Absorption of equations for non-parents for an animal model with maternal effects and genetic groups

Rules for forming the mixed-model equations for the reduced animal model with all relationships and including maternal effects have been set out by Quaas and Pollak. They also have shown how to simplify the mixed-model equations when genetic group effects are included in the model with what has become known as the Q-P transformation. Westell has given rules for calculating the coefficients for the Q-P transformed equations that are associated with the inverse of the numerator relationship matrix and genetic group effects. Those rules can be extended to include maternal effects and genetic groups for maternal as well as direct effects. As with the rules of Quaas and Pollak for the equations for the reduced animal model, a similar set of rules can be obtained for the genetic groups model after the Q-P transformation. The rules are derived easily by examining the algebraic results of absorbing the direct and maternal breeding value equations for non-parents into the parent breeding value, group and fixed effects equations. These rules involve Westell's rules and the inverse elements of the genetic (co)variance matrix for direct and maternal additive genetic effects. The rules make calculation of breeding values for parents for models including direct and maternal genetic group effects nearly as easy as for models without genetic group effects. Back solution for direct and maternal breeding values of non-parents similarly is as simple as when genetic group effects are not in the model.     

Alternative animal models with maternal effects and foster dams

Effects of foster dams can be included in genetic evaluations using animal models with maternal effects in several ways. The alternatives discussed involve minor changes in computing strategies from strategies used with reduced animal models that predict breeding values for direct and maternal effects. The easiest alternative is to assign foster dams to groups by breed and time period and add equations for fixed effects of breed-period. Random and, assumed, independent effects of foster dams can be nested in breed-period groups. If foster dams do not repeat, then those effects can be absorbed into equations for other fixed effects, additive direct breeding value and breed-period effects by slightly modifying least squares contributions to coefficients of those equations. A third alternative for foster dams of the same breed is to add breeding values for foster dams for direct and maternal effects to solution vectors for breeding values. Equations are similar to those without foster dams, except that least squares contributions to coefficient matrix and right-hand sides are to equations for maternal breeding values and nongenetic maternal effects of foster dams rather than biological dams. Relationships and covariance between direct and maternal effects contribute mixed-model coefficients to direct and maternal breeding value equations of biological dams. This alternative basically requires only larger solution vectors for direct and maternal breeding values to accommodate foster dams that might not be included. The fourth alternative includes a vector of maternal breeding values for foster dams of each breed of foster dams and would require using rules of Westell to calculate coefficients due to relationships and fixed maternal genetic groups within each breed of foster dam. These alternatives do not require much additional computational effort compared with full or reduced animal model equations when the transformation to predict breeding values is used with Westell's rules to calculate coefficients due to relationships and genetic group effects due to prior genetic selection.     

Illustration of the maternal animal model used for genetic evaluation of beef cattle

National cattle evaluation programs for weaning weight in most beef breed associations involve implementation of the maternal animal model to predict direct and maternal EPD. With this model, direct breeding values are predicted for all animals with records or pedigree ties to animals with records, or both. Even though maternal genetic value is expressed only in animals that become dams, these effects are transmitted by all parents and inherited from parents by all animals, leading to maternal breeding values being predicted for all animals as well. A small example data set was simulated involving 12 parents, 8 nonparents, and 13 animals with weaning weight records. The pedigree was developed to include paternal and maternal half-sib families, full-sibs, and some inbreeding, similar to field populations of beef cattle. Assembly of the mixed model equations and solutions for the maternal animal model are illustrated explicitly to assist animal breeding students in their understanding of the properties of the maternal animal model and to explicitly implement the model. Model parameters and moments, fixed contemporary group solutions, adjustment of breeding values for merit of mates, interpretation of maternal permanent environmental effect solutions, and alternatives for the assembly of the equations are shown. This example should lead to increased student and producer understanding of genetic improvement programs for weaning weight in beef cattle.     

