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Effect of Sire of Fetus on Subsequent 90-Day Production of the Holstein Dam
Effect of Sire of Fetus on Subsequent 90-Day Production of the Holstein Dam J. F. HAYES, B. R. M A N G U R K A R , 1 and R. I. CUE Department of Animal Science Macdonald College of McGill University 21,111 Lakeshore Road Ste. Anne de Bellevue, Qu6bec, Canada H9X 1C0
ABSTRACT
Data on calving ease and 90-day milk, fat, protein, fat percent, and protein percent were available on 8,817 Holstein cows enrolled in the Quebec Dairy Herd Analysis Service. These observations were distributed in 802 herds, and they represented 124 service sires. Estimates of the variance components associated with the service sire (sire of fetus) were obtained by Minimum Norm Quadratic Unbiased Estimation procedures. Two models were fitted; one model included herd, month of calving, age of cow, and sire of cow as fixed effects and service sire and residual as random effects, whereas a second model included calving ease as an additional fixed effect. Variance components and percentages of variance accounted for by service sire were similar under both models. Proportions of variance accounted for by service sire were 1.1, .3, .5, .2, and .3% for 90-day milk, fat, protein, fat percent, and protein percent. INTRODUCTION
In dairy cattle, variability of production traits associated with sire of fetus has received considerable attention. There are differences in the estimates that partly reflect different methodologies and different degrees of nonrandomness of association of sire of fetus with sire of cow. By Henderson's Method 1, estimates of 8.2, 1.0, and 3.0% for the proportion of the phenotypic variance o f milk yield ac-
Received June 14, 1984. t The Bharatiya Agro-Industries Foundation, UruliKanchan, District Puna, India 412202. 1985 J Dairy Sci 68:1286-1289
counted for by sire of fetus in Holsteins were reported in (1, 5, 11). An estimate of .8% b y Henderson's Method 3 was in (5), and .9% b y Wickham with Minimum Norm Quadratic Unbiased Estimation (MINQUE) was in (16). As pointed out in (16), estimates are smaller the more nearly association of sire of fetus with sire of cow is random or the more the method of estimation accounts for any nonrandomness. The influence of sire of calf ( f e t u s ) o n the difficulty with which the calf is born in Holsteins has been reported in several studies. Heritabilities (from sire of fetus variance) of calving difficulty in heifers, later lactations, or all lactations combined were in (2, 9, 12), for example. Estimates of genetic correlation between calving ease in different parities were in (2, 8, 14), and evidence is in (14) that calving ease in different parities is essentially the same trait. To utilize variability of calving difficulty associated with sire of calf, sire evaluation for calving difficulty (of mates) was considered in (3,4,9). Additionally, there is evidence that calving difficulty affects subsequent production of the Holstein cow (6, 13, 15). Therefore, because variability of calving difficulty is associated with sire of calf (fetus), it is worth investigating if the effect of sire of fetus on production is operating partially through its effect on calving difficulty or if its effect on production is independent of, or over and above, its effect on calving difficulty. Our objective was 1) to estimate variance components associated with sire of fetus and the proportion of variance accounted for by sire of fetus for 90-day milk, fat, protein, fat percent, and protein percent in Holstein cows in Quebec by MINQUE procedures and 2) to compare these estimates under a model with and without calving ease-calf survival as fixed.
