The consequences of gene flow for local adaptation and differentiation: a two-locus two-deme model (Q2447547)

The purpose of this paper is to improve the understanding of how genetic architecture (recombination and locus effects) as well as the pattern and amount of migration determine polymorphism, local adaptation, and differentiation in a subdivided population inhabiting a heterogeneous environment. Two linked, diallelic loci and the migration between two demes are considered. If the alleles are beneficial in only one environment and detrimental in the other, local adaptation of subpopulations, and differentiation between them can be obtained only if a (multilocus) polymorphism is maintained. Therefore, the authors prove the existence and stability of polymorphic equilibrium and the dependence of the equilibrium configurations on the model parameters (migration rates, selection coefficients, recombination rate). Three evolutionarily stable states occur: (i) existence of a single specialist optimally adapted to one deme and poorly to the other, (ii) existence of a single generalist type which has higher average fitness in the whole population than than any of the specialists, and (iii) existence of a set of specialists each adapted to its deme, i.e., coexistence in a polymorphism then local adaptation and differentiation occur only in case (iii). The analytical results presented complement the numerical findings by \textit{S. Yeaman} and \textit{M. C. Whitlock} [``The genetic architecture of adaptation under migration-selection balance'', Evolution 65, No. 7, 1897--1911 (2011; \url{doi:10.1111/j.1558-5646.2011.01269.x})] for a multilocus quantitative-genetic model with clusters of locally adaptive mutations, or concentrated genetic architectures, build up in spatially structured populations with opposing selection pressures in two demes. Because tighter linkage is required for invasion under increasingly asymmetric migration rates, more concentrated architectures and a greater advantage for recombination-reducing mechanisms (such as chromosome inversions) should be expected for highly asymmetric migration. In finite populations, invasion of new mutants occurs only with a certain probability, and the genetic drift may erase polymorphism. In the absence of epistasis and under the present form of balancing selection, reduced recombination between selected loci is favored, except when migration rates are sufficiently symmetric and high. Because in the present model, polymorphism at the selected loci is maintained by balancing selection, the effective migration rate may be greatly reduced compared with the actual migration rate. Therefore, strong barriers against gene flow may build up at such neutral sites and enhance (neutral) differentiation.