Estimation of correlation between maternal permanent environmental effects of related dams in beef cattle

Weaning weights from Gelbvieh (GV; n = 82,138) and Limousin (LM; n = 88,639) calves were used to estimate genetic and environmental variance components with models that included different values for the correlation (lambda) between permanent environmental effects of dams and their daughters. Each analysis included fixed discrete effects of contemporary group, sex of calf, age of dam at calving, and month of calving, a fixed continuous effect of age of calf, random direct and maternal additive genetic effects, permanent environmental effects due to dams, and residual effects. The REML procedure was employed with a "grid search," in which the likelihood was computed for a series of values for lambda. For both breeds, models that included a nonzero value for lambda fitted the data significantly better than the model that did not include lambda. The maximum restricted likelihood was obtained for lambda of approximately -0.2 for both breeds. Estimates of residual and direct genetic variances were similar for all values of lambda, including zero; however, estimates of maternal genetic variance and maternal heritability increased slightly, and maternal permanent environmental variance and the proportion of the maternal variance to the total (phenotypic) variance decreased slightly, when the correlated structure for permanent environmental effects was assumed. As the value of lambda became more negative, absolute values of the direct-maternal genetic covariance and direct-maternal correlation estimates were decreased. Pearson and rank correlations for direct genetic, maternal genetic, and maternal environmental effects estimated with and without lambda were very high (>0.99). These results indicated that the linear relationship between maternal permanent environmental effects of dams and their daughters for weaning weight is negative but low in both breeds. Considering this relationship in the operational model did not significantly affect estimated breeding values, and thus, it may not be important in genetic evaluations.     

Influence of population structure on estimates of direct and maternal parameters.

The estimation of (co)variance components for multiple traits with maternal genetic effects was found to be influenced by population structure. Two traits in a closed breeding herd with random mating were simulated over nine generations. Population structures were simulated on the basis of different proportions of dams not having performance records (0, 0.1, 0.5, 0.8 and 0.9): three genetic correlations (-0.5, 0.0 and +0.5) between direct and maternal effects and three genetic correlations (0, 0.3 and 0.8) between two traits. Three ratios of direct to maternal genetic variances, (1:3, 1:1, 3:1), were also considered. Variance components were estimated by restricted maximum likelihood. The proportion of dams without records had an effect on the SE of direct-maternal covariance estimates when the proportion was 0.8 or 0.9 and the true correlation between direct and maternal effects was negative. The ratio of direct to maternal genetic variances influenced the SE of the (co)variance estimates more than the proportion of dams with missing records. The correlation between two traits did not have an effect on the SE of the estimates. The proportion of dams without records and the correlation between direct and maternal effects had the strongest effects on bias of estimates. The largest biases were obtained when the proportion of dams without records was high, the correlation between direct and maternal effects was positive, and the direct variance was greater than the maternal variance, as would be the situation for most growth traits in livestock. Total bias in all parameter estimates for two traits was large in the same situations. Poor population structure can affect both bias and SE of estimates of the direct-maternal genetic correlation, and can explain some of the large negative estimates often obtained.     

The impact of data structure on genetic (co)variance components of early growth in sheep,estimated using an animal model with maternal effects

Several studies have noted high negative correlations between maternal genetic and direct additive effects and their influence on additive and maternal heritability of early growth traits in sheep. Multigeneration data from the Suffolk Sire Reference Scheme (SSRS) were used to investigate the effect of data structure on estimates of direct and maternal (co)variances for lamb 8-wk weight. In all analyses the additive, maternal genetic, maternal environmental, and residual effects were fitted along with the covariance between direct and maternal additive effects. The contributions of particular genetic relationships to the estimates were studied by analyzing subsets of the SSRS data. A further eight subsets were formed having 10% or 50% of the dams with their own records and having one or two, three or four, five or six, and more than six offspring per dam. Analysis of data having only 10% of the dams with their own record and one or two offspring records yielded a high negative correlation (-0.99) between direct and maternal genetic effects. However, the seven other data sets with more records per dam or a higher proportion of dams with their own records produced values of -0.35 to -0.51. Data structure and the number of dams and granddams with records are important determinants of estimated direct and maternal effects in early growth traits.     

