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Animal Improvement Programs Laboratory, Agricultural Research Service, USDA, Beltsville, MD 20705-2350
Corresponding author: J. B. Cole; e-mail: jcole{at}aipl.arsusda.gov.
| ABSTRACT |
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Key Words: calving ease crossbred genetic evaluation purebred
Abbreviation key: BH = dataset with BS-sired pure-bred calvings, HO-sired purebred calvings, and BS-sired calvings from HO dams, BS = Brown Swiss, BSc = dataset with all calvings in BSp plus BS-sired crossbred calvings, BSp = dataset with BS-sired purebred calvings, CE = calving ease, %DBH = percentage of difficult births in heifers, DCE = daughter calving ease, HO = Holstein, JE = Jersey, JEc = dataset with all calvings in JEp plus JE-sired crossbred calvings, JEp = dataset with JE-sired purebred calvings, MGS = maternal grandsire, SCE = service-sire calving ease, S-MGS = sire-maternal grandsire.
| INTRODUCTION |
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Dairy producers are increasingly interested in cross-breeding. In a recent survey of US dairy producers using crossbreeding, almost all respondents indicated a desire to improve calving ease as well as health, fertility, and longevity (Weigel and Barlass, 2003). Holstein-Brown Swiss (BS) and HO-Jersey (JE) F1 both outperformed purebred HO for Net Merit and Cheese Merit, although no cross outperformed HO for Fluid Merit (VanRaden and Sanders, 2003). The authors also reported a small, favorable (1.2%) heterotic benefit for productive life. Heins et al. (2003b) reported that JE-HO crossbred heifers and cows had significantly lower phenotypic dystocia scores than purebred HO contemporaries, 1.32 vs. 1.94. A related study (Heins et al., 2003a) reported that JE-sired calves were born with significantly lower dystocia scores than BS-sired calves, and BS-sired calves had significantly lower dystocia scores than HO-sired calves in a population of HO, HO-JE, and HO-Normande cows. McClintock et al. (2004) presented further evidence that JE-HO crossbreds have a lower incidence of dystocia than purebred HO. No difference for dystocia was found between HO-JE and JE-HO calvings (Cassell et al., 2004), although the sample size was very small. Heins et al. (2004) reported that HO-sired calvings had significantly more dystocia than JE-sired calvings; HO cows also had higher rates of dystocia than Normande-HO, Montbeliarde-HO, and Scandinavian-HO cows. These results suggest that the use of sires from several non-HO breeds in a crossbreeding program may result in reduced incidence of dystocia.
In response to interest in calving ease from the Brown Swiss Association and the announcement of an Interbull pilot study of CE for breeds other than HO, the Animal Improvement Programs Laboratory studied CE in the BS and JE breeds. The objectives of this research were: 1) to determine the extent to which CE data are recorded in the BS and JE breeds; 2) to characterize the available CE data for BS and JE, as well as for BS- and JE-sired crosses; 3) to perform preliminary prediction of PTA for these breeds, as well as for BS-and JE-sired crosses, using the available data; and 4) to develop a procedure for routine national evaluations for the BS and JE breeds, if appropriate.
| MATERIALS AND METHODS |
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Difficult births, indicated by a CE score of 4 or 5, were combined into a single category for the JEp and JEc evaluations to attain convergence. Records from herds with only difficult calvings, or with only one calving record in the database, were omitted from the BS and JE datasets.
Genetic Evaluation Models
Purebred and crossbred evaluation.
The same S-MGS model as used for the routine HO genetic evaluation (Van Tassell et al., 2003) was used to analyze BSp and JEp datasets:
![]() | ([1]) |
where yijklnopr = CE score, hyi = random effect of herd-year i, YSj = fixed effect of year-season j, PSk = fixed effect of parity-sex k, SBl = fixed effect of sire birth year l, BMn = fixed effect of maternal grandsire (MGS) birth year n, slo = random effect of sire o in birth-year group l, mnp = random effect of MGS n in birth-year group p, and eijklnopr = random residual effect.