1286
SERVICE SIRE EFFECTS ON PRODUCTION MATERIALS AND METHODS
Records on 90-day production and calving ease were available for 35,773 Holsteins on the official test option of the Quebec Dairy Herd Analysis Service (DHAS) for calvings from September 1979 to December 1980. Calving ease was scored b y the dairy farmer as 1) unassisted or unobserved, 2) easy pull, 3) hard pull, 4) surgical, or 5) malpresentation; calf survival was scored as 1) alive (in first 24 h after birth) or 2) dead. There remained 8,817 observations after deletion of records because of malpresentations, lack of sire of cow identification, birth dates of cows, breeding and drying dates, service sire identification, and restriction of data to service sires having at least five pregnancies in at least two herds. These observations were distributed in 802 herds and represented 124 service sires. Estimates of sire of fetus and residual variances for 90-day milk, fat, protein yield, percent fat, and percent protein were obtained by fitting each of the following models by iterative MINQUE procedures (10): Yijklmno = I/+ hi + mj + a k + c 1 + dm + Sn + Eijklmno
[1]
Yijklrnn =/2 + hi + mj + a k + c1 + Sm + Eijklmn
[2]
where: /2 = population mean, h i = effect of the ith herd (absorbed), mj = effect of the mth month of calving, a k = effect of the kth age subclass (<3, 3 to 4, 4 to 5, 5 to 6, and > 6 yr), c I = effect of the lth sire of cow, d m = effect of the mth calving easecalf survival subclass (m = 1 . . . . 8), S m (or Sn) = effect of the mth sire of fetus (0, as2), and E = residual ~, (0, oe2). Standard errors for estimates of sire of fetus and residual variances, and o f the proportion of total variance accounted for by sire of fetus, were constructed from asymptotic variances
1287
and covariances of estimates of sire of fetus and residual variances. RESULTS A N D DISCUSSION
The residual variance component, the variance component associated with the sire of fetus, and the proportion of total variance accounted for b y sire of fetus ignoring calving ease are in Table 1 for 90-day milk fat and protein yields and 90-day fat and protein percent. The proportion of variance of 90-day milk yield accounted for b y sire of fetus was 1.1%. This estimate agrees in general with those in (5, 11, 16) for either complete or partial lactations and suggests that sire of fetus is not an important source of variation in 90-day milk yield. The variation associated with sire of fetus in 90-day fat and protein yields and 90-day fat and protein percent was .3, .5, .2, and .3%. Therefore, the effect of sire of fetus on 90-day production traits is small and of little consequence. Calving difficulty in cows often is considered a trait of the sire of the fetus that initiated the lactation. Efforts have been made to investigate the effect of sire of fetus, on the one hand, and calving difficulty, on the other hand, on the cow's production, It is interesting, therefore, to see if the effect of sire of fetus on production is changed when calving ease is included in the model. Results for sire of fetus when calving ease is included in the model are in Table 2.Results for variance components and proportions of variance accounted for by sire of fetus are similar to those in Table 1. Therefore, the effect of sire of fetus on production traits is independent of the effect of calving ease on production traits. This suggests that factors in the calf (contributed by its sire) that influence milk yield of its dam differ from factors in the calf that influence the difficulty with which it is born. The mechanisms b y which the sire of fetus influence subsequent production of the cow are not clear; variability of secretion of placental lactogen and estrogens b y the fetus or associated placenta has been pointed out as a possible mechanism (2). It also has been suggested that dosage of placental lactogen, related to the size of the calf, is involved. Factors that affect size of calf, which in turn affects calving difficulty, therefore would affect the dosage of placental lactogen and subsequent production. However, the contribution of sire of fetus to Journal of Dairy Science Vol. 68, No. 5, 1985
1288
HAYES ET AL.
TABLE 1. Residual (o~) and sire of fetus (o]) variance components and proportion of total variance accounted for by sire of fetus for 90-day production traits. Trait
o~
Milk yield Fat yield Protein yield Fat % Protein %
122,678.52 1,888,685.35 1,075,044.60 1,464.21 446.06
(1,982.39) 1 (30,467.20) (17,355.10) (23.61) (7.19)
1,398.28 4,897.55 5,503.00 2.52 1.11
o~
og/(o~ + a~)
(511.60) (4,026.80) (3,024.63) (2.71) (.94)
.011 .003 .005 .002 .003
(.004) (.002) (.003) (.002) (.002)
1 Standard errors in parentheses.
TABLE 2. Residual (of) and sire of fetus (o~) variance components and proportion of total variance accounted for by sire of fetus for 90-day production traits after adjusting for the effects of calving ease.
a~
Trait Milk yield Fat yield Protein yield Fat % Protein %
122,205.63 1,885,859.17 1,O71,867.87 1,464.00 445.54
(1,975.70)' (30,435.98) (17,312.O0) (23.61) (7.19)
1,399.51 4,997.70 5,582.06 2.56 1.12
o~
a~/(a~ + o~)
(511.24) (4,057.10) (3,040.90) (2.73) (.94)
.011 .003 .005 .002 .002
(.004) (.002) (.003) (.002) (.002)
Standard errors in parentheses.
