Israeli Holstein Population Research Articles

Genetic and Genomic Analysis of Cow Mortality in the Israeli Holstein Population

"Livability" was defined as the inverse of the probability of death. The objectives of this study were to estimate the heritability, genetic and phenotypic trends for the livability of Israeli Holstein cows; estimate the genetic and environmental correlations between livability and the nine traits included in the Israeli breeding index; estimate the effect of the inclusion of livability in the Israeli breeding index on expected genetic gains; and compute a genome-wide association study (GWAS) for livability. Seven data sets were analyzed. All data were derived from the database of the Israeli dairy cattle herd-book. The mean livability for the complete data set of 523,954 cows born from 2000 through 2016 was 89.6%. Pregnancy reduced livability by 15%. Livability generally increased with parity and days in milk within parity. Heritability of livability was 0.0082. Phenotypic and genetic trends over the 14-year period from 2000 through 2013 were -0.42% and -0.22% per year. If livability is included in the Israeli breeding index, accounting for 9% of the index, livability would increase by 1.3% and protein production would decrease by 11 kg over the next decade, as compared to the current index. A marker in proximity to the oxytocin-vasopressin locus had the greatest effect in the GWAS. Oxytocin activity in cattle affects calving-associated pathologies and maternal death. Inclusion of livability in the Israeli breeding index is not recommended.

Breeding Dairy Cattle for Female Fertility and Production in the Age of Genomics.

Phenotypic and genetic changes for female fertility and production traits in the Israeli Holstein population over the last three decades were studied in order to determine if long term selection has resulted in reduced heritability and negative genetic correlations. Annual means for conception status, defined as the inverse of the number of inseminations to conception in percent, decreased from 55.6 for cows born in 1983 to 46.5 for cows born in 2018. Mean estimated breeding values increased by 1.8% for cow born in 1981 to cows born in 2018. Phenotypic records from 1988 to 2016 for the nine Israeli breeding index traits were divided into three time periods for multi-trait REML analysis by the individual animal model. For all traits, heritabilities increased or stayed the same for the later time periods. Heritability for conception status was 0.05. The first parity genetic correlation between conception status and protein yield was -0.38. Heritabilities decreased with the increase in parity for protein but remained the same for conception status. Realized genetic trends were greater than expected for cows born from 2008 through 2016 for somatic cell score, conception status and herd-life, and lower than expected for the production traits.

Short communication: Selection for persistency by changing the shape of lactation curves with geometric morphometrics

The objective was to simulate the response to selection by modifying the shape of the lactation curve to improve persistency in Israeli first-parity Holstein cattle using geometric morphometrics and multivariate quantitative genetics. The 2-dimensional Cartesian coordinates representing the shape of the lactation curve were characterized using geometric morphometrics. Genetic parameters of daily milk yield were estimated for the individual animal model using REML. Results of selection were simulated using the multivariate breeders' equation. We defined 6 selection regimens: (1) decrease in the third month in milk (MIM), (2) increase in the ninth MIM, (3) decrease in the third MIM and increase in the ninth MIM, (4) decrease in the first MIM, (5) decrease in the second MIM, and (6) decrease in the fourth MIM. The first 3 selection criteria targeted changes in the MIM of previous measurements of persistency in the Israeli Holstein population, and the last 3 targeted the months with the highest variability. All selection regimens attempted to modify the shape of the lactation curve while keeping total milk yield constant. The first 3 selection regimens showed greater total responses, with the highest response for decrease in the third MIM and increase in the ninth MIM. Magnitudes of correlated responses consistently surpassed direct responses in all selection criteria, suggesting that selection on specific parts of the lactation curve would produce correlated responses in other parts. Predicted responses to selections were depicted as shape changes from the mean shape in the data set. Most predicted responses demonstrated a fairly similar shape compared with the lactation curve. The predicted responses of lactation shapes of the last 3 selection regimens were overall lower in the first 3 MIM and decreased less in the last MIM, demonstrating a much flatter shape and thus improved persistency.

