Copula miss-specification in REML multivariate genetic animal model estimation

Abstract

Abstract Background In animal genetics, linear mixed models are used to deal with genetic and environmental effects. The variance and covariance terms of these models are usually estimated by restricted maximum likelihood (REML), which provides unbiased estimators. A strong hypothesis of REML estimation is the multi-normality of the response variables. However, in practice, even if the marginal distributions of each phenotype are normal, the multi-normality assumption may be violated by non-normality of the cross-sectional dependence structure, that is to say when the copula of the multivariate distribution is not Gaussian. This study uses simulations to evaluate the impact of copula miss-specification in a bivariate animal model on REML estimations of variance components. Result Bivariate phenotypes were simulated for populations undergoing selection, considering different copulas for the dependence structure between the error components. Two multi-trait situations were considered: two phenotypes were measured on the selection candidates, or only one phenotype was measured on the selection candidates. Three generations with random selection and five generations with truncation selection based on estimated breeding values were simulated. When selection was performed at random, no significant differences were observed between the REML estimations of variance components and the true parameters even for the non-Gaussian distributions. For the truncation selections, when two phenotypes were measured on candidates, biases were systematically observed in the variance components for high residual dependence in the case of non-Gaussian distributions, especially in the case of a heavy-tailed or asymmetric distribution when the two traits were measured. Conversely, when only one phenotype was measured on candidates, no difference was observed between the Gaussian and non-Gaussian distributions in REML estimations. Conclusions This study confirms that REML can be used by geneticists to evaluate breeding values in the multivariate case even if the multivariate phenotypes deviate from normality in the situation of random selection or if one trait is not measured for the candidate under selection. Nevertheless, when the two traits are measured, the violation of the normality assumption may lead to non-negligible biases in the REML estimations of the variance-covariance components.