p a r t u r i t i o n of t h e calf is n o t solely t h r o u g h calf size b u t also t h r o u g h h o r m o n e s s e c r e t e d b y t h e p l a c e n t a t h a t i n f l u e n c e p a r t u r i t i o n . T h e s e horm o n e s are n o t necessarily t h e same as t h o s e that influence subsequent production through d e v e l o p m e n t of t h e m a m m a r y system, i.e., p l a c e n t a l lactogen, w h i c h o c c u r s early in pregrmncy. F o r e x a m p l e , calving d i f f i c u l t y m a y b e i n f l u e n c e d b y s e c r e t i o n o f estrogenic h o r m o n e s late in p r e g n a n c y b y f e t a l m e m b r a n e s (7). ACKNOWLEDGMENTS
B. R. M a n g u r k a r a c k n o w l e d g e s receipt o f a C I D A - N S E R C fellowship. This w o r k was supp o r t e d in p a r t b y a g r a n t f r o m t h e Minist~re de l ' A g r i c u l t u r e , des Pecheries, et d e L ' A l i m e n t a t i o n o f Qu6bec.
REFERENCES
1 Adkinson, R. W., C. J. Wilcox, and W. W. Thatcher. 1977. Effects of sire of fetus upon subsequent production and days open of the darn. J. Dairy Sci. 60:1964. 2 Bar-Anan, R., M. Soller, and J. C. Bowman. 1976. Genetic and environmental factors affecting the Journal of Dairy Science Vol. 68, No. 5, 1985
incidence of calving difficulty and perinatal calf mortality in Israd-Friesian dairy herds. Anim. Prod. 22:299. 3 Berger, P. J., and A. E. Freeman. 1978. Prediction of sire merit for calving difficulty. J. Dairy Sci. 61: 1146. 4 Cady, R. A. 1980. Evaluation of Holstein bulls for dystoci~ Ph.D. Diss., Cornell Univ., Ithaca, NY. 5 Johnson, L. P., and L. D. Van Vleck. 1979. Components of variance associated with service sire for milk yield and reproductive traits. J. Dairy Sci. 62: 754. 6 Mangurkar, B. R., J. F. Hayes, and J. E. Moxley. 1984. Effects of calving ease-calf survival on production and reproduction in Holsteins. J. Dairy Sci. 67:1496. 70singa, A., and W. Hazelegee. 1977. The influence of the sire on the estrogen production of the bovine fetus. Page 7 in Eur. Econ. Commun. Seminar on calving problems and early viability of the calf. 8 Philipsson, J. 1976. Studies on calving difficulty, stillbirth and associated factors in Swedish cattle breeds. II. Effects of non genetic factors. Acta Agric. Scand. 26:165. 9 Pollak, E. J., and A. E. Freeman. 1976. Parameter estimation and sire evaluation for dystocia and calf size in Holsteins. J. Dairy Sci. 59:1817. 10 Rao, C. R. 1971. Estimation of variance and covariance components-MINQUE theory. J. Multivaro Anal. 1:257.
SERVICE SIRE EFFECTS ON PRODUCTION 11 Skjervold, H., and E. Fimland. 1975. Evidence for a possible influence of the fetus on the milk yield of the dam. Z. Tierz. Zfichtungsbiol. 92:245. 12 St. Arnaud, H., J. F. Hayes, and J. E. Moxley. 1980. A study of calving ease and calf survival in Quebec DHAS herds. Can. J. Anim. Sci. 60:1033. 13 Thompson, J. R., A. E. Freeman, and P. J. Berger. 1980. Relationship of dystocia transmitting ability with type and production transmitting abilities in Holstein bulls. J. Dairy Sci. 63:1462.
1289
14 Thompson, J. R., A. E. Freeman, and P. J. Berger. 1981. Age of dam and maternal effects for dystocia in Holsteins. J. Dairy Sci. 64:1603. 15 Thompson, J. R., A. E. Freeman, P. J. Berger, and M. L. Martinez. 1981. A survey of dystocia and calf mortality in five dairy breeds. J. Dairy Sci. 81(Suppl. 1):81. (Abstr.) 16 Van Vleck, L. D., and L. P. Johnson. 1980. Genetic and economic implications on fetal effects of the dam. J. Dairy Sci. 63:1483.
