Evolution of genetic variability and the advantage of sex and recombination in changing environments

The role of recombination and sexual reproduction in enhancing adaptation and population persistence in temporally varying environments is investigated on the basis of a quantitative-genetic multilocus model. Populations are finite, subject to density-dependent regulation with a finite growth rate, diploid, and either asexual or randomly mating and sexual with or without recombination. A quantitative trait is determined by a finite number of loci at which mutation generates genetic variability. The trait is under stabilizing selection with an optimum that either changes at a constant rate in one direction, exhibits periodic cycling, or fluctuates randomly. It is shown by Monte Carlo simulations that if the directional-selection component prevails, then freely recombining populations gain a substantial evolutionary advantage over nonrecombining and asexual populations that goes far beyond that recognized in previous studies. The reason is that in such populations, the genetic variance can increase substantially and thus enhance the rate of adaptation. In nonrecombining and asexual populations, no or much less increase of variance occurs. It is explored by simulation and mathematical analysis when, why, and by how much genetic variance increases in response to environmental change. In particular, it is elucidated how this change in genetic variance depends on the reproductive system, the population size, and the selective regime, and what the consequences for population persistence are

Rate of adaptation in sexuals and asexuals: A solvable model of the Fisher-Muller effect

The adaptation of large asexual populations is hampered by the competition between independently arising beneficial mutations in different individuals, which is known as clonal interference. Fisher and Muller proposed that recombination provides an evolutionary advantage in large populations by alleviating this competition. Based on recent progress in quantifying the speed of adaptation in asexual populations undergoing clonal interference, we present a detailed analysis of the Fisher-Muller mechanism for a model genome consisting of two loci with an infinite number of beneficial alleles each and multiplicative fitness effects. We solve the infinite population dynamics exactly and show that, for a particular, natural mutation scheme, the speed of adaptation in sexuals is twice as large as in asexuals. Guided by the infinite population result and by previous work on asexual adaptation, we postulate an expression for the speed of adaptation in finite sexual populations that agrees with numerical simulations over a wide range of population sizes and recombination rates. The ratio of the sexual to asexual adaptation speed is a function of population size that increases in the clonal interference regime and approaches 2 for extremely large populations. The simulations also show that the imbalance between the numbers of accumulated mutations at the two loci is strongly suppressed even by a small amount of recombination. The generalization of the model to an arbitrary number $L$ of loci is briefly discussed. If each offspring samples the alleles at each locus from the gene pool of the whole population rather than from two parents, the ratio of the sexual to asexual adaptation speed is approximately equal to $L$ in large populations. A possible realization of this scenario is the reassortment of genetic material in RNA viruses with $L$ genomic segments.Comment: Title has been changed. Supporting Information (animation) can be found in the source file. 53 pages. 10 figures. To appear in Genetic

Demographic viability of populations of \u3cem\u3eSilene regis\u3c/em\u3e in midwestern prairies: relationships with fire management, genetic variation, geographic location, population size and isolation

We studied the demographic viability of populations of a long-lived iteroparous prairie perennial, Silene regia, in relation to management regimes, population sizes, geographical region (Ohio and Indiana vs. Missouri and Arkansas), degree of isolation and amount of genetic variation. Demographic data were collected from 16 populations for up to 7 years. This species has high survivorship, slow growth, frequent flowering and episodic seedling recruitment. Matrix projection methods were used to summarize population performance with and without recruitment. Median finite rates of increase by population varied from 0.57 to 1.82 and from 0.44 to 0.99, respectively. Populations with the highest rates of increase had been burned. Six of eight populations, for which stochastic modelling predicted persistence for 1000 years, included fire in their management. None of the five populations with predicted 100-year extinction probabilities of 100% was managed for conservation or burned. An intermediate group of three populations with at least 10% probability of extinction between 100 and 1000 years was not managed, but was none the less kept open by mowing and herbicide application. Analysis of composite elasticities showed that growth and fecundity terms were higher for growing (vs. declining) populations and that growth elasticity was higher in burned than unburned populations. Lack of burning shifts the elasticity spectrum from that typical of open habitat herbs (higher growth and fecundity elasticities) to values usually found for closed habitat herbs (higher survival elasticities). In multivariate analyses predicting finite rates of increase (with and without recruitment), fire management and region were the strongest predictors, followed by genetic variation, population size, isolation and interactions of population size and fire, and region and fire. Populations with the highest rates of increase were burned, eastern, more genetically diverse, larger and less isolated. Discrimination of populations with different extinction risks (three classes) was related mainly to fire, genetic variation and region. Most of these conclusions support conservation biology predictions that population viability will be highest in larger, less-isolated, more genetically diverse populations. However, management and geographic trends have overriding roles affecting demographic viability. Habitat fragmentation and genetic depletion have the potential to threaten residual prairie populations of S. regia, but lack of fire management appears to be the primary short-term threat

Biological evolution through mutation, selection, and drift: An introductory review

Motivated by present activities in (statistical) physics directed towards biological evolution, we review the interplay of three evolutionary forces: mutation, selection, and genetic drift. The review addresses itself to physicists and intends to bridge the gap between the biological and the physical literature. We first clarify the terminology and recapitulate the basic models of population genetics, which describe the evolution of the composition of a population under the joint action of the various evolutionary forces. Building on these foundations, we specify the ingredients explicitly, namely, the various mutation models and fitness landscapes. We then review recent developments concerning models of mutational degradation. These predict upper limits for the mutation rate above which mutation can no longer be controlled by selection, the most important phenomena being error thresholds, Muller's ratchet, and mutational meltdowns. Error thresholds are deterministic phenomena, whereas Muller's ratchet requires the stochastic component brought about by finite population size. Mutational meltdowns additionally rely on an explicit model of population dynamics, and describe the extinction of populations. Special emphasis is put on the mutual relationship between these phenomena. Finally, a few connections with the process of molecular evolution are established.Comment: 62 pages, 6 figures, many reference