Investigating maternal effects on production traits in Duroc pigs using animal and sire models

Variance components for production traits were estimated using different models to evaluate maternal effects. Data analysed were records from the South African pig performance testing scheme on 22 224 pigs from 18 herds, tested between 1990 and 2008. The traits analysed were backfat thickness (BFAT), test period weight gain (TPG), lifetime weight gain (LTG), test period feed conversion ratio (FCR) and age at slaughter (AGES). Data analyses were performed by REML procedures in ASREML, where random effects were successively fitted into animal and sire models to produce different models. The first animal model had one random effect, the direct genetic effects, while the additional random effects were maternal genetic and maternal permanent environmental effects. In the sire model, the random effects fitted were sire and maternal grand sire effects. The best model considered the covariance between direct and maternal genetic effects or between sire and maternal grand sire effects. Fitting maternal genetic effects into the animal model reduced total additive variance, while the total additive variance increased when maternal grand sire effects were fitted into the sire model. The correlations between direct and maternal genetic effects were all negative, indicating antagonism between these effects, hence the need to consider both effects in selection programmes. Direct genetic correlations were higher than other correlations, except for maternal genetic correlations of FCR with TPG, LTG and AGES. There has been direct genetic improvement and almost constant maternal ability in production traits as shown by trends for estimated (EBVs) and maternal breeding values (MBVs), while phenotypic trends were similar to those for EBVs. These results suggest that maternal genetic effects should be included in selection programmes for these production traits. Therefore, the animal–maternal model may be the most appropriate model to use when estimating genetic parameters for production traits in this population.     

Direct and maternal (co)variance components, genetic parameters, and annual trends for growth traits of Makooei sheep in Iran

Genetic parameters and genetic trends for birth weight (BW), weaning weight (WW), 6-month weight (6MW), and yearling weight (YW) traits were estimated by using records of 5,634 Makooei lambs, descendants of 289 sires and 1,726 dams, born between 1996 and 2009 at the Makooei sheep breeding station, West Azerbaijan, Iran. The (co)variance components were estimated with different animal models using a restricted maximum likelihood procedure and the most appropriate model for each trait was determined by Akaike’s Information Criterion. Breeding values of animals were predicted with best linear unbiased prediction methodology under multi-trait animal models and genetic trends were estimated by regression mean breeding values on birth year. The most appropriate model for BW was a model including direct and maternal genetic effects, regardless of their covariance. The model for WW and 6MW included direct additive genetic effects. The model for YW included direct genetic effects only. Direct heritabilities based on the best model were estimated 0.15?±?0.04, 0.16?±?0.03, 0.21?±?0.04, and 0.22?±?0.06 for BW, WW, 6MW, and YW, respectively, and maternal heritability obtained 0.08?±?0.02 for BW. Genetic correlations among the traits were positive and varied from 0.28 for BW–YW to 0.66 for BW–WW and phenotypic correlations were generally lower than the genetic correlations. Genetic trends were 8.1?±?2, 67.4?±?5, 38.7?±?4, and 47.6?±?6 g per year for BW, WW, 6MW, and YW, respectively.     

Genetic parameters for direct and maternal calving ease in Walloon dairy cattle based on linear and threshold models

Calving ease scores from Holstein dairy cattle in the Walloon Region of Belgium were analysed using univariate linear and threshold animal models. Variance components and derived genetic parameters were estimated from a data set including 33 155 calving records. Included in the models were season, herd and sex of calf × age of dam classes × group of calvings interaction as fixed effects, herd × year of calving, maternal permanent environment and animal direct and maternal additive genetic as random effects. Models were fitted with the genetic correlation between direct and maternal additive genetic effects either estimated or constrained to zero. Direct heritability for calving ease was approximately 8% with linear models and approximately 12% with threshold models. Maternal heritabilities were approximately 2 and 4%, respectively. Genetic correlation between direct and maternal additive effects was found to be not significantly different from zero. Models were compared in terms of goodness of fit and predictive ability. Criteria of comparison such as mean squared error, correlation between observed and predicted calving ease scores as well as between estimated breeding values were estimated from 85 118 calving records. The results provided few differences between linear and threshold models even though correlations between estimated breeding values from subsets of data for sires with progeny from linear model were 17 and 23% greater for direct and maternal genetic effects, respectively, than from threshold model. For the purpose of genetic evaluation for calving ease in Walloon Holstein dairy cattle, the linear animal model without covariance between direct and maternal additive effects was found to be the best choice.     