Parities were first, second, and third and later. Year-season groups begin in October and May. The model used to analyze BSc, JEc, and BH datasets was similar to [1] but included a fixed effect to account for breed composition (BCq). The breed composition effect had 2 levels in the BSc(JEc) data set to differentiate between births of purebred and crossbred calves. There were 3 levels of breed composition in the BH data set to differentiate between breeds of MGS (BS, HO, and all other). The (co)variance components estimated by Wiggans et al. (2003) were used for all analyses.
The same sire birth-year groups were defined for the BSp(JEp) and BSc(JEc) datasets:
1990, 1991 to 1995, and 1996, 1997, ..., 2003. Identical MGS birth-year group definitions were used for the BSp(JEp) datasets. Maternal grandsire birth years ranged from 1964 to 2001 for BS and 1958 to 2001 for JE. Different MGS groupings for animals with known MGS ID and with unknown MGS ID were used. For animals with known MGS ID, MGS birth years were
1985, 1986 to 1990, 1991 to 1995, and 1996, and 1997 for BSc and
1990, 1991 to 1995, and 1996 to 2000 for JEc. Records without valid MGS ID were assigned to birth-year groups based on dam birth year. When dam birth years were not recorded, they were approximated as calving year parity 1. Maternal grandsire birth-year groups for bulls without valid ID were:
1995 and >1995.
Genetic bases for service-sire CE (SCE) and daughter CE (DCE) were defined by bulls born in 1995 and in 1990, respectively. Sire and MGS solutions on the underlying scale were adjusted such that the mean of the base bulls on the observed scale was approximately equal to the mean percentage of difficult births in heifers (%DBH; CE scores of 4 or 5 for first-calf heifers giving birth to male calves) observed in the appropriate offspring (Van Tassell et al., 2003). Mean %DBH was estimated separately for each data set. The BH evaluation used the HO base for all animals although mean %DBH was slightly higher for HO than BS (8.1 vs. 7.6%). The choice of base does not affect within-breed rankings.
Joint Brown Swiss-Holstein evaluation.
The inclusion of HO records in the evaluation was expected to benefit BS sires with a large number of crossbred calvings, as well as BS sires whose daughters have HO contemporaries. The joint evaluation used the same sire and MGS birth-year group definitions as the routine HO evaluation and groups included sires of both breeds. Different groups were not used for each breed due to the small number of BS sires and similar trends over birth years in individual breed datasets. Groupings by breed may be necessary in the future to account for differing amounts of selection pressure on calving ease.
The approach used to calculate %DBH was modified to account for breed-of-MGS effects. Van Tassell et al. (2003) computed %DBH as:
![]() | ([2]) |
![]() | ([3]) |
where F = standard normal cumulative density function, T3 = the threshold between CE scores of 3 and 4 on the observed scale,
= the solution on the underlying scale with fixed MGS birth year and breed-of-MGS solutions added to the MGS solution, and * denotes the group of animals used to define the base. A constant, c, is used to achieve the desired base, and c in [2] is replaced with
in [3]. In a single-breed evaluation,
consists only of the random MGS solution plus the fixed MGS birth-year group solution. In a multiple-breed evaluation,
should be computed as the sum of the random MGS solution, the fixed MGS birth year solution, and the fixed breed-of-MGS solution.
| RESULTS |
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Distributions of calving ease scores by data set are presented in Table 1
. The purebred and crossbred datasets had similar distributions of scores within sire breeds and very different distributions between sire breeds. Jersey sires produced calves that resulted in fewer difficult births (CE scores
4) than did BS sires. Brown Swiss sires produced calves with higher frequencies of difficult births than JE sires but with slightly lower frequencies than HO sires. Similar trends were seen in score distributions by parity. In all breeds, the frequency of difficult births was highest in first parity. The difference between first and second parity was much greater than the difference between second and later parities.
