Variance and covariance components for weaning weight for Herefords in three countries

Records from the Hereford Associations of the United States (USA), Canada, and Uruguay were used to estimate genetic and phenotypic variances and covariances for weaning weight. Estimation was done using a complete animal model, relatively large data sets, and the same methodology for the three countries in order to determine whether genetic parameters for weaning weight were homogeneous across environments. Data were composed of 2,322,722, 487,661, and 102,986 edited weaning weight records for USA, Canada, and Uruguay, respectively. Ten samples were obtained from each country by eliminating data from small herds with fewer than 500 records, selecting herds at random from the entire data set after removing the small herds, and then retaining the direct-sire-connected contemporary groups within each sample. The final sample sizes ranged from 9,832 to 46,377 records. An accelerated EM-REML algorithm was used in estimating the (co)variance components in each sample. The estimates were pooled by calculating the arithmetic mean of the 10 samples from within each country. Direct and maternal (in parentheses) heritability estimates were .24 (.16), .20 (.16), and .23 (.18) for USA, Canada, and Uruguay, respectively. Maternal heritabilities reported here are nearly 50% smaller than the values currently used in national genetic evaluation for the breed, which were estimated using sire-maternal grandsire models. Covariance between direct and maternal was negative in all countries, accounting for 6, 8, and 10% of the total phenotypic variation, and the total dam effect was 32.5, 37.0, and 34.0% in USA, Canada, and Uruguay, respectively. Total heritabilities were similar among the countries, with values of .19, .19, and .17 for the three respective countries. The similarity of genetic and environmental parameters across the three countries suggests that joint genetic evaluation is feasible across environments provided that the genotype x environment interaction is negligible and can be ignored.     

Estimation of variance and covariance components to determine heritabilities and repeatability of weaning weight in American Simmental cattle

Components of (co)variance for weaning weight were estimated from field data provided by the American Simmental Association. These components were obtained for the observational components of variance corresponding to a sire, maternal grandsire, and dam within maternal grandsire model. From these estimates, direct additive genetic variance (Sigma2A), maternal additive genetic variance (Sigma2M), covariance between direct and maternal additive genetic effects (SigmaAM), variance of permanent environment(Sigma2pe) and temporary environment variance(Sigma2te) were determined. A procedure to approximate restricted maximum likelihood (REML) estimates of the observational components of variance based on the expectation-maximization (EM) algorithm is described. From these results, phenotypic variance ( ) of weaning weight was 667.88 kg2. Values forSigma2A, Sigma2M, Sigma2pe and Sigma2te were 79,30,58,38,49.45, and 469.97 kg2, respectively. Genetic correlation between direct and maternal additive genetic effects was .16.     

Sire x herd interactions for weaning weight in beef cattle.

Weaning weight records of 44,357 Australian Angus calves produced by 1,020 sires in 90 herds were used to evaluate the importance of sire x herd interactions. Models fitted fixed effects of contemporary group (herd-year-date of weighing subclass), sex, calf age, and dam age and random effects of sire or of sire and sire x herd interaction using REML. Effects of standardizing the data, including sire relationships and including dam maternal breeding values (MBV) as a covariate were also investigated. Sire x herd interactions were found (P less than .05) in all cases and, in the most complete model, accounted for 3.3% of phenotypic variance. Across-herd heritabilities ranged from .19 to .28. Differential nonrandom mating among herds seemed to occur in the data. Significant sire x herd effects were observed for dam MBV, and adjustment for dam MBV yielded the smallest estimates of interaction variance and across-herd heritability. If sire x herd interactions were due only to genotype x environment interaction, within-herd heritabilities would range from .33 to .49. These estimates are larger than previously reported estimates. Thus, unreported environmental effects common to progeny of individual sires may also be involved in the observed interaction but could not be disentangled from true genotype x environment interaction effects using these data. Results of these analyses suggest that some accommodation of sire x herd interaction effects on weaning weight may be needed in beef cattle genetic evaluations, but a compelling case for development of herd-specific breeding value prediction cannot be made.     