Journal of Dairy Science Vol. 68, No. 5, 1985
ABSTRACT
Data on calving ease and 90-day milk, fat, protein, fat percent, and protein percent were available on 8,817 Holstein cows enrolled in the Quebec Dairy Herd Analysis Service. These observations were distributed in 802 herds, and they represented 124 service sires. Estimates of the variance components associated with the service sire (sire of fetus) were obtained by Minimum Norm Quadratic Unbiased Estimation procedures. Two models were fitted; one model included herd, month of calving, age of cow, and sire of cow as fixed effects and service sire and residual as random effects, whereas a second model included calving ease as an additional fixed effect. Variance components and percentages of variance accounted for by service sire were similar under both models. Proportions of variance accounted for by service sire were 1.1, .3, .5, .2, and .3% for 90-day milk, fat, protein, fat percent, and protein percent. INTRODUCTION
In dairy cattle, variability of production traits associated with sire of fetus has received considerable attention. There are differences in the estimates that partly reflect different methodologies and different degrees of nonrandomness of association of sire of fetus with sire of cow. By Henderson's Method 1, estimates of 8.2, 1.0, and 3.0% for the proportion of the phenotypic variance o f milk yield ac-
Received June 14, 1984. t The Bharatiya Agro-Industries Foundation, UruliKanchan, District Puna, India 412202. 1985 J Dairy Sci 68:1286-1289
counted for by sire of fetus in Holsteins were reported in (1, 5, 11). An estimate of .8% b y Henderson's Method 3 was in (5), and .9% b y Wickham with Minimum Norm Quadratic Unbiased Estimation (MINQUE) was in (16). As pointed out in (16), estimates are smaller the more nearly association of sire of fetus with sire of cow is random or the more the method of estimation accounts for any nonrandomness. The influence of sire of calf ( f e t u s ) o n the difficulty with which the calf is born in Holsteins has been reported in several studies. Heritabilities (from sire of fetus variance) of calving difficulty in heifers, later lactations, or all lactations combined were in (2, 9, 12), for example. Estimates of genetic correlation between calving ease in different parities were in (2, 8, 14), and evidence is in (14) that calving ease in different parities is essentially the same trait. To utilize variability of calving difficulty associated with sire of calf, sire evaluation for calving difficulty (of mates) was considered in (3,4,9). Additionally, there is evidence that calving difficulty affects subsequent production of the Holstein cow (6, 13, 15). Therefore, because variability of calving difficulty is associated with sire of calf (fetus), it is worth investigating if the effect of sire of fetus on production is operating partially through its effect on calving difficulty or if its effect on production is independent of, or over and above, its effect on calving difficulty. Our objective was 1) to estimate variance components associated with sire of fetus and the proportion of variance accounted for by sire of fetus for 90-day milk, fat, protein, fat percent, and protein percent in Holstein cows in Quebec by MINQUE procedures and 2) to compare these estimates under a model with and without calving ease-calf survival as fixed.
