Abstract

We consider a population subdivided into two demes connected by migration in which selection acts in opposite direction. We explore the effects of recombination and migration on the maintenance of multilocus polymorphism, on local adaptation, and on differentiation by employing a deterministic model with genic selection on two linked diallelic loci (i.e., no dominance or epistasis). For the following cases, we characterize explicitly the possible equilibrium configurations: weak, strong, highly asymmetric, and super-symmetric migration, no or weak recombination, and independent or strongly recombining loci. For independent loci (linkage equilibrium) and for completely linked loci, we derive the possible bifurcation patterns as functions of the total migration rate, assuming all other parameters are fixed but arbitrary. For these and other cases, we determine analytically the maximum migration rate below which a stable fully polymorphic equilibrium exists. In this case, differentiation and local adaptation are maintained. Their degree is quantified by a new multilocus version of F_mathrm{ST} and by the migration load, respectively. In addition, we investigate the invasion conditions of locally beneficial mutants and show that linkage to a locus that is already in migration-selection balance facilitates invasion. Hence, loci of much smaller effect can invade than predicted by one-locus theory if linkage is sufficiently tight. We study how this minimum amount of linkage admitting invasion depends on the migration pattern. This suggests the emergence of clusters of locally beneficial mutations, which may form ‘genomic islands of divergence’. Finally, the influence of linkage and two-way migration on the effective migration rate at a linked neutral locus is explored. Numerical work complements our analytical results.

Highlights

  • Migration in a geographically structured population may have opposing effects on the genetic composition of that population and, on its evolutionary potential

  • What are the consequences of the genetic architecture, such as linkage between loci, relative magnitude of locus effects, or epistasis, on the degree of local adaptation and of differentiation achieved for a given amount of gene flow? What are the consequences for genetic variation at linked neutral sites? What genetic architectures can be expected to evolve under various forms of spatially heterogeneous selection?

  • Symbols give the exact numerical value of meff = −λN. The purpose of this investigation was to improve our understanding of how genetic architecture, in particular 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

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Summary

Introduction

Migration in a geographically structured population may have opposing effects on the genetic composition of that population and, on its evolutionary potential. The focus of this work was on the properties of the evolutionary dynamics and the conditions for the maintenance of multilocus polymorphism in limiting or special cases, such as weak or strong migration (Bürger 2009a,b), or in the Levene model (Nagylaki 2009; Bürger 2009c, 2010; Barton 2010; Chasnov 2012). Using a continent-island-model framework, Bürger and Akerman (2011) and Bank et al (2012) analyzed the effects of gene flow on local adaptation, differentiation, the emergence of Dobzhansky-Muller incompatibilities, and the maintenance of polymorphism at two linked diallelic loci. They obtained analytical characterizations of the possible equilibrium configurations and bifurcation patterns for wide ranges of parameter combinations.

The model
Equilibria and their stability
Existence of boundary equilibria
Stability of monomorphic equilibria
Stability of single-locus polymorphisms
Weak migration
Linkage equilibrium
Strong recombination: quasi-linkage equilibrium
No recombination
Highly asymmetric migration
3.10 The super-symmetric case
3.11 General case
Bifurcation patterns and maintenance of polymorphism
Important quantities
Admissibility of SLPs
Strong migration
The following diagrams occur for an open set of parameters
Weak recombination
4.10 Highly asymmetric migration
4.11 Maintenance of polymorphism
Migration load and local adaptation
FST and differentiation
Invasion of a locally beneficial mutant
The effective migration rate at a linked neutral site
Discussion
The super-symmetric case
Findings
Important quantities and relations
Full Text
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