Optimization of a genomic breeding program for a moderately sized dairy cattle population

Although it now standard practice to genotype thousands of female calves, genotyping of bull calves is generally limited to progeny of elite cows. In addition to genotyping costs, increasing the pool of candidate sires requires purchase, isolation, and identification of calves until selection decisions are made. We economically optimized via simulation a genomic breeding program for a population of approximately 120,000 milk-recorded cows, corresponding to the Israeli Holstein population. All 30,000 heifers and 60,000 older cows of parities 1 to 3 were potential bull dams. Animals were assumed to have genetic evaluations for a trait with heritability of 0.25 derived by an animal model evaluation of the population. Only bull calves were assumed to be genotyped. A pseudo-phenotype corresponding to each animal's genetic evaluation was generated, consisting of the animal's genetic value plus a residual with variance set to obtain the assumed reliability for each group of animals. Between 4 and 15 bulls and between 200 and 27,000 cows with the highest pseudo-phenotypes were selected as candidate bull parents. For all progeny of the founder animals, genetic values were simulated as the mean of the parental values plus a Mendelian sampling effect with variance of 0.5. A probability of 0.3 for a healthy bull calf per mating, and a genomic reliability of 0.43 were assumed. The 40 bull calves with the highest genomic evaluations were selected for general service for 1 yr. Costs included genotyping of candidate bulls and their dams, purchase of the calves from the farmers, and identification. Costs of raising culled calves were partially recovered by resale for beef. Annual costs were estimated as $10,922 + $305 × candidate bulls. Nominal profit per cow per genetic standard deviation was $106. Economic optimum with a discount rate of 5%, first returns after 4 yr, and a profit horizon of 15 yr were obtained with genotyping 1,620 to 1,750 calves for all numbers of bull sires. However, 95% of the optimal profit can be achieved with only 240 to 300 calves. The higher reliabilities achieved through addition of genomic information to the selection process contribute not only in obtaining higher genetic gain, but also in obtaining higher absolute profits. In addition, the optimal profits are obtained for a lower number of calves born in each generation. Inbreeding, as allowed within genomic selection for the Israeli herd, had virtually no effect on genetic gain or on profits, when compared with the case of exclusion of all matings that generate inbreeding. Annual response to selection ranged from 0.35 to 0.4 genetic standard deviation for 4 to 15 bull sires, as compared with 0.25 to 0.3 for a comparable half-sib design without genomic selection.

Maternity validation using sire‐only BovineSNP50 BeadChip data

Based on pairwise identity-by-state (IBS) distances and whole-genome SNP data, kinship was investigated in the Israeli Holstein population. A total of 789 bulls, including most of the artificial insemination sires in service since 1987, were genotyped by the BovineSNP50 BeadChip. This sample included up to five generations. For each bull-by-bull combination, three states are possible for each marker: no match, a single match and both alleles match. Summing over all markers, the 932598 IBS scores (three match frequencies*310866 bull-by-bull combinations) were visualized using three-dimensional coordinates that corresponded to the frequencies of the three possible states. Results were reduced to two dimensions using the transformations x'=0.7071(1+freq1-freq2) and y'=1.2247freq0. Bull-by-bull pairs were grouped according to their level of kinship, and canonical scores were calculated using discriminant analysis and the x' and y' features. Of the 474 pairs of recorded maternal grandsire-grandson with both individuals genotyped, the probability for 28 pairs to belong to this level of kinship was low (P<0.05), suggesting an error rate of around 3% per generation in pedigree determination.

Genetic parameters for paratuberculosis infection and effect of infection on production traits inIsraeliHolsteins

Paratuberculosis (Johne's disease) is an infectious enteric disease in dairy cattle and other species that causes substantial economic loss worldwide. In this study, two recursive Gaussian-threshold models were employed in order to infer the effects of Johne's disease on milk yield, fat yield, and protein yield while simultaneously estimating genetic parameters (i.e. heritability and genetic correlation) in an Israeli Holstein population. Disease diagnosis was based on ELISA serum antibody tests. Data were available for 4694 daughters of 361 sires; 3.5% were positive; and 1.6% were suspect for the disease test. Disease status was coded either as a binary character (negative vs. positive) or as an ordered categorical trait (negative, suspect, and positive) in the two recursive models and as a binary trait in a linear model. Among sires with ≥ 50 daughters, predicted probability of Mycobacterium avium ssp. paratuberculosis (MAP) infection in future daughters ranged from <1% to 16.5%. Heritability estimates for Johne's disease were near 0.15, confirming a genetic contribution to disease susceptibility. Genetic correlation estimates for Johne's disease with the three yield traits were 0.15-0.22. Residual correlations for Johne's disease with the yield traits were between -0.01 and -0.10. For the linear regression model, yield losses associated with a positive disease diagnosis during 305 days of lactation were 300 kg milk and around 10 kg for fat and protein. Yield loss estimates from the recursive models were 25-50% less than linear model estimates. Recursive modeling has theoretical advantages over linear models for these phenotypes, but the estimated genetic parameters in this study did not differ significantly between the two types of models.