Selection for weaning weight and postweaning gain in Hereford cattle. II. Response to selection

Single trait selection was practiced in three lines of Hereford cattle at two locations. Bulls were selected within sire families for increased weaning weight (WW) in the WW line (WWL), for postweaning gain (PG) in the PG line (PGL) and at random in the control line (CTL). Data include the performance of 2,467 calves produced from 1967 to 1981. Environmental effects were estimated from CTL (method I) and from multiple regression procedures (method II). Phenotypic and environmental time trends were negative for WW and generally were positive for PG. Estimated genetic gains for WW in WWL were 1.07 +/- .51 kg/yr in bulls and .62 +/- .36 kg/yr in heifers using method I and .50 +/- .31 kg/yr in bulls and .10 +/- .17 kg/yr in heifers using method II. Corresponding values for PG in PGL were .85 +/- .40 and 1.03 +/- .24 kg/yr in bulls and .30 +/- .28 and .37 +/- .12 kg in heifers. Correlated genetic gains for WW in PGL were larger than direct WW gains, whereas genetic gains for PG in WWL were smaller than direct PG gains. From method I, estimates of realized heritability (h2R) for WW were .31 +/- .18 in bulls and .22 +/- .13 in heifers. For PG, h2R was .31 +/- .13 in bulls and .06 +/- .12 in heifers. Using method II, h2R for WW was .09 +/- .08 in bulls and .02 +/- .07 in heifers. Corresponding values for PG were .29 +/- .10 and .11 +/- .08. Joint estimates of the realized genetic correlation between WW and PG were .69 +/- .18 and .46 +/- .31 for methods I and II, respectively. Variation in selection response was evaluated using quasi-replicates. Results of this study indicate that selection for PG improved both WW and PG faster than selection for WW.     

Selection for increased weaning or yearling weight in Hereford cattle. II. Direct and correlated responses

Selection was applied from 1964 to 1978 for increased weaning weight (WWL) or yearling weight (YWL) in two Hereford lines. An Angus line was maintained as an unselected control line (CL). Each line was maintained with 50 cows and four sires each year (two sires selected each year and used for 2 yr). Primary traits measured in the lines were birth weight (BW), preweaning daily gain (WDG), weaning weight (WW), weaning conformation grade (WG), weaning condition score (WC), weaning to yearling daily gain (YDG), yearling weight (YW), yearling conformation grade (YG) and yearling condition score (YC). Averaged over two methods, estimated genetic responses/generation (in standard deviation units) in WWL and YWL were: BW, .29, .26; WDG, .17, .15; WW, .22, .19; WG, .19, .26; WC, .12, .12; YDG, -.02, .04; YW, .08, .14; YG, .19, .16; YC, -.13, -.03. The realized heritability estimates were .23 and .15 for WW and YW, respectively. The realized genetic correlation between WW and YW was .69. Progeny from crosses of selected WWL and YWL sires to Angus cows had similar feedlot and carcass performance. At the end of the study, milk yield and composition were similar for mature cows in WWL and YWL.     

Selection for weaning weight and postweaning gain in Hereford cattle. I. Population structure and selection applied

Single trait selection was practiced in three lines of Hereford cattle derived from a common base population. Selection was practiced on males only within sire families for increased weaning weight (WW) in the WW line (WWL), for postweaning gain (PG) in the PG line (PGL) and at random in the control line (CTL). Females were culled on the basis of age or reproductive failure. Progeny of selected bulls were produced in two herds from 1970 through 1981. The data consisted of records on 2,467 progeny of 125 sires and 922 dams. Generations of selection to produce the 1981 calf crop were 1.96, 1.85 and 1.80 for WWL, PGL and CTL, respectively. For calves born in 1981, mean cumulative selection differentials (CSD) were 54.5 kg in WWL and 37.8 kg in PGL. Corresponding values in standard deviation units (SDU) were 2.31 and 1.68, respectively. Secondary selection differentials were 25 to 40% as large as selection differentials for the primary traits. Unintentional selection in the CTL in 1981 was 16.2 kg or .68 SDU for WW and .2 kg or .01 SDU for PG, respectively. Regressions of CSD on year were 4.1 kg or .17 SDU in WWL and 3.2 kg or .14 SDU in PGL. Realized selection differentials were approximately 88% of the potential selection differentials in both lines. Inbreeding coefficients of dam and calves in 1981 were 2.0 and 3.5% in WWL, 2.1 and 3.5% in PGL and 2.9 and 5.8% in CTL.     

