The Hill-Robertson effect reduces effective population size through linked selection.
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Peer-reviewed population genetics literature confirms that selection at linked sites, known as the Hill-Robertson effect, reduces the effective population size experienced by genomic sites.
(i) A computer simulation study has been made of selection on two linked loci in small populations, where both loci were assumed to have additive effects on the character under selection with no interaction between loci. If N is the effective population size, i the intensity of selection in standard units, α and β measure the effects of the two loci on the character under selection as a proportion of the pheno-typic standard deviation and c is the crossover distance between them, it was shown that the selection process can be completely specified by Ni α, Ni βand Nc and the initial gene frequencies and linkage disequilibrium coefficient. It is then easily possible to generalize from computer runs at only one population size. All computer runs assumed an initial population at linkage equilibrium between the two loci. Analysis of the results was greatly simplified by considering the influence of segregation at the second locus on the chance of fixation at the first (defined as the proportion of replicate lines in which the favoured allele was eventually fixed). (ii) The effects of linkage are sufficiently described by Nc. The relationship between chance of fixation at the limit and linkage distance (expressed as 2Nc /( 2Nc + 1)) was linear in the majority of computer runs. (iii) When gene frequency changes under independent segregation were small, linkage had no effect on the advance under selection. In general, segregation at the second locus had no detectable influence on the chance of fixation at the first if the gene effects at the second were less than one-half those at the first. With larger gene effects at the second locus, the chance of fixation passed through a minimum and then rose again. For two loci to have a mutual influence on one another, their effects on the character under selection should not differ by a factor of more than two. (iv) Under conditions of suitable relative gene effects, the influence of segregation at the second locus was very dependent on the initial frequency of the desirable allele. The chance of fixation at the first, plotted against initial frequency of the desirable allele at the second, passed through a minimum when the chance of fixation at the second locus was about 0·8. (v) A transformation was found which made the influence of segregation at the second locus on the chance of fixation at the first almost independent of initial gene frequency at the first and of gene effects at the first locus when these are small. (vi) In the population of gametes at final fixation, linkage was not at equilibrium and there was an excess of repulsion gametes. (vii) The results were extended to a consideration of the effect of linkage on the limits under artificial selection. Linkage proved only to be of importance when the two loci had roughly equal effects on the character under selection. The maximum effect on the advance under selection occurred when the chance of fixation at both of the loci was between 0·7 and 0·8. When the advance under selection is most sensitive to changes in recombination value, a doubling of the latter in no case increased the advance under selection by more than about 6%. The proportion selected to give maximum advance under individual selection (0·5 under independent segregation) was increased, but only very slightly, when linkage is important. (viii) These phenomena could be satisfactorily accounted for in terms of the time scale of the selection process and the effective size of the population within which changes of gene frequency at the locus with smaller effect must take place.
The accurate estimation of the distribution of fitness effects (DFE) of new mutations is critical for population genetic inference but remains a challenging task. While various methods have been developed for DFE inference using the site frequency spectrum of putatively neutral and selected sites, their applicability in species with diverse life history traits and complex demographic scenarios is not well understood. Selfing is common among eukaryotic species and can lead to decreased effective recombination rates, increasing the effects of selection at linked sites, including interference between selected alleles. We employ forward simulations to investigate the limitations of current DFE estimation approaches in the presence of selfing and other model violations, such as linkage, departures from semidominance, population structure, and uneven sampling. We find that distortions of the site frequency spectrum due to Hill-Robertson interference in highly selfing populations lead to mis-inference of the deleterious DFE of new mutations. Specifically, when inferring the distribution of selection coefficients, there is an overestimation of nearly neutral and strongly deleterious mutations and an underestimation of mildly deleterious mutations when interference between selected alleles is pervasive. In addition, the presence of cryptic population structure with low rates of migration and uneven sampling across subpopulations leads to the false inference of a deleterious DFE skewed towards effectively neutral/mildly deleterious mutations. Finally, the proportion of adaptive substitutions estimated at high rates of selfing is substantially overestimated. Our observations apply broadly to species and genomic regions with little/no recombination and where interference might be pervasive.
Abstract The process of evolution at a given site in the genome can be influenced by the action of selection at other sites, especially when these are closely linked to it. Such selection reduces the effective population size experienced by the site in question (the Hill–Robertson effect), reducing the level of variability and the efficacy of selection. In particular, deleterious variants are continually being produced by mutation and then eliminated by selection at sites throughout the genome. The resulting reduction in variability at linked neutral or nearly neutral sites can be predicted from the theory of background selection, which assumes that deleterious mutations have such large effects that their behavior in the population is effectively deterministic. More weakly selected mutations can accumulate by Muller’s ratchet after a shutdown of recombination, as in an evolving Y chromosome. Many functionally significant sites are probably so weakly selected that Hill–Robertson interference undermines the effective strength of selection upon them, when recombination is rare or absent. This leads to large departures from deterministic equilibrium and smaller effects on linked neutral sites than under background selection or Muller’s ratchet. Evidence is discussed that is consistent with the action of these processes in shaping genome-wide patterns of variation and evolution.
Abstract The process of evolution at a given site in the genome can be influenced by the action of selection at other sites, especially when these are closely linked to it. Such selection reduces the effective population size experienced by the site in question (the Hill–Robertson effect), reducing the level of variability and the efficacy of selection. In particular, deleterious variants are continually being produced by mutation and then eliminated by selection at sites throughout the genome. The resulting reduction in variability at linked neutral or nearly neutral sites can be predicted fr
Recombination can impede ecological speciation with gene flow by mixing locally adapted genotypes with maladapted migrant genotypes from a divergent population. In such a scenario, suppression of recombination can be selectively favoured. However, in finite populations evolving under the influence of random genetic drift, recombination can also facilitate adaptation by reducing Hill–Robertson interference between loci under selection. In this case, increased recombination rates can be favoured. Although these two major effects on recombination have been studied individually, their joint effect
Sex and recombination are widespread, but explaining these phenomena has been one of the most difficult problems in evolutionary biology. Recombination is advantageous when different individuals in a population carry different advantageous alleles. By bringing together advantageous alleles onto the same chromosome, recombination speeds up the process of adaptation and opposes the fixation of harmful mutations by means of Muller's ratchet. Nevertheless, adaptive substitutions favour sex and recombination only if the rate of adaptive mutation is high, and Muller's ratchet operates only in small or asexual populations. Here, by tracking the fate of modifier alleles that alter the frequency of sex and recombination, we show that background selection against deleterious mutant alleles provides a stochastic advantage to sex and recombination that increases with population size. The advantage arises because, with low levels of recombination, selection at other loci severely reduces the effective population size and genetic variance in fitness at a focal locus (the Hill-Robertson effect), making a population less able to respond to selection and to rid itself of deleterious mutations. Sex and recombination reveal the hidden genetic variance in fitness by combining chromosomes of intermediate fitness to create chromosomes that are relatively free of (or are loaded with) deleterious mutations. This increase in genetic variance within finite populations improves the response to selectio
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