Short communication: Association of an OLR1 polymorphism with milk production traits in the Israeli Holstein population

Previous studies have reported significant associations between haplotypes of the oxidized low-density lipoprotein (lectin-like) receptor 1 (OLR1) gene and a single nucleotide polymorphism (SNP) in its 3′-untranslated region with milk composition and health traits in different cattle populations. However, to provide a better estimation of the impact and size of the association of OLR1 with production traits, a need exists to validate its genomic association in additional cattle populations. Thus, the objective of this study was to perform association analysis of the OLR1 SNP with milk traits in the Israeli Holstein population. Estimated breeding values of milk composition traits and somatic cell score were obtained for a total of 1,211 Holstein cows, which were genotyped for a SNP in the 3′-untranslated region. Statistical analysis revealed significant association between the OLR1 SNP and protein percentage and somatic cell score. Thus, the validation of association of this SNP with milk traits in independent cattle populations suggests OLR1 as a candidate gene for further functional studies.

Fine Mapping of a QTL for Fertility on BTA7 and Its Association With a CNV in the Israeli Holsteins

A quantitative trait locus (QTL) affecting female fertility, scored as the inverse of the number of inseminations to conception, on Bos taurus chromosome 7 was detected by a daughter design analysis of the Israeli Holstein population (P < 0.0003). Sires of five of the 10 families analyzed were heterozygous for the QTL. The 95% confidence interval of the QTL spans 27 cM from the centromere. Seven hundred and four SNP markers on the Illumina BovineSNP50 BeadChip within the QTL confidence interval were tested for concordance. A single SNP, NGS-58779, was heterozygous for all the five QTL heterozygous patriarchs, and homozygous for the remaining five QTL homozygous sires. A significant effect on fertility was associated with this marker in the sample of 900 sires genotyped (P < 10−6). Haplotype phase was the same for four of the five segregating sires. Thus concordance was obtained in nine of the ten families. We identified a common haplotype region associated with the rare and economically favorable allele of the SNP, spanning 270 kbp on BTA7 upstream to 4.72 Mbp. Eleven genes found in the common haplotype region should be considered as positional candidates for the identification of the causative quantitative trait nucleotide. Copy number variation was found in one of these genes, KIAA1683. Four gene variants were identified, but only the number of copies of a specific variant (V1) was significantly associated with breeding values of sires for fertility.

Differential expression of ruminant ZNF496 variants: Association with quantitative trait locus affecting bovine milk concentration and fertility1,2

A single nucleotide polymorphism in the intergenic region upstream of the ZNF496 gene on Bos taurus chromosome 7 displayed significant population-wide linkage disequilibrium with milk protein percentage in the Israeli Holstein population. The frequency of the allele associated with increased protein concentration was 10%. This single nucleotide polymorphism was located in the promoter region from which a 10-exon transcript of the bovine and the ovine ZNF496 genes are transcribed. The gene architecture was similar to the mouse ortholog Zkscan17. A 5-exon murine antisense transcript was complementary to the 5′ untranslated Zkscan17 region that included a sequence domain conserved between mouse and ruminants, suggesting a regulatory function. In the bovine ZNF496 chromosomal region, segregation of a quantitative trait locus (QTL) for milk protein percentage was confirmed in a daughter design sire family. Concordance was not obtained between QTL status of bulls and any of the polymorphisms in the functional elements of ZNF496. This excludes these variations as the causative polymorphism under the assumption of no epigenetic effect for this locus. However, ZNF496 variants were differentially expressed in bovine ovaries, and only the paternal variant was expressed in liver and kidney in a sheep family with polymorphic ZNF496 sequence. Thus, the search for the mutation underlying the minor QTL allele, which is a top economically favorable allele in Israeli Holstein cattle, may be complicated by the presence of an imprinting center in this QTL confidence interval.