Selection for increased weaning or yearling weight in Hereford cattle. I. Measurement of selection applied

Selection was applied from 1964 to 1978 for increased weaning weight (WWL) or yearling weight (YWL) in two Hereford lines with an Angus line maintained as an unselected control line (CL). Each line was maintained with 50 cows and four sires (two sires selected each year and each used for 2 yr). Traits analyzed were birth weight (BW), preweaning daily gain (WDG), weaning weight (WW), weaning conformation grade (WG), weaning condition score (WC), weaning to yearling daily gain (YDG), yearling weight (YW), yearling conformation grade (YG) and yearling condition score (YC). After 15 yr of selection, a total of 3.22 generations of selection had occurred in both WWL and YWL. Average selection differentials in standard measure per generation for WWL, YWL and CL, respectively, were: BW, .44, .51, .0; WDG, .95, .81, .09; WW, .97, .85, .09; WG, .66, .57, .09; WC, .60, .38, -.02; YDG, .30, .79, .38; YW, .80, 1.05, .25; YG, .63, .62, .34 and YC, .45, .64, .24. The proportionate contribution of sire selection (delta S) to the average midparent selection differential per generation (delta M) was 70% in WWL and 76% in YWL. Selection indexes in retrospect were also calculated.     

Bayesian estimation in maternally ancestral animal models for weaning weight of beef cattle

The Bayesian approach was implemented for fitting several maternally ancestral models for weaning weight data of Angus calves. The goal was to evaluate to what extent genetic evaluation models with additive grand maternal effects (G), or with an ancestrally structured covariance matrix for maternal environmental effects (E), or with a sire × year interaction (ISY), or combinations thereof (GE, GSY, ESY, GESY), redistribute the additive variability and reduce the negative magnitude of the additive correlation between direct and maternal effects (r(AoAm)), when compared with the regular maternal animal model (I). All animals with records had known dams and maternal granddams. The sampling scheme induced low autocorrelations among all variables and tended to converge quickly. The signs of the estimates of r(AoAm) were consistently negative for all models fitted. The magnitudes of the estimates of r(AoAm) from models E, G, GE, ESY, and GESY were almost one-third of those from models I and ISY. Inclusion of the sire × year interaction had some effect in reducing the negative magnitude of r(AoAm), but also reduced the size of the estimates of direct (h(0)(2)) and maternal (h(m)(2)) heritabilities. In comparison, models E or G reduced the negative magnitude of r(AoAm) by 0.50 units and produced more favorable estimates of H(0)(2) and h(m)(2) than models I and ISY. The estimate of h(0)(2) from G was similar to the one from I; however, the estimated h(m)(2) was 0.04 units greater, whereas the estimate of r(AoAm) was much less negative (-0.21 vs. -0.71) than the respective estimates from I. The environmental correlation between the weaning weights of dams and their daughters (λ) was estimated to be -0.28 ± 0.03 in E and ESY, and -0.21 ± 0.03 in GE and GESY. Inclusion of the sire × year interaction effect by itself did not have much of an impact in the reduction of the estimated magnitude of r(AoAm). Rank correlations among EBV for direct effects were larger than 0.94 and did not show any appreciable difference among models, whereas the rank correlation among maternal breeding values displayed differences in the ranking between I and the other models. Models E and ESY recovered the largest amount of total additive variability with maternal effects.     