Estimation of genetic effects in the presence of multicollinearity in multibreed beef cattle evaluation

Breed additive, dominance, and epistatic loss effects are of concern in the genetic evaluation of a multibreed population. Multiple regression equations used for fitting these effects may show a high degree of multicollinearity among predictor variables. Typically, when strong linear relationships exist, the regression coefficients have large SE and are sensitive to changes in the data file and to the addition or deletion of variables in the model. Generalized ridge regression methods were applied to obtain stable estimates of direct and maternal breed additive, dominance, and epistatic loss effects in the presence of multicollinearity among predictor variables. Preweaning weight gains of beef calves in Ontario, Canada, from 1986 to 1999 were analyzed. The genetic model included fixed direct and maternal breed additive, dominance, and epistatic loss effects, fixed environmental effects of age of the calf, contemporary group, and age of the dam x sex of the calf, random additive direct and maternal genetic effects, and random maternal permanent environment effect. The degree and the nature of the multicollinearity were identified and ridge regression methods were used as an alternative to ordinary least squares (LS). Ridge parameters were obtained using two different objective methods: 1) generalized ridge estimator of Hoerl and Kennard (R1); and 2) bootstrap in combination with cross-validation (R2). Both ridge regression methods outperformed the LS estimator with respect to mean squared error of predictions (MSEP) and variance inflation factors (VIF) computed over 100 bootstrap samples. The MSEP of R1 and R2 were similar, and they were 3% less than the MSEP of LS. The average VIF of LS, R1, and R2 were equal to 26.81, 6.10, and 4.18, respectively. Ridge regression methods were particularly effective in decreasing the multicollinearity involving predictor variables of breed additive effects. Because of a high degree of confounding between estimates of maternal dominance and direct epistatic loss effects, it was not possible to compare the relative importance of these effects with a high level of confidence. The inclusion of epistatic loss effects in the additive-dominance model did not cause noticeable reranking of sires, dams, and calves based on across-breed EBV. More precise estimates of breed effects as a result of this study may result in more stable across-breed estimated breeding values over the years.     

Evaluation of animal models to explore the influence of maternal genetic and maternal permanent environment effect on reproductive performance of Jersey crossbred cattle

The present study was conducted on 1,002 reproductive records of 430 Jersey crossbred cattle, descended from 57 sires and 198 dams, maintained at the Eastern Regional Station of ICAR-National Dairy Research Institute, Kalyani, Nadia, West Bengal, India to investigate the influence of direct genetic, maternal genetic and maternal permanent environmental effect on three most important reproductive traits viz., number of service per conception (NSPC), days open (DO) and calving interval (CI) of Jersey crossbred cattle. Six single-trait animal models (including or excluding maternal genetic or permanent environmental effects) were fitted to analyse these traits, and the best model was chosen after testing the significant increase in the log-likelihood values when additional parameters were added in the model. Direct heritability estimates for NSPC, DO and CI from the best model were 0.10, 0.14 and 0.20, respectively. The maternal permanent environmental (c²) effects on reproductive traits accounted for almost negligible fraction of the total phenotypic variance in this study. The maternal genetic effects (m²) also contributed very little (0%–3%) to the total phenotypic variance except for CI where it was important and accounted for 20% of phenotypic variance. A significantly large negative genetic correlation was observed between direct and maternal genetic effects for all traits, suggesting the presence of antagonistic relationship between dam's direct additive component and daughter's additive genetic component. Results suggest that both direct and maternal effects were important only for CI but not for other traits. Therefore, both direct additive effects and maternal genetic effect need to be considered for improving this trait by selection.     

Variance components and breeding values for growth traits from different statistical models.

Estimates of genetic parameters resulting from various analytical models for birth weight (BWT, n = 4,155), 205-d weight (WWT, n = 3,884), and 365-d weight (YWT, n = 3,476) were compared. Data consisted of records for Line 1 Hereford cattle selected for postweaning growth from 1934 to 1989 at ARS-USDA, Miles City, MT. Twelve models were compared. Model 1 included fixed effects of year, sex, age of dam; covariates for birth day and inbreeding coefficients of animal and of dam; and random animal genetic and residual effects. Model 2 was the same as Model 1 but ignored inbreeding coefficients. Model 3 was the same as Model 1 and included random maternal genetic effects with covariance between direct and maternal genetic effects, and maternal permanent environmental effects. Model 4 was the same as Model 3 but ignored inbreeding. Model 5 was the same as Model 1 but with a random sire effect instead of animal genetic effect. Model 6 was the same as Model 5 but ignored inbreeding. Model 7 was a sire model that considered relationships among males. Model 8 was a sire model, assuming sires to be unrelated, but with dam effects as uncorrelated random effects to account for maternal effects. Model 9 was a sire and dam model but with relationships to account for direct and maternal genetic effects; dams also were included as uncorrelated random effects to account for maternal permanent environmental effects. Model 10 was a sire model with maternal grandsire and dam effects all as uncorrelated random effects. Model 11 was a sire and maternal grandsire model, with dams as uncorrelated random effects but with sires and maternal grandsires assumed to be related using male relationships. Model 12 was the same as Model 11 but with all pedigree relationships from the full animal model for sires and maternal grandsires. Rankings on predictions of breeding values were the same regardless of whether inbreeding coefficients for animal and dam were included in the models. Heritability estimates were similar regardless of whether inbreeding effects were in the model. Models 3 and 9 best fit the data for estimation of variances and covariances for direct, maternal genetic, and permanent environmental effects. Other models resulted in changes in ranking for predicted breeding values and for estimates of direct and maternal heritability. Heritability estimates of direct effects were smallest with sire and sire-maternal grandsire models.     