Incorporation of genotype effects into animal model evaluations when only a small fraction of the population has been genotyped

The method of Israel and Weller (Estimation of candidate gene effects in dairy cattle populations. Journal of Dairy Science 1998, 81, 1653–1662) to estimate quantitative trait locus (QTL) effects when only a small fraction of the population was genotyped was investigated by simulation. The QTL effect was underestimated in all cases, but bias was greater for extreme allelic frequencies, and increased with the number of generations included in the simulations. Apparently, as the fraction of animals with inferred genotypes increases, the genotype probabilities tend to ‘mimic’ the effect of relationships. Unbiased estimates of QTL effects were derived by a modified ‘cow model’ without the inclusion of the relationship matrix on simulated data, even though only a small fraction of the population was genotyped. This method yielded empirically unbiased estimates for the effects of the genes DGAT1 and ABCG2 on milk production traits in the Israeli Holstein population. Based on these results, an efficient algorithm for marker-assisted selection in dairy cattle was proposed. Quantitative trait loci effects are estimated and subtracted from the cows’ records. Genetic evaluations are then computed for the adjusted records. Animals are then selected based on the sum of their polygenic genetic evaluations and QTL effects. This scheme differs from a traditional dairy cattle breeding scheme in that all bull calves were considered candidates for selection. At year 10, total genetic gain was 20% greater by the proposed algorithm as compared to the selection based on a standard animal model for a locus with a substitution effect of 0.5 phenotypic standard deviations. The proposed method is easy to apply, and all required software are ‘on the shelf.’ It is only necessary to genotype breeding males, which are a very small fraction of the entire population. The method is flexible with respect to the model used for routine genetic evaluation. Any number of genetic markers can be easily incorporated into the algorithm, and the reduction in genetic gain due to incorrect QTL determination is minimal.

Genetic Analysis of Persistency in the Israeli Holstein Population by the Multitrait Animal Model

Persistency was defined as the predicted milk production 180 d after peak divided by peak production (in %). Heritability of persistency in a multitrait analysis including parities 1 through 5 increased from 0.16 to 0.27 from first through third parity, and then declined through fifth parity. Genetic correlations for persistency between consecutive parities were all>0.8. First-parity genetic correlations of the traits included in the Israeli selection index with persistency were all<0.1, except for fertility and herdlife, which were 0.20 and 0.25; whereas second-parity genetic correlations of persistency with the 3 milk production traits were all>0.34, and the genetic correlation with fertility was only 0.10. The genetic correlation between second-parity persistency and herdlife was 0.58. Persistency in the Israeli Holstein population was analyzed by the multitrait animal model. The genetic trend since 1985 for the multiparity index was 0.22% persistency/yr, even though there was no direct selection on persistency.

Correcting for bias in estimation of quantitative trait loci effects

Estimates of quantitative trait loci (QTL) effects derived from complete genome scans are biased, if no assumptions are made about the distribution of QTL effects. Bias should be reduced if estimates are derived by maximum likelihood, with the QTL effects sampled from a known distribution. The parameters of the distributions of QTL effects for nine economic traits in dairy cattle were estimated from a daughter design analysis of the Israeli Holstein population including 490 marker-by-sire contrasts. A separate gamma distribution was derived for each trait. Estimates for both the α and β parameters and their SE decreased as a function of heritability. The maximum likelihood estimates derived for the individual QTL effects using the gamma distributions for each trait were regressed relative to the least squares estimates, but the regression factor decreased as a function of the least squares estimate. On simulated data, the mean of least squares estimates for effects with nominal 1% significance was more than twice the simulated values, while the mean of the maximum likelihood estimates was slightly lower than the mean of the simulated values. The coefficient of determination for the maximum likelihood estimates was five-fold the corresponding value for the least squares estimates.

Correcting for bias in estimation of quantitative trait loci effects

Estimates of quantitative trait loci (QTL) effects derived from complete genome scans are biased, if no assumptions are made about the distribution of QTL effects. Bias should be reduced if estimates are derived by maximum likelihood, with the QTL effects sampled from a known distribution. The parameters of the distributions of QTL effects for nine economic traits in dairy cattle were estimated from a daughter design analysis of the Israeli Holstein population including 490 marker-by-sire contrasts. A separate gamma distribution was derived for each trait. Estimates for both the α and β parameters and their SE decreased as a function of heritability. The maximum likelihood estimates derived for the individual QTL effects using the gamma distributions for each trait were regressed relative to the least squares estimates, but the regression factor decreased as a function of the least squares estimate. On simulated data, the mean of least squares estimates for effects with nominal 1% significance was more than twice the simulated values, while the mean of the maximum likelihood estimates was slightly lower than the mean of the simulated values. The coefficient of determination for the maximum likelihood estimates was five-fold the corresponding value for the least squares estimates.

Factors Affecting Incorrect Paternity Assignment in the Israeli Holstein Population

A total of 6040 Israeli Holstein cows from 181 Kibbutz herds listed as progeny of 11 sires were genotyped for 104 microsatellites. Seventeen markers were deleted due to a frequency of erroneous genotypes >1%, leaving 160,470 valid genotypes. Conflicts between the putative sire and daughter in at least 2 markers and for at least 10% of the markers genotyped per cow were required to reject paternity. Cows that did not meet the requirements for paternity confirmation or rejection were deleted from further analysis. The frequency of rejected paternity was 11.7%. The effects of recorded sire, birth year, geographical region, herd, and inseminator on the frequency of paternity rejection were analyzed with linear and nonlinear models. Only the effects of inseminator and recorded sire were significant in all models tested that included these effects. The main causes of incorrect paternity recording appear to be inseminator recording mistakes, and possibly mistakes with respect to semen labeling at the AI institutes. Incorrect paternity recording due to multiple inseminations by different sires could explain, at most, 20% of the paternity mistakes. Instituting a system of quality control, especially at the level of the inseminator, should reduce paternity errors to no more than 8%, and increase genetic progress by at least 1%.

A complete genome scan of the Israeli Holstein population for quantitative trait loci by a daughter design.

Eleven Israeli Holstein families including 5221 cows were analyzed by a daughter design for eight economic traits: milk, fat and protein production, fat and protein percentage, somatic cell score (SCS), herd-life, and female fertility. The cows were genotyped for 73 microsatellites with maximum spacing between markers of 53 cM. There were 86,304 informative genotypes. Preliminary analysis was by ANOVA of each trait, with the marker effect nested within sire. Significance was determined by controlling the false discovery rate at 0.4, after excluding markers with genome-wide significance for at least a single trait, and traits without any significant effects at this level. Thus, four markers on chromosomes 6 and 14 and female fertility were excluded. There remained 40 significant marker-trait combinations, and it is expected that 24 of these are true effects. To perform interval mapping for the families with significant contrasts, 21 additional markers were genotyped on chromosomes 2, 7, and 27. The bootstrap confidence intervals for gene effect did not include zero for protein percent on chromosome 2 and fat yield, protein yield, and SCS on chromosome 7. Quantitative trait locus heterozygosity was 33%, which is consistent with the hypothesis that only two alleles are segregating with unequal allele frequency.

Population-Wide Analysis of a QTL Affecting Milk-Fat Production in the Israeli Holstein Population

A population-wide linkage disequilibrium on bovine chromosome 14 between microsatellite ILSTS039 and DGAT1, a putative quantitative trait locus affecting milk production traits, was found in the Israeli Holstein population. A total of 394 bulls were genotyped for both DGAT1 and ILSTS039, and 1747 cows were genotyped for ILSTS039. The ILSTS039 allele termed “225,” and the DGAT1 K allele (substitution of a lysine residue with alanine), were associated with decreased milk production, and increased fat production and fat and protein percent. The number of 225 ILSTS039 and K DGAT1 alleles per individual were the same for 80% of the bulls genotyped. From the effects associated with cows homozygous for the 225 allele, the effect of the quantitative trait locus appears to be approximately codominant. The substitution effect was 0.16% fat. Genotype probabilities for the quantitative gene were determined for the entire Israeli Holstein milk-recorded population, including 507,725 cows and 1442 bulls, using segregation analysis. Overall frequency of the allele that increased fat percent was 8.9% in cows and 15.5% in bulls. The frequency of this allele decreased from 1981 until 1990, from 15 to 5%, and since has increased to 10%. The effects estimated on the population-wide analyses of both cows and bulls were similar to the effect associated with DGAT1 in the daughters of genotyped bulls. Modified animal model evaluations were computed for the entire population with the effect of this gene included in the model. The correlations between the modified and standard animal model evaluations for all traits were>0.99.

Multiple quantitative trait locus analysis of bovine chromosome 6 in the Israeli Holstein population by a daughter design.