Preweaning breed-of-sire comparisons involving the Senepol breed of cattle

Preweaning data collected at two locations (Kentucky, Louisiana) were utilized to evaluate breed-of-sire comparisons involving the Senepol breed of cattle. For the Kentucky study, calves sired by Senepol bulls were 1.3 kg heavier (P less than .05) at birth than calves sired by Hereford bulls; however, weaning weights were similar for the two sire groups. For the Louisiana study, calves sired by Longhorn bulls were 5.3 kg lighter (P less than .01) at birth, 20 kg lighter (P less than .01) at weaning and had weaning condition scores .5 unit less (P less than .01) than the average of calves sired by Red Poll and Senepol bulls. Also, heifers exposed to Longhorn bulls weaned 23 kg less (P less than .01) calf per heifer exposed than the average of heifers exposed to Red Poll and Senepol bulls. Calves sired by Red Poll bulls were 1.2 kg heavier (P less than .01) at birth and 12 kg heavier (P less than .01) at weaning than those sired by Senepol bulls; however, the Senepol-sired calves received higher (P less than .01) condition scores at weaning. Heifers exposed to Red Poll bulls weaned 20 kg more (P less than .05) calf per heifer exposed than did heifers exposed to Senepol bulls.     

Effects of sire misidentification on estimates of genetic parameters for birth and weaning weights in Hereford cattle

Birth weights (4,155) and weaning weights (3,884) of Line 1 Herefords collected at the Fort Keogh Livestock and Range Research Laboratory in Miles City, MT, between the years of 1935 to 1989 were available. To study the effect of misidentification on estimates of genetic parameters, the sire identification of calf was randomly replaced by the identification of another sire based on the fraction of progeny each sire contributed to a yearly calf crop. Misidentification rates ranged from 5 to 50% with increments of 5%. For each rate of misidentification, 100 replicates were obtained and analyzed with single-trait and two-trait analyses with a restricted maximum likelihood (REML) algorithm. Two different models were used. Both models contained year x sex combinations and ages of dam as fixed effects, calendar birth date as a fixed covariate, and random animal and maternal genetic effects and maternal permanent environment effects. Model 2 also included sire x year combinations as random effects. As the rate of misidentification increased, estimates of the direct-maternal genetic correlation increased for both traits, with both models, for all analyses. With singletrait analyses, estimates of the fraction of variance that were due to sire x year interaction effects increased slightly for birth weight (near zero) and decreased slightly (0.015 to 0.004) for weaning weight as misidentification increased. With two-trait analyses, estimates of fraction of variance that were due to sire x year effects gradually decreased for weaning weight as misidentification increased. With the two-trait analyses, and with both models, as the level of sire misidentification increased, estimates of the genetic correlation between direct effects gradually increased, and estimates of the correlation between maternal effects gradually decreased. Estimates of the direct-maternal genetic correlation were more positive with Model 2 than with Model 1 for all levels of misidentification. Results of this study indicate that misidentification of sires would severely bias estimates of genetic parameters and would reduce genetic gain from selection.     

Sire X environment interactions in beef cattle weaning weight field data

Weaning weight field records, supplied by the American Polled Hereford Association, were used to examine sire X environment interactions. Sire X herd/region and sire X contemporary group/herd interactions were evaluated from a data set containing 19,503 records. Sire X region interaction was evaluated from a data set containing 8,659 records. The genetic correlations of sire progeny performance across contemporary groups/herd were .59 and .37 across herds and contemporary groups/region. The average genetic correlation of sire progeny performance across regions was .64. Heritability of weaning weight was .11 across regions, .17 within region and .28 within herd. Mixed-model sire analyses of Polled Hereford weaning weight field records should include sire X herd/region and sire X contemporary group/herd random effects to reduce the sire X environment effects particular to any herd or contemporary group, and to account for the distribution of sire progeny across herds and contemporary groups in the estimation of prediction error variance. It may be necessary to perform separate sire analyses for some regions to evaluate the breeding values of sires in regions where rank changes are likely to occur.     

