Gavrilets, S., and G. de Jong. 1993. ``Pleiotropic models of polygenic variation, stabilizing selection, and epistasis.'' Genetics 134: 609-625.
ABSTRACT
We show that in polymorphic populations many polygenic traits
pleiotropically related to fitness are expected to be under apparent
"stabilizing selection" independently of the real selection acting on the
population. This occurs, for example, if the genetic system is at a stable
polymorphic equilibrium determined by selection and the nonadditive
contributions of the loci to the trait value either are absent, or are
random and independent of those to fitness. Stabilizing selection is also
observed if the polygenic system is at an equilibrium determined by a
balance between selection and mutation (or migration) when both additive
and nonadditive contributions of the loci to the trait value are random and
independent of those to fitness. We also compare different viability models
that can maintain genetic variability at many loci with respect to their
ability to account for the strong stabilizing selection on an additive
trait. Let Vm be the genetic variance supplied by mutation (or migration)
each generation, Vg be the genotypic variance maintained in the population,
and n be the number of the loci influencing fitness. We demonstrate that in
mutation (migration)-selection balance models the strength of apparent
stabilizing selection is order Vm/Vg. In the overdominant model and in the
symmetric viability model the strength of apparent stabilizing selection is
approximately 1/(2n) that of total selection on the whole phenotype. We
show that a selection system that involves pairwise additive by additive
epistasis in maintaining variability can lead to a lower genetic load and
genetic variance in fitness (approximately 1/(2n) times) than an equivalent
selection system that involves overdominance. We show that, in the
epistatic model, the apparent stabilizing selection on an additive trait
can be as strong as the total selection on the whole phenotype.