Nine Israeli Holstein sire families with 2978 daughters were analyzed for quantitative trait loci effects on chromosome 6 for five milk production traits by a daughter design. All animals were genotyped for 2 markers. The three families with significant effects were genotyped for up to 10 additional markers spanning positions 0-122 cM of BTA6. Two sires were segregating for a locus affecting protein and fat percentage near position 55 cM with an estimated substitution effect of 0.18% protein, which is equivalent to one phenotypic standard deviation. This locus was localized to a confidence interval of 4 cM. One of these sires was also heterozygous for a locus affecting milk, fat, and protein production near the centromere. The hypothesis of two segregating loci was verified by multiple regression analysis. A third sire was heterozygous for a locus affecting milk and protein percentage near the telomeric end of the chromosome. Possible candidates for the major quantitative gene near position 55 cM were determined by comparative mapping. IBSP and SSP1 were used as anchors for the orthologous region on human chromosome 4. Twelve genes were detected within a 2-Mbp sequence. None of these genes have been previously associated with lactogenesis.

Multiple genotype analysis and sexing of IVF bovine embryos

Twenty-one in vitro-fertilized bovine blastocysts were quartered, lysed and subjected to primer elongation preamplification (PEP) procedure, allowing for the analysis of up to 40 genotypes per quarter embryo. The quarter-embryos were sexed by polymerase chain reaction (PCR) using BRY.1, Bov97M and ZFX/ZFY loci, and then genotyped for k-casein, bovine leukocyte adhesion deficiency (BLAD) and microsatellite D9S1. The mitochondrial cytochrome B locus was used as an internal control with a 95% success rate. The PEP procedure amplified genomic fragments in 93% of all cases. The embryos were identified to be 11 males and 10 females. Sexing accuracy was 87% for BRY.1, 97% for ZFX/ZFY and 100% for Bov97M. False genotyping was due mostly to amplification of BRY.1 in the female embryos and to the nonamplification of the ZFY locus in the male embryos. The results indicate that the combined use of Bov97M and ZFX/ZFY loci is a highly accurate procedure for sexing bovine embryos. Genotyping for κ-casein, D9S1 and BLAD was successful in 94, 99 and 91% of assays, respectively. Sex ratios and allele frequencies of embryos for gk-casein, BLAD and D9S1 were all close to the observed frequencies in the Israeli Holstein population. These results support the conclusion that the genotyping of embryos is as accurate as that of mature animals. Thus, marker-assisted selection can be efficiently applied at the preimplantation embryo level for loci of economic importance.

Sequence variation in D‐loop mtDNA of cow lineages selected for high and low maternal effects on milk production

The four highest and four lowest cow lineages for milk production were selected from 324 maternal lineages of the Israeli Holstein population. For a single cow from each lineage, the mitochondrial D-loop region was amplified using the polymerase chain reaction and directly sequenced for 719bp. Eighteen polymorphic sites were found of which 17 were substitutions and one was an insertion. Only two alleles were found for each polymorphic site, one of which corresponded to the sequence of Anderson et al. (1982), which was the more frequent allele, in all but two sites. None of the polymorphic sites had an apparent effect on milk production, even though an effect > or = 0.33 phenotypic standard deviations, equivalent to 470 kg milk, could be detected by this experimental design. Furthermore, there was no indication that moderately increasing the sample size would have altered the result, since all rare alleles found in more than a single lineage were found in both the high and low subgroups.

Sire and Maternal Grandsire Effects on Calving Difficulty and Calf Mortality in Israeli Holsteins

Associations among sire and maternal grandsire evaluations for calving difficulty and calf mortality were investigated in the Israeli Holstein population. Primiparous and multiparous calvings were analysed separately. Genetic correlations between calving difficulty and calf mortality were close to unity for primiparous calvings but lower for multiparous calvings. Genetic correlations between primiparous and multiparous sire evaluations were .94 and .72 for calving difficulty and calf mortality. Genetic correlations between sire and maternal grandsire evaluations were .6 for primiparous calvings and not-significant for multiparous calvings for both traits. These results correspond to findings that the direct genetic effect is larger than the maternal effect but that there is a slight negative correlation between them.We suggest that sire evaluations for calving difficulty of multiparous cows be used as a preliminary prediction of heifer calving difficulty and calf mortality and that the separate calving trait evaluations be combined into an overall calving selection index.