1286
SERVICE SIRE EFFECTS ON PRODUCTION MATERIALS AND METHODS
Records on 90-day production and calving ease were available for 35,773 Holsteins on the official test option of the Quebec Dairy Herd Analysis Service (DHAS) for calvings from September 1979 to December 1980. Calving ease was scored b y the dairy farmer as 1) unassisted or unobserved, 2) easy pull, 3) hard pull, 4) surgical, or 5) malpresentation; calf survival was scored as 1) alive (in first 24 h after birth) or 2) dead. There remained 8,817 observations after deletion of records because of malpresentations, lack of sire of cow identification, birth dates of cows, breeding and drying dates, service sire identification, and restriction of data to service sires having at least five pregnancies in at least two herds. These observations were distributed in 802 herds and represented 124 service sires. Estimates of sire of fetus and residual variances for 90-day milk, fat, protein yield, percent fat, and percent protein were obtained by fitting each of the following models by iterative MINQUE procedures (10): Yijklmno = I/+ hi + mj + a k + c 1 + dm + Sn + Eijklmno
[1]
Yijklrnn =/2 + hi + mj + a k + c1 + Sm + Eijklmn
[2]
where: /2 = population mean, h i = effect of the ith herd (absorbed), mj = effect of the mth month of calving, a k = effect of the kth age subclass (<3, 3 to 4, 4 to 5, 5 to 6, and > 6 yr), c I = effect of the lth sire of cow, d m = effect of the mth calving easecalf survival subclass (m = 1 . . . . 8), S m (or Sn) = effect of the mth sire of fetus (0, as2), and E = residual ~, (0, oe2). Standard errors for estimates of sire of fetus and residual variances, and o f the proportion of total variance accounted for by sire of fetus, were constructed from asymptotic variances
1287
and covariances of estimates of sire of fetus and residual variances. RESULTS A N D DISCUSSION
The residual variance component, the variance component associated with the sire of fetus, and the proportion of total variance accounted for b y sire of fetus ignoring calving ease are in Table 1 for 90-day milk fat and protein yields and 90-day fat and protein percent. The proportion of variance of 90-day milk yield accounted for b y sire of fetus was 1.1%. This estimate agrees in general with those in (5, 11, 16) for either complete or partial lactations and suggests that sire of fetus is not an important source of variation in 90-day milk yield. The variation associated with sire of fetus in 90-day fat and protein yields and 90-day fat and protein percent was .3, .5, .2, and .3%. Therefore, the effect of sire of fetus on 90-day production traits is small and of little consequence. Calving difficulty in cows often is considered a trait of the sire of the fetus that initiated the lactation. Efforts have been made to investigate the effect of sire of fetus, on the one hand, and calving difficulty, on the other hand, on the cow's production, It is interesting, therefore, to see if the effect of sire of fetus on production is changed when calving ease is included in the model. Results for sire of fetus when calving ease is included in the model are in Table 2.Results for variance components and proportions of variance accounted for by sire of fetus are similar to those in Table 1. Therefore, the effect of sire of fetus on production traits is independent of the effect of calving ease on production traits. This suggests that factors in the calf (contributed by its sire) that influence milk yield of its dam differ from factors in the calf that influence the difficulty with which it is born. The mechanisms b y which the sire of fetus influence subsequent production of the cow are not clear; variability of secretion of placental lactogen and estrogens b y the fetus or associated placenta has been pointed out as a possible mechanism (2). It also has been suggested that dosage of placental lactogen, related to the size of the calf, is involved. Factors that affect size of calf, which in turn affects calving difficulty, therefore would affect the dosage of placental lactogen and subsequent production. However, the contribution of sire of fetus to Journal of Dairy Science Vol. 68, No. 5, 1985
1288
HAYES ET AL.
TABLE 1. Residual (o~) and sire of fetus (o]) variance components and proportion of total variance accounted for by sire of fetus for 90-day production traits. Trait
o~
Milk yield Fat yield Protein yield Fat % Protein %
122,678.52 1,888,685.35 1,075,044.60 1,464.21 446.06
(1,982.39) 1 (30,467.20) (17,355.10) (23.61) (7.19)
1,398.28 4,897.55 5,503.00 2.52 1.11
o~
og/(o~ + a~)
(511.60) (4,026.80) (3,024.63) (2.71) (.94)
.011 .003 .005 .002 .003
(.004) (.002) (.003) (.002) (.002)
1 Standard errors in parentheses.
TABLE 2. Residual (of) and sire of fetus (o~) variance components and proportion of total variance accounted for by sire of fetus for 90-day production traits after adjusting for the effects of calving ease.
a~
Trait Milk yield Fat yield Protein yield Fat % Protein %
122,205.63 1,885,859.17 1,O71,867.87 1,464.00 445.54
(1,975.70)' (30,435.98) (17,312.O0) (23.61) (7.19)
1,399.51 4,997.70 5,582.06 2.56 1.12
o~
a~/(a~ + o~)
(511.24) (4,057.10) (3,040.90) (2.73) (.94)
.011 .003 .005 .002 .002
(.004) (.002) (.003) (.002) (.002)
Standard errors in parentheses.
