in age between parent and offspring was computed in months to the nearest month. In order to determine at what age the superior cows made their records, it was necessary to take this data directly from the Advanced Registry Year book and record it on the pedi- gree. RESULTS—THE DIFFERENCE IN AGE COMPARED Table I gives the average age of the ancestry of the Superior and the Infer- ior Classes by groups and for the classes TABLE I. Average Age, in Months, of Ancestry of Superior and of Inferior Classes of Cows Inferior Cows Superior Cows Ancestor Group 1|Group 2|Group 3 SiLereecsemsc a 44.3 Sil 7 46.3 | DE eva gyetinge, oa cages etd Ih CSc) 4) Wily Paternal grand sire.|. 48.6 49.6 49.6 Paternal grand dam.| 64.9 65.2 67.6 Maternal grand sire.| 42.3 41.4 42.3 |42 Maternal grand dam| 65.2 63.0 59.6 Great grand sire!...| 44.3 44.8 48.1 Great grand dam!..| 61.3 59.0 63.8 Number of pedigrees in each group and Glass yaad: 86 121 404 1 Great grand sire refers to sire of paternal grand sire. the maternal grand dam. Class Group 1/Group 2|Group 3 Class 14.647] 45.7 | 44.6 | 41.6 [43.14 .529 .9+.890] 60.0 | 62.2 6457 63-0 2944 .5+.717] 45.4 | 46.4 | 46.7 |46.6+ .646 .7+.929] 66.5 | 69.4 | 65.8 |66.7+1.034 14.507] 44.8 | 43.0 | 43.4 143.6+ .508 SOEE Shop Oo 0 | 6954. 16278) O47 OS .9+.640] 45.4 | 47.9 | 43.8 |45.1+ .602 poe WOOO) Olen UPA GAN 64.7 |65.7+1.006 611 111 114 285 510 Great grand dam refers to the dam of 170 as a whole. This table allows us to make comparisons and determine any significant difference between the aver- age age of the superior cows and the (comparatively) inferior cows. It will be noted in table I that the greatest difference in age is four months and that this occurs in the case of the sires. This difference might seem to be somewhat significant in the light of the probable error as a measure of its probable significance. The difference is more than three times the probable error but the prob- able error in this case is small. We must remember, however, that four months is not a very marked difference and this difference loses a large part of its significance when we study the group averages. There is more differ- ence between the groups within the classes even here than there is between the classes themselves. This is still more noticeable in other places in the table where the class difference is not so great. Take for example, the case of the dams, the great granddams, or the maternal grand sire. In each of these cases, the difference is probably insignificant in the light of the prob- able error, and in each case, the differ- ence between the groups within the class is greater than the difference between the classes. The real difference in age between the various parents and generations is more clearly shown by means of a pedigree. Such a pedigree is given in Figure 12. It is evident that there can be no increased hereditary value due to the age of the dams, for in every case ex- cept one, the dams of the Inferior Class average older, though not significantly so, than the dams of the Superior Class. In the one case, that of the paternal granddam, they average exactly alike. From a study of Table I and the pedigree, Figure 12, it is apparent that there is no significant difference between the ages of parents of the Inferior and of the Superior Classes. In only one case is the difference out- side the limits defined by three times the probable error. In this case, the The Journal of Heredity small difference between the class averages can hardly be considered sig- nificant when there is so great a differ- ence between the group averages within the class. ° BULLS ARE NO MORE VALUABLE PARENTS AS THEY GROW OLD Turning our attention again to the pedigree, Figure 12, we note that in every case the Superior sires on the top line of the pedigree are older than the corresponding inferior sires. The dif- ference is probably not significant even in the case of the sires, but the tendency persists. This led to a study to determine whether a _ bull produced superior offspring as he grew older. From a study of four bulls having a total of five hundred and six daughters, it seems that bulls may or may not have better daughters as they grow older, depending on the quality of animals to which they are bred. Asa bull proves his superior worth, he is very likely to be bred to better cows and produce better offspring. This probably ac- counts for the difference shown in the pedigree. This whole matter will be fully discussed in a later paper. Thus the difference in age of sires can be very readily assigned to the use of the sires and there is no indication that the sires are any better when they are old than when young and immature. In fact the frequency distribution, Figures 13 and 14, tends to show that the young parent of either sex is just as valuable as a parent when immature as it is in later life. The frequency curves include only the sires and dams of the first generation. FREQUENCY DISTRIBUTION ACCORD- ING TO AGE OF SIRE In the frequency curves, Figures 13 and 14, it will be observed that the curves run