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Jersey.
Distributions of CE scores by parity for the purebred (data not shown) and crossbred (Table 3
) JE datasets indicated more difficult births in first than later parities. For first parity, the incidence of calving difficulty (scores of 4 or 5) in the JEc data set was 1.1%, the highest value for any parity in either data set. Scores of 1 (no problem) were recorded for more than 92% of all calvings. This is consistent with evidence that JE are easy calvers (Thompson et al., 1981). Statistics of solutions to the S-MGS model for the crossbred JE evaluations are presented in Table 4
. The range and SD of the solutions are similar to results from the purebred JE evaluation (data not shown) and the routine HO evaluation (Van Tassell et al., 2003), although the range and SD of the herd-year solution is smaller than for HO. This may reflect the fact that there was a greater number of herd-years in the HO data than either JE data set. The sire and MGS birth-year group effects are consistent with the lack of genetic trend for %DBH (Figure 1
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Distributions of service sire and daughter %DBH (Figure 2
) indicated %DBH of 2 and 1 are most common for SCE and DCE, respectively, in the JEc data set. Only 2 JE bulls had a PTA > 2 for SCE, and only one had a PTA > 2 for DCE. No JE bull had a PTA > 3 for any trait in either data set. Clearly, relative to other breeds, there are very small genetic differences among JE sires and MGS for either SCE or DCE. There are adequate CE data available in both breeds for genetic evaluation. Although there is sufficient genetic variability in the BS to make a routine genetic evaluation worthwhile, that is not the case with the JE.
Joint Brown Swiss-Holstein Evaluation
Results of the BS and JE evaluations indicate that routine evaluation of BS sires is possible. Many BS records come from herds that also provide HO records, and F1 calvings with non-HO sires and HO dams have reduced dystocia as compared with purebred HO calvings, suggesting that a joint evaluation using both the BS and HO data is desirable. The Netherlands (Interbull, 2004b) and New Zealand (Interbull, 2004c) use records from multiple breeds in a joint evaluation in their national genetic evaluations for CE.
Genetic evaluation.
Distribution of CE scores by parity for the BH data set is shown in Table 3
. Difficult births accounted for 8, 3.3, and 3.1% of all calvings in first, second, and third parities, respectively. Scores of 4 and 5 occurred more often in this data set than in the BSc and had frequencies similar to the routine HO evaluation (data not shown). Addition of BS data to HO data did not change the distribution of scores. Statistics of solutions to the S-MGS model for the joint evaluation are presented in Table 4
and are very similar to reported results (Van Tassell et al., 2003). The range of the herd-year solution is larger than that reported for Holsteins in Van Tassell et al. (2003) but is comparable to the value (7.06) obtained in the routine August 2004 HO evaluation. The sire and MGS birth-year group effects are consistent with the lack of genetic trend for %DBH presented in Figure 3
. Breed-of-MGS solutions were small (BS: 0.51, HO: 0.55, other: 0.48) and consistent with expectations; smaller values are more favorable.
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![]() | ([4]) |
where reli,t is the reliability of sire i for trait t (sire or MGS effect), diagi,t is the reciprocal of the diagonal element from the coefficient matrix, and
t is the genetic standard deviation of trait t. The approximation assumes that all relatives and fixed effects in the model are perfectly estimated. Because of this assumption, reli,t computed in [4] is independent of the number of contemporaries. Berger (1994) showed sire model evaluation of calving ease requires a large number of effective progeny to achieve high reliabilities. Given the simplification used in [4], a larger number of effective progeny is probably required under the S-MGS model than a sire model. Although an improved method for computing reliabilities is desirable, it may not be of great practical significance. Berger (1991) found that all of the reliability approximations reviewed overestimated prediction error variances but were highly correlated with true prediction error variances. As long as reliabilities are consistently overestimated and selection pressure on CE remains low, consequences of this overestimation are minimal.