Genetic parameters for birth weight,weaning weight and age at first calving in Brown Swiss cattle in Mexico

Heritabilities and genetic correlations between birth weight (n = 13,741), adjusted 240-day weaning weight (WW, n = 8,806) and age at first calving (AFC, n = 3,955) of Brown Swiss cattle in Mexico were estimated. Data from 91 herds located in 19 of 32 states of Mexico from 1982 to 2006 were provided by the Mexican Brown cattle Breeder Association. Components of (co)variance, direct and maternal heritabilities were estimated for birth weight, WW and AFC using bivariate animal models. Direct and maternal heritabilities were 0.21 and 0.05 for birth weight, 0.40 and 0.05 for WW, whereas direct heritability for AFC was 0.08. The correlations between direct and maternal effects for birth weight and WW were −0.49 and −0.64, respectively. The genetic correlations between birth weight–WW and WW–AFC were 0.36 and −0.02, respectively. Under the conditions of this study, selection for increasing birth weight would increase WW, but increasing WW will not change AFC.     

The impact of direct-maternal genetic correlations on international beef cattle evaluations for Limousin weaning weight

In beef cattle maternally influenced traits, estimates of direct-maternal genetic correlations (r_dm) are usually reported to be negative. In international evaluations, r_dm can differ both within countries (r_{dm_WC}) and between countries (r_{dm_BC}). The r_{dm_BC} are difficult to estimate and are assumed to be zero in the current model for international beef cattle evaluations (Interbeef). Our objective was to investigate re-ranking of international estimated breeding values (IEBVs) in international beef cattle evaluations between models that either used estimated values for r_dm or assumed them to be 0. Age-adjusted weaning weights and pedigree data were available for Limousin beef cattle from ten European countries. International EBVs were obtained using a multi-trait animal model with countries modeled as different traits. We compared IEBVs from a model that uses estimated r_{dm_BC} (ranging between −0.14 and +0.14) and r_{dm_WC} (between −0.33 and +0.40) with IEBVs obtained either from the current model that assumes r_{dm_BC} to be 0, or from an alternative model that assumes both r_{dm_BC} and r_{dm_WC} to be 0. Direct and maternal IEBVs were compared across those three scenarios for different groups of animals. The ratio of population accuracies from the linear regression method was used to further investigate the impact of r_dm on international evaluations, for both the whole set of animals in the evaluation and the domestic ones. Ignoring r_{dm_BC}, i.e., replacing estimated values with 0, resulted in no (rank correlations > 0.99) or limited (between 0.98 and 0.99) re-ranking for direct and maternal IEBVs, respectively. Both r_{dm_BC} and r_{dm_WC} had less impact on direct IEBVs than on maternal IEBVs. Re-ranking of maternal IEBVs decreased with increasing reliability. Ignoring r_{dm_BC} resulted in no re-ranking for sires with IEBVs that might be exchanged across countries and limited re-ranking for the top 100 sires. Using estimated r_{dm_BC} values instead of considering them to be 0 resulted in null to limited increases in population accuracy. Ignoring both r_{dm_BC} and r_{dm_WC} resulted in considerable re-ranking of animals’ IEBVs in all groups of animals evaluated. This study showed the limited impact of the current practice of ignoring r_{dm_BC} in international evaluations for Limousin weaning weight, most likely because the estimated r_{dm_BC} was close to 0. We expect that these conclusions can be extended to other traits that have reported r_dm values in the range of r_{dm_WC} values for weaning weight in Limousin.     

Estimation of direct, maternal, and grandmaternal genetic effects for weaning weight in several breeds of beef cattle.