very nearly parallel throughout. The curves start high, indicating that a large percentage of cows in both the Superior and Inferior Classes are sired by very young bulls. In this connection, it will be remembered from Table I that Allen: Effect of Parents’ Age on Quality of Dairy Cows 49.5 46.0 GrandiSines3-nes0.8e Grands Damen eee 66.7 60.7 42.1 43.6 iy A 46.9 45.1 Great Grand Sire GrandsSire= eee Grands Damen: 61.0 64.7 | Great Grand Dam 62.5 65.7 AN AGE-PEDIGREE OF THE TWO CLASSES OF COWS FiGurE 12. This shows the average age of the ancestors of the superior cows (Bold-face type), and of the inferior cows (Italics). of the forebears of the two classes. the sires of the Superior Class average less than four years of age, 46.4 months to be exact, and that the sires of the Inferior Class average somewhat younger. The largest number of off- spring were born when their sires were from two and one-half to three years of age. This means that these bulls sired a larger percentage of their off- spring between twenty-two and twenty- seven months of age than at any other time. Table II shows very clearly just how our purebred sires are used. It indicates that improvement has been based on the selection of young sires rather than on the use of tried sires. THE PROVEN SIRE A bull cannot be called a proven sire until his first crop of heifers come in milk and can. hardly be so called until these heifers have milked at least one year. Thus a bull can hardly It is evident that there are no significant differences in the age (See text, p. 169.) be called a proven sire until he is five years of age. In general practice, the percentage of cows sired by unproven bulls is probably much higher than the percentage shown for the animals in- cluded in this study. This means that even the breeders who are making records use a tried sire for only a small percentage of their cows. That the tried sire is more valuable is indi- cated by the fact that 21.1 percent of the Superior cows are daughters of tried sires, whereas only 15.9 percent of the Inferior cows are daughters of bulls of an equal age. This may seem to conflict with the idea that the bull when old is not more valuable, but the paper referred to above shows that he is not more valuable as a parent. - FREQUENCY DISTRIBUTION ACCORD- ING TO AGE OF DAM The frequency curve for the dams, Figure 13, is very much more irregular than the curve for the sires but on the PERCENTAGE OF YOUNG BORN 172 The Journal of Heredity _ a BSL Obeemer ec meee - SSS 24 30 36 42 48 54 60 66 72 78 84 390 96 102 108 114 '20 126 132 138 144 150 156 162 168 174 180 186 192 198 204 AGE OF DAMS IN MONTHS FREQUENCY CURVE FOR DAMS FiGuRE 13. The percentage of young born by dams of various ages is shown in this chart. The age of the dams is computed for six-months periods, and the upper age limit of these groups is shown. That is, all the dams 24 months and under are considered 24 months old, those between 24 and thirty months of age, as 30 months old, etc. It is evident that much the greater number of oe are born by very young cows, the quality of whose offspring cannot be known. (See text, p. 1 Allen: Effect of Parents’ Age on Quality of Dairy Cows 173 whole, even here, the curves run nearly parallel. There is, however, a distinctly larger percentage of high producers born of very young dams than there is of low producers. Itis hard to account for this unless the Superior Class have received better care and are bred younger on the average than the In- ferior Class. It will be noted in connection with Table DM that a. slightly. larger percentage of the Superior Class were born from dams under five years of age than there were of the Inferior Class. In later life, the two classes are very near- ly alike and almost exactly as many of the Superior Class were born from dams over eight years of age as were born in the Inferior Class brome, Gams: of. a like age. The per- centage having dams over twelve years is exactly alike for both classes. VALUE OF OFFSPRING OF VERY YOUNG AND VERY OLD PARENTS From a study of the frequency curves and Tables II and III, it is evident that the young born from very young parents or from very old parents are as valuable as any for productive SUPERIOR CLASS) = INFERIOR CLASS 0.0 24 30 36 42 48 54 60 66 72 at 84 90 96 102 108 114 120 126 132 138 144 AGE OF Backes IN MONTHS FREQUENCY CURVE FOR SIRES FiGurE 14. The co-ordinates are the same as in the preceding chart. The same conclusions apply even more strikingly in this case, as the sires are not used for breeding purposes as long as the dams. A sire cannot be considered a proven sire until his first crop of daughters have been in milk at least a year, which means that he must be five years of age. These figures are for a selected class of sires and show that even the breeders who are making records use tried sires for only a small percentage of their cows. (See text, p. 170.) 