Validation.
The joint evaluation was validated by comparing results with those from the BSc and routine HO evaluations. Brown Swiss records made up less than 0.15% of total records evaluated and therefore should have little impact on HO evaluations. Product-moment correlations among service sire and daughter evaluations (Table 5
) confirm notable changes in service sire and daughter evaluations for BS sires with negligible changes for HO sires. Rank correlations were similar to product-moment correlations in all cases. Characteristics of the changes in solutions and PTA between the single-breed and joint evaluations are presented in Table 6
. Brown Swiss evaluations changed moderately for both traits when moving from the single-breed to the joint evaluation; some reranking of sires also occurred. Changes in BS rankings are attributable to changes in sire and MGS solutions as well as sire and MGS birth year solutions.
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Genetic trend validation (Boichard et al., 1995) was performed on the BSc and BH datasets (results not shown). Method 3 validation, which compares sire evaluations over time as daughter records are added, was used. There should be no systematic change in sire PTA over time. The BSc validation did not produce a significant result due to the small amount of data available. The BH data set passed trend validation using the specifications provided by Interbull (Interbull, 2004a).
| DISCUSSION |
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It was assumed the true values of (co)variance components used in the analysis were the same for all breeds and were equal to HO (co)variances (Wiggans et al., 2003). This seems reasonable for BS and HO, but may not be valid for JE. If this assumption is incorrect, the (co)variances used are probably overestimates for the JE population. The PTA from the JE analyses are very small, and if overestimated should not result in erroneous conclusions. Results suggest essentially no genetic variation exists for calving ease in the JE breed whether the heritability in the model was correct.
Heterosis was not directly accounted for in the crossbred evaluations. However, only F1 were included in the BSc, JEc, and BH datasets. The breed composition effects in the BSc and JEc models were equivalent to a heterosis effect with two levels, 0% (for purebreds) and 100% (for F1) because they distinguished between purebred and crossbred calvings. The breed composition effect used in the BH analysis differentiated between different breeds of MGS. Although this is not directly equivalent to general heterosis, it does account for some of the same variation because the model only considers purebreds and F1. If the joint evaluation model is extended to accommodate crossbreds other than F1, it will be necessary to include a covariate to account for general heterosis (VanRaden, 1992). This is a less sophisticated approach than the crossbred model proposed by Lo et al. (1997), but Lutaaya et al. (2002) found that the use of a crossbred model in swine is not justified when the evaluation of purebreds is the principal goal and the number of crossbred records is small relative to the number of purebred records. It is reasonable to assume the same would be true for dairy cattle.
The reliability approximation currently used does not increase when the number of daughter herdmates increases. A key argument in favor of developing a joint evaluation is that BS sires with many crossbred daughters, and whose crossbred daughters are found predominantly in HO herds, benefit from the inclusion of those data in the evaluation. There is substantial reranking of BS sires when comparing the BSc and BH evaluations, suggesting the joint evaluation does impact BS sires. Distributions of reliabilities from the BSc and BH evaluations were very similar for BS bulls.
The development and implementation of a routine national CE evaluation for BS bulls allowed the US to contribute those data to the international CE evaluation under development by Interbull. Contribution of BS data to the Interbull effort is desirable because US sires provide pedigree connections among several European populations.
| CONCLUSIONS |
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A joint evaluation for calving ease of BS and HO in the US has been developed. The joint evaluation utilizes information from purebred and crossbred (BS-HO) calvings of BS bulls, as well as purebred calvings of HO bulls. The joint evaluation does not adversely affect HO sires but does result in reranking of BS bulls as compared with an evaluation of BS data only. Evaluations from data for the August 2004 run were provided to Interbull for inclusion in the international pilot run for calving ease.
| ACKNOWLEDGEMENTS |
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Received for publication September 30, 2004. Accepted for publication December 19, 2004.
| REFERENCES |
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