MAXIMUM-LIKELIHOOD APPROACHES APPLIED TO QUANTITATIVE GENETICS OF NATURAL POPULATIONS.

Growing interest in adaptive evolution in natural populations has spurred efforts to infer genetic components of variance and covariance of quantitative characters. Here, I review difficulties inherent in the usual least-squares methods of estimation. A useful alternative approach is that of maximum likelihood (ML). Its particular advantage over least squares is that estimation and testing procedures are well defined, regardless of the design of the data. A modified version of ML, REML, eliminates the bias of ML estimates of variance components. Expressions for the expected bias and variance of estimates obtained from balanced, fully hierarchical designs are presented for ML and REML. Analyses of data simulated from balanced, hierarchical designs reveal differences in the properties of ML, REML, and F-ratio tests of significance. A second simulation study compares properties of REML estimates obtained from a balanced, fully hierarchical design (within-generation analysis) with those from a sampling design including phenotypic data on parents and multiple progeny. It also illustrates the effects of imposing nonnegativity constraints on the estimates. Finally, it reveals that predictions of the behavior of significance tests based on asymptotic theory are not accurate when sample size is small and that constraining the estimates seriously affects properties of the tests. Because of their great flexibility, likelihood methods can serve as a useful tool for estimation of quantitative-genetic parameters in natural populations. Difficulties involved in hypothesis testing remain to be solved.