Weaning weights from nine parental breeds and three composites were analyzed to estimate variance due to grandmaternal genetic effects and to compare estimates for variance due to maternal genetic effects from two different models. Number of observations ranged from 794 to 3,465 per population. Number of animals in the pedigree file ranged from 1,244 to 4,326 per population. Two single-trait animal models were used to obtain estimates of covariance components by REML using an average information method. Model 1 included random direct and maternal genetic, permanent maternal environmental, and residual environmental effects as well as fixed sex x year and age of dam effects. Model 2 in addition included random grandmaternal genetic and permanent grandmaternal environmental effects to account for maternal effects of a cow on her daughter's maternal ability. Non-zero estimates of proportion of variance due to grandmaternal effects were obtained for 7 of the 12 populations and ranged from .03 to .06. Direct heritability estimates in these populations were similar with both models. Existence of variance due to grandmaternal effects did not affect the estimates of maternal heritability (m2) or the correlation between direct and maternal genetic effects (r(am)) for Angus and Gelbvieh. For the other five populations, magnitude of estimates increased for both m2 and r(am) when estimates of variance due to grandmaternal effects were not zero. Estimates of the correlation between maternal and grandmaternal genetic effects were large and negative. These results suggest that grand-maternal effects exist in some populations, that when such effects are ignored in analyses maternal heritability may be underestimated, and that the correlation between direct and maternal genetic effects may be biased downward if grandmaternal effects are not included in the model for weaning weight of beef cattle.     

Comparison of models to estimate genetic effects of weaning weight of Angus cattle

Weaning weights from nine sets of Angus field data from three regions of the United States were analyzed. Six animal models were used to compare two approaches to account for an environmental dam-offspring covariance and to investigate the effects of sire x herd-year interaction on the genetic parameters. Model 1 included random direct and maternal genetic, maternal permanent environmental, and residual effects. Age at weaning was a covariate. Other fixed effects were age of dam and a herd-year-management-sex combination. Possible influence of a dam's phenotype on her daughter's maternal ability was modeled by including a regression on maternal phenotype (fm) (Model 3) or by fitting grandmaternal genetic and grandmaternal permanent environmental effects (Model 5). Models 2, 4, and 6 were based on Models 1, 3, and 5, respectively, and additionally included sire x herd-year (SH) interaction effects. With Model 3, estimates of fm ranged from -.003 to .014, and (co)variance estimates were similar to those from Model 1. With Model 5, grandmaternal heritability estimates ranged from .02 to .07. Estimates of maternal heritability and direct-maternal correlation (r(am)) increased compared with Model 1. With models including SH, estimates of the fraction of phenotypic variance due to SH interaction effects were from .02 to .10. Estimates of direct and maternal heritability were smaller and estimates of r(am) were greater than with models without SH interaction effects. Likelihood values showed that SH interaction effects were more important than fm and grandmaternal effects. The comparisons of models suggest that r(am) may be biased downward if SH interaction and(or) grandmaternal effects are not included in models for weaning weight.     

Genotype by region and season interactions on weaning weight in United States Angus cattle

The objective of this study was to determine if weaning weight performance is genetically consistent across different environments in the United States. The American Angus Association provided weight and pedigree data. Weaning weights observed in the Southeast (SoE) and Northwest (NW) were the focus of this study, as these regions are perceived as opposite extremes in climate. The 2 most represented calving seasons in each region were fall and winter in the SoE and winter and spring in the NW. The original data were edited to remove weaning weight records outside of 3 SD from the respective region-season mean, contemporary groups smaller than 20, and single-sire contemporary groups. The final dataset included 884,465 weaning weight records with 64,907 from fall-born calves in the SoE, 74,820 from winter-born calves in the SoE, 346,724 from winter-born calves in the NW and 398,014 from spring-born calves in the NW. Weaning weights of calves born in different region-season classes adjusted to 205 d of age were considered different but genetically correlated traits in a multivariate analysis. The sole fixed effect was weaning contemporary group and random effects included direct, maternal, maternal permanent environment, and a residual. Direct heritability estimates differed little across environments: 0.31 and 0.35 for weight in fall- and winter-born calves in the SoE, and 0.29 and 0.32 for winter- and spring-born calves in NW. Maternal heritability estimates ranged from 0.12 in the NW to 0.16 the SoE. Genetic correlations spanned from 0.69 to 0.93 among direct effects and from 0.65 to 0.95 among maternal effects. All heritability estimates had small (0.01 to 0.04) SE. The most distinct environments appeared to be winter in SoE and spring in NW (correlations of 0.69 and 0.65 for the direct and maternal effects). Different choices of sires for different environments might be justified to achieve the growth performance expected.