Genetic variation in heterogeneous environments.

A model of population structure in heterogeneous environments is described and conditions sufficient for maintaining a polymorphism are derived. The absolute fitness of any genotype is regarded as a function of location in the niche space and the population density at that location. Two modes of habitat selection are examined: (1) organisms are distributed uniformly over the environment; and (2) each organism selects to occupy that habitat in which it is most fit ("optimal habitant selection"). Sufficient conditions for maintenance of genetic polymorphisms are derived for both models. In populations which do not practice habitat selection heterozygote superiority averaged over the environment is sufficient to guarantee the existence of polymorphisms. Comparable conditions for populations which practice optimal habitat selction are much less restrictive. If the heterozygotes are superior to one homozygote in any one part of the niche and to the other homozygote in any other part of the niche then a polymorphism will be defined. A positive correlation between genetic and environmental variation follows from the model with habitat selection, but not from the other. The adaptive significance of polymorphisms thus depends on how animals behave.