scholarly journals Selection, Load and Inbreeding Depression in a Large Metapopulation

Genetics ◽  
2002 ◽  
Vol 160 (3) ◽  
pp. 1191-1202 ◽  
Author(s):  
Michael C Whitlock
Keyword(s):  
Population Structure ◽  
Inbreeding Depression ◽  
Population Subdivision ◽  
Response To Selection ◽  
Mutation Load ◽  
Deleterious Alleles ◽  
Subdivided Population ◽  
Soft Selection ◽  
Mean Fitness ◽  
The Mean

Abstract The subdivision of a species into local populations causes its response to selection to change, even if selection is uniform across space. Population structure increases the frequency of homozygotes and therefore makes selection on homozygous effects more effective. However, population subdivision can increase the probability of competition among relatives, which may reduce the efficacy of selection. As a result, the response to selection can be either increased or decