Gavrilets, S., and Hastings, A. 1994. ``Dynamics of genetic variability in two-locus models of stabilizing selection.'' Genetics 138: 519-532.
ABSTRACT
We study a two locus model, with additive contributions to the phenotype,
to explore the dynamics of different phenotypic characteristics under
stabilizing selection and recombination. We demonstrate that the interaction of selection and recombination results in constraints on the mode of phenotypic
evolution. Let $V_{g}$ be the genic variance of the trait and $C_{L}$ be the contribution of linkage disequilibrium to the genotypic variance. We demonstrate that, independent of the initial conditions, the dynamics of the
system on the plane $(V_{g},C_{L})$ are typically characterized by a
quick approach to a straight line with slow evolution along this line
afterwards. We analyze how the mode and the rate of phenotypic
evolution depend on the strength of selection relative to
recombination, on the form of fitness function, and the difference in
allelic effect. We argue that if selection is not extremely weak
relative to recombination, linkage disequilibrium generated by
stabilizing selection influences the dynamics significantly. We
demonstrate that under these conditions, which are plausible in nature and certainly the case in
artificial stabilizing selection experiments, the model can have
a polymorphic equilibrium with positive linkage disequilibrium that is stable simultaneously with monomorphic equilibria.