Gillespie J H
Department of Genetics, University of California, Davis 95616.
Theor Popul Biol. 1990 Feb;37(1):91-109. doi: 10.1016/0040-5809(90)90028-t.
This paper provides a theoretical description of the distribution of the number of mutations that separate alleles that are held in a population by balancing selection. Two models of nucleotide site epistasis are described: parity models and additive site models. Parity models are shown to result in a more uniform distribution of mutations across alleles than the neutral model, while additive sites models show a more extreme distribution. The analytic approach uses strong-selection, weak-mutation approximations to constant-fitness and random-environment diffusion models.