174 The Journal of Heredity WATSON SEGIS PONTIAC HOMESTEAD FiGurRE 15. She was born when her dam, Watson Segis Pontiac, was just 2 years, 2 days old, when her sire, Piebe Laura Allie Homestead King, was a senior two-year-old. Her record is 25,360 pounds of milk and 928 pounds of fat. Her mother has a record of 844 pounds of fat, and she is the daughter of King Segis Pontiac Count, a great breeding sire, who was only a little over two years old when she was born. Photo by U.S. Department of Agriculture. The Percentage of Daughters Born on or before a Given Age of Sire, and the Age of the Sire at the Time the Dams of the Daughters were Bred TABLE II. Percentage of Daughters Age of Sire when Daughters} Age of Sire when the Dams of were born Daughters were Bred Superior Inferior 30 Months and Under 21 Months and Under PXf as 28.0 36 Months and Under 27 Months and Under 45.3 45.3 60 Months and Under 51 Months and Under 78.9 84.1 Over 60 Months Over 51 Months Mil Ah 15.9 TABLE III. Showing the Percentage of Young Born Before and After a Given Age of Dam Age of Dam when Daughters Percentage of Superior Percentage of Inferior were born Daughters Daughters 30sMonthsjandsUndersese.. sess. ti 2 IDS JOeMonthsrand (Winders oneness ae ZR 2 21.6 60 Months and Under............. Si. 54.9 OversSuvearsen caer aee aie athe tke on 14.8 15.0 Ver eVieanss «Scere ass 2.8 2.8 Allen: Effect of Parents’ Age on Quality of Dairy Cows TS A SIRE OF RECORD-MAKING DAUGHTERS FicureE 16. King Hengeveld Aaggie Fayne (56635) has shown remarkable ability to transmit production. He was slaughtered when only three years of age and asa result has only 31 A. R. O. daughters. Eleven of these daughters have records in excess of 30 pounds of butter (80% fat) in 7 days. Four of his daughters have yearly records. Three of these records made by two and three year old heifers average 21,762.2 pounds of milk and 978.93 pounds of butterfat. The other yearly record was made by a mature daughter and is 15,486.1 pounds of milk and 530.89 pounds of butterfat. King Hengeveld Aaggie Fayne was born when his parents were quite young. His sire was 22 months of age and his dam 28 months of age when he was born. Photo by the Holstein-Fresian World. purposes. An unusually large percent- age of the Superior Class were born before their dams were two and one- half years of age and must have been first calves. The sires of a large per- centage of the Superior Class were also under two and one-half years when the dams were bred. On the other hand, just as large a percentage of the Superior Class as of the Inferior Class is from very old parents. AGE WHEN THE BEST RECORDS ARE MADE Table IV was arranged for the Supe- rior Class only. The best record was used in every case. If any of the cows herein considered have made a better record since the collection of this data this fact introduces an error. It is recognized that only a very few cows have ever made more than one TABLE IV. The Average Age of All the Superior Cows at the Time Their Records Were Made | Group I Group II Group III Superior Class Number of Cows........ 86 121 404 611 Average Age in Months.. 69.7 Hj Al Teer! UPd! 176 record above thirty pounds of butter in a week and most of those that have done so are here considered. This error, then, is in all probability very small. The group averages and the class average all show that the best records of the animals herein considered were made at an average age of about six years. Of course many were much older than this and many were younger, but it is well known that animals vary greatly in rapidity of development. It would seem that the numbers con- sidered are large enough to make the average significant even though it is recognized that most cows do not have a chance to make a record every year. The cow is most likely to be tested at the time she shows greatest promise. Since these high producers made their best records at six years of age, it seems only fair to assume that they were at the height of their productive capacity at this time. It would also indicate that cows generally reach the 1 PEARL RAyMonpD, The Change of Milk Flow with Age. Station Bulletin, No. 262. 2 GOWAN, Bulletin, No. 293. 1917. 1920. J. W., Studies in Milk Secretion VIII. The Journal of Heredity limit of their possibilities for high production at about this age. This contention is supported from other sources,! although one investigator,” who attacked the problem”™ in an entirely different manner, maintains that the highest production is not reached until much later in life. SUMMARY The parentage of superior or high producing dairy cows is no older than the parentage of comparatively inferior or low producing cows and thus the old animal is no more valuable as a parent than the young animal. As far as the age of either parent is concerned, the young born at one time in life seem to be just as valuable as the young born at any other time. A very large percentage of our cows are born of immature parents but the proven animal at any age is much more valuable for breeding purposes. On the average, cows make their best records at about six years of age. Maine Agricultural Experiment Maine Agricultural Expiriment Station pt danke > + 4 ey he pas Aes Wl