p a r t u r i t i o n of t h e calf is n o t solely t h r o u g h calf size b u t also t h r o u g h h o r m o n e s s e c r e t e d b y t h e p l a c e n t a t h a t i n f l u e n c e p a r t u r i t i o n . T h e s e horm o n e s are n o t necessarily t h e same as t h o s e that influence subsequent production through d e v e l o p m e n t of t h e m a m m a r y system, i.e., p l a c e n t a l lactogen, w h i c h o c c u r s early in pregrmncy. F o r e x a m p l e , calving d i f f i c u l t y m a y b e i n f l u e n c e d b y s e c r e t i o n o f estrogenic h o r m o n e s late in p r e g n a n c y b y f e t a l m e m b r a n e s (7). ACKNOWLEDGMENTS
B. R. M a n g u r k a r a c k n o w l e d g e s receipt o f a C I D A - N S E R C fellowship. This w o r k was supp o r t e d in p a r t b y a g r a n t f r o m t h e Minist~re de l ' A g r i c u l t u r e , des Pecheries, et d e L ' A l i m e n t a t i o n o f Qu6bec.
REFERENCES
1 Adkinson, R. W., C. J. Wilcox, and W. W. Thatcher. 1977. Effects of sire of fetus upon subsequent production and days open of the darn. J. Dairy Sci. 60:1964. 2 Bar-Anan, R., M. Soller, and J. C. Bowman. 1976. Genetic and environmental factors affecting the Journal of Dairy Science Vol. 68, No. 5, 1985
incidence of calving difficulty and perinatal calf mortality in Israd-Friesian dairy herds. Anim. Prod. 22:299. 3 Berger, P. J., and A. E. Freeman. 1978. Prediction of sire merit for calving difficulty. J. Dairy Sci. 61: 1146. 4 Cady, R. A. 1980. Evaluation of Holstein bulls for dystoci~ Ph.D. Diss., Cornell Univ., Ithaca, NY. 5 Johnson, L. P., and L. D. Van Vleck. 1979. Components of variance associated with service sire for milk yield and reproductive traits. J. Dairy Sci. 62: 754. 6 Mangurkar, B. R., J. F. Hayes, and J. E. Moxley. 1984. Effects of calving ease-calf survival on production and reproduction in Holsteins. J. Dairy Sci. 67:1496. 70singa, A., and W. Hazelegee. 1977. The influence of the sire on the estrogen production of the bovine fetus. Page 7 in Eur. Econ. Commun. Seminar on calving problems and early viability of the calf. 8 Philipsson, J. 1976. Studies on calving difficulty, stillbirth and associated factors in Swedish cattle breeds. II. Effects of non genetic factors. Acta Agric. Scand. 26:165. 9 Pollak, E. J., and A. E. Freeman. 1976. Parameter estimation and sire evaluation for dystocia and calf size in Holsteins. J. Dairy Sci. 59:1817. 10 Rao, C. R. 1971. Estimation of variance and covariance components-MINQUE theory. J. Multivaro Anal. 1:257.
SERVICE SIRE EFFECTS ON PRODUCTION 11 Skjervold, H., and E. Fimland. 1975. Evidence for a possible influence of the fetus on the milk yield of the dam. Z. Tierz. Zfichtungsbiol. 92:245. 12 St. Arnaud, H., J. F. Hayes, and J. E. Moxley. 1980. A study of calving ease and calf survival in Quebec DHAS herds. Can. J. Anim. Sci. 60:1033. 13 Thompson, J. R., A. E. Freeman, and P. J. Berger. 1980. Relationship of dystocia transmitting ability with type and production transmitting abilities in Holstein bulls. J. Dairy Sci. 63:1462.
1289
14 Thompson, J. R., A. E. Freeman, and P. J. Berger. 1981. Age of dam and maternal effects for dystocia in Holsteins. J. Dairy Sci. 64:1603. 15 Thompson, J. R., A. E. Freeman, P. J. Berger, and M. L. Martinez. 1981. A survey of dystocia and calf mortality in five dairy breeds. J. Dairy Sci. 81(Suppl. 1):81. (Abstr.) 16 Van Vleck, L. D., and L. P. Johnson. 1980. Genetic and economic implications on fetal effects of the dam. J. Dairy Sci. 63:1483.
Journal of Dairy Science Vol. 68, No. 5, 1985