Breeding value prediction with maternal genetic groups

For models with only additive direct genetic effects, the rules of Westell combined with the Q-P transformation can be used to calculate the coefficients of mixed-model equations corresponding to the inverse elements of the numerator relationship matrix and group effects that are used to account for selection on ancestors that do not have records. Groups generally can be assigned on the basis of most recent ancestors without records. When maternal effects are in the model, most recent female ancestors without records contribute maternal effects to their progeny. If the vectors for additive direct and maternal effects do not include the same animals, numerator relationship matrices for direct and maternal effects and between direct and maternal effects are different. Even if they are the same, the Q-P transformation and Westell's rules do not lead to simplification for calculation of the coefficient matrix unless group assignment is the same for direct and maternal effects. This result can be achieved by including each female ancestor with offspring having records in both vectors and by assigning both of her parents to the same group she would have been assigned for a model including only direct effects. This strategy is equivalent to assigning group effects similarly for both direct and maternal effects and allows making use of the computational efficiency available from the Q-P transformation and Westell's rules, which are similar to Henderson's rules for calculating the inverse of the numerator relationship matrix.     

Nuclear, cytoplasmic, and environmental effects on growth, fat, and muscle traits in suffolk lambs from a sire referencing scheme

Maternal effects are an important source of variation in early growth and body traits in sheep but are often excluded from genetic analyses. Maternal additive genetic, maternal environmental, and cytoplasmic effects were investigated in a large Suffolk breeding scheme using a range of models involving different combinations of these effects with the direct additive genetic effect. Weights at 8 wk of age and at scanning (mean age 146 d) and ultrasonically measured muscle and fat depth were analyzed using an animal model on 55,683 (8-wk weight) and 28,947 (scanning traits) lamb records. Simple additive models always overestimated the heritability of all traits when compared to more complex models. The successive inclusion of maternal environmental, maternal genetic, and the covariance between direct and maternal additive effects in the model significantly improved the fit for almost all models and all traits, as indicated by a likelihood ratio test. Under the full model, the heritability of both weight traits was low (0.14 and 0.20 for 8-wk and scanning weight, respectively). The maternal additive and maternal environmental effects, as a proportion of the phenotypic variance, were similar (0.10 and 0.08 for 8-wk weight and 0.07 and 0.06 for scanning weight). The two scanning traits had higher heritabilities (0.29 and 0.27 for muscle depth and fat depth, respectively) with low levels of maternal genetic and maternal environmental variance. No evidence was found of a cytoplasmic effect on any of the traits studied under the full model. Breeding schemes for early growth and body traits in sheep should account for maternal effects in their genetic evaluations in order to improve their accuracy. The exact model to use will depend on the trait and individual circumstances of the scheme.     

Variance and covariance estimates for weaning weight of Senepol cattle.

Variance and covariance components were estimated for weaning weight from Senepol field data for use in the reduced animal model for a maternally influenced trait. The 4,634 weaning records were used to evaluate 113 sires and 1,406 dams on the island of St. Croix. Estimates of direct additive genetic variance (sigma 2A), maternal additive genetic variance (sigma 2M), covariance between direct and maternal additive genetic effects (sigma AM), permanent maternal environmental variance (sigma 2PE), and residual variance (sigma 2 epsilon) were calculated by equating variances estimated from a sire-dam model and a sire-maternal grandsire model, with and without the inverse of the numerator relationship matrix (A-1), to their expectations. Estimates were sigma 2A, 139.05 and 138.14 kg2; sigma 2M, 307.04 and 288.90 kg2; sigma AM, -117.57 and -103.76 kg2; sigma 2PE, -258.35 and -243.40 kg2; and sigma 2 epsilon, 588.18 and 577.72 kg2 with and without A-1, respectively. Heritability estimates for direct additive (h2A) were .211 and .210 with and without A-1, respectively. Heritability estimates for maternal additive (h2M) were .47 and .44 with and without A-1, respectively. Correlations between direct and maternal (IAM) effects were -.57 and -.52 with and without A-1, respectively.     

Choice of statistical model for estimating genetic parameters using restricted maximum likelihood in swine

Summary Restricted maximum likelihood (REML) was used to determine the choice of statistical model, additive genetic maternal and common litter effects and consequences of ignoring these effects on estimates of variance–covariance components under random and phenotypic selection in swine using computer simulation. Two closed herds of different size and two traits, (i) pre‐weaning average daily gain and (ii) litter size at birth, were considered. Three levels of additive direct and maternal genetic correlations (r_dm) were assumed to each trait. Four mixed models (denoted as GRM1 through GRM4) were used to generate data sets. Model GRM1 included only additive direct genetic effects, GRM2 included only additive direct genetic and common litter effects, GRM3 included only additive direct and maternal genetic effects and GRM4 included all the random effects. Four mixed animal models (defined as EPM1 through EPM4) were defined for estimating genetic parameters similar to GRM. Data from each GRM were fitted with EPM1 through EPM4. The largest biased estimates of additive genetic variance were obtained when EPM1 was fitted to data generated assuming the presence of either additive maternal genetic, common litter effects or a combination thereof. The bias of estimated additive direct genetic variance (VA_d) increased and those of recidual variance (VE) decreased with an increase in level of r_dm when GRM3 was used. EPM1, EPM2 and EPM3 resulted in biased estimation of the direct genetic variances. EPM4 was the most accurate in each GRM. Phenotypic selection substantially increased bias of estimated additive direct genetic effect and its mean square error in trait 1, but decreased those in trait 2 when ignored in the statistical model. For trait 2, estimates under phenotypic selection were more biased than those under random selection. It was concluded that statistical models for estimating variance components should include all random effects considered to avoid bias.     

Additive, dominance, and epistatic loss effects on preweaning weight gain of crossbred beef cattle from different Bos taurus breeds

(Co)variance components, direct and maternal breed additive, dominance, and epistatic loss effects on preweaning weight gain of beef cattle were estimated. Data were from 478,466 animals in Ontario, Canada, from 1986 to 1999, including records of both purebred and crossbred animals from Angus, Blonde d'Aquitaine, Charolais, Gelbvieh, Hereford, Limousin, Maine-Anjou, Salers, Shorthorn, and Simmental breeds. The genetic model included fixed direct and maternal breed additive, dominance, and epistatic loss effects, fixed environmental effects of age of the calf, contemporary group, and age of the dam x sex of the calf, random additive direct and maternal genetic effects, and random maternal permanent environment effects. Estimates of direct and maternal additive genetic, maternal permanent environmental and residual variances, expressed as proportions of the phenotypic variance, were 0.32, 0.20, 0.12, and 0.52, respectively. Correlation between direct and maternal additive genetic effects was -0.63. Breed ranking was similar to previous studies, but estimates showed large SE. The favorable effects of direct and maternal dominance (P < 0.05) on preweaning gain were equivalent to 1.3 and 2.3% of the phenotypic mean of purebred calves, respectively. The same features for direct and maternal epistatic loss effects were -2.2% (P < 0.05) and -0.1% (P > 0.05). The large SE of breed effects were likely due to multicollinearity among predictor variables and deficiencies in the dataset to separate direct and maternal effects and may result in a less reliable ranking of the animals for across breed comparisons. Further research to identify the causes of the instability of estimates of breed additive, dominance, and epistatic loss genetic effects, and application of alternative statistical methods is recommended.     

Estimation of Genetic Parameters and Variance Components for Growth Traits of Nguni Cattle in Limpopo Province,South Africa

Estimates of (co)variance and genetic parameters of birth, weaning (205 days) and yearling (365 days) weight were obtained using single-trait animal models. The data were analysed by restricted maximum likelihood, fitting an animal model that included direct and maternal genetic and permanent environmental effects. The data included records collected between 1976 and 2001. The pedigree information extended as far back as early 1960s. The heritabilities for direct effects of birth, weaning and yearling weights were 0.36, 0.29 and 0.25, respectively. Heritability estimates for maternal effects were 0.13, 0.16 and 0.15 for birth, weaning and yearling weights, respectively. The correlations between direct and maternal additive genetic effects were negative for all traits analysed. The results indicate that both direct and maternal effects should be included in a selection programme for all the traits analysed.     

Investigating pig survival in different production phases using genomic models

Pig survival is an economically important trait with relevant social welfare implications, thus standing out as an important selection criterion for the current pig farming system. We aimed to estimate (co)variance components for survival in different production phases in a crossbred pig population as well as to investigate the benefit of including genomic information through single-step genomic best linear unbiased prediction (ssGBLUP) on the prediction accuracy of survival traits compared with results from traditional BLUP. Individual survival records on, at most, 64,894 crossbred piglets were evaluated under two multi-trait threshold models. The first model included farrowing, lactation, and combined postweaning survival, whereas the second model included nursery and finishing survival. Direct and maternal breeding values were estimated using BLUP and ssGBLUP methods. Furthermore, prediction accuracy, bias, and dispersion were accessed using the linear regression validation method. Direct heritability estimates for survival in all studied phases were low (from 0.02 to 0.08). Survival in preweaning phases (farrowing and lactation) was controlled by the dam and piglet additive genetic effects, although the maternal side was more important. Postweaning phases (nursery, finishing, and the combination of both) showed the same or higher direct heritabilities compared with preweaning phases. The genetic correlations between survival traits within preweaning and postweaning phases were favorable and strong, but correlations between preweaning and postweaning phases were moderate. The prediction accuracy of survival traits was low, although it increased by including genomic information through ssGBLUP compared with the prediction accuracy from BLUP. Direct and maternal breeding values were similarly accurate with BLUP, but direct breeding values benefited more from genomic information. Overall, a slight increase in bias was observed when genomic information was included, whereas dispersion of breeding values was greatly reduced. Combined postweaning survival presented higher direct heritability than in the preweaning phases and the highest prediction accuracy among all evaluated production phases, therefore standing out as a candidate trait for improving survival. Survival is a complex trait with low heritability; however, important genetic gains can still be obtained, especially under a genomic prediction framework.     

Comparison of models for estimation of genetic parameters for mature weight of Hereford cattle

Genetic parameters of mature weight are needed for effective selection and genetic evaluation. Data for estimating these parameters were collected from 1963 to 1985 and consisted of 32,018 mature weight records of 4,175 Hereford cows that were in one control and three selection lines that had been selected for weaning weight, for yearling weight, or for an index combining yearling weight and muscle score for 22 yr. Several models and subsets of the data were considered. The mature weight records consisted of a maximum of three seasonal weights taken each year, at brand clipping (February and March), before breeding (May and June), and at palpation (August and September). Heritability estimates were high (0.49 to 0.86) for all models considered, which suggests that selection to change mature weight could be effective. The model that best fit the data included maternal genetic and maternal permanent environmental effects in addition to direct genetic and direct permanent environmental effects. Estimates of direct heritability with this model ranged from 0.53 to 0.79, estimates of maternal heritability ranged from 0.09 to 0.21, and estimates of the genetic correlation between direct and maternal effects ranged from -0.16 to -0.67 for subsets of the data based on time of year that mature weight was measured. For the same subsets, estimates of the proportions of variance due to direct permanent environment and maternal permanent environment ranged from 0.00 to 0.09 and 0.00 to 0.06, respectively. Using a similar model that combined all records and included an added fixed effect of season of measurement of mature weight, direct heritability, maternal heritability, genetic correlation between direct and maternal effects, proportion of variance due to direct permanent environmental effects, and proportion of variance due to maternal permanent environmental effects were estimated to be 0.69, 0.13, -0.65, 0.00, and 0.04, respectively. Mature weight is a highly heritable trait that could be included in selection programs and maternal effects should not be ignored when analyzing mature weight data.