Bürger Reinhard, Gimelfarb Alexander
Department of Mathematics, University of Vienna, A-1090 Vienna, Austria.
Genetics. 2004 Jul;167(3):1425-43. doi: 10.1534/genetics.103.018986.
The equilibrium properties of an additive multilocus model of a quantitative trait under frequency- and density-dependent selection are investigated. Two opposing evolutionary forces are assumed to act: (i) stabilizing selection on the trait, which favors genotypes with an intermediate phenotype, and (ii) intraspecific competition mediated by that trait, which favors genotypes whose effect on the trait deviates most from that of the prevailing genotypes. Accordingly, fitnesses of genotypes have a frequency-independent component describing stabilizing selection and a frequency- and density-dependent component modeling competition. We study how the equilibrium structure, in particular, number, degree of polymorphism, and genetic variance of stable equilibria, is affected by the strength of frequency dependence, and what role the number of loci, the amount of recombination, and the demographic parameters play. To this end, we employ a statistical and numerical approach, complemented by analytical results, and explore how the equilibrium properties averaged over a large number of genetic systems with a given number of loci and average amount of recombination depend on the ecological and demographic parameters. We identify two parameter regions with a transitory region in between, in which the equilibrium properties of genetic systems are distinctively different. These regions depend on the strength of frequency dependence relative to pure stabilizing selection and on the demographic parameters, but not on the number of loci or the amount of recombination. We further study the shape of the fitness function observed at equilibrium and the extent to which the dynamics in this model are adaptive, and we present examples of equilibrium distributions of genotypic values under strong frequency dependence. Consequences for the maintenance of genetic variation, the detection of disruptive selection, and models of sympatric speciation are discussed.
研究了在频率和密度依赖选择下数量性状的加性多基因座模型的平衡性质。假设存在两种相反的进化力量起作用:(i)对性状的稳定选择,它有利于具有中间表型的基因型;(ii)由该性状介导的种内竞争,它有利于其对性状的影响与主流基因型偏差最大的基因型。因此,基因型的适合度有一个描述稳定选择的频率独立成分和一个模拟竞争的频率和密度依赖成分。我们研究稳定平衡的平衡结构,特别是数量、多态性程度和遗传方差,如何受到频率依赖强度的影响,以及基因座数量、重组量和种群统计学参数起什么作用。为此,我们采用一种统计和数值方法,并辅以分析结果,探讨在具有给定基因座数量和平均重组量的大量遗传系统上平均的平衡性质如何依赖于生态和种群统计学参数。我们确定了两个参数区域,中间有一个过渡区域,其中遗传系统的平衡性质明显不同。这些区域取决于相对于纯稳定选择的频率依赖强度和种群统计学参数,但不取决于基因座数量或重组量。我们进一步研究在平衡时观察到的适合度函数的形状以及该模型中动力学的适应程度,并给出强频率依赖下基因型值平衡分布的例子。讨论了对遗传变异维持、间断选择检测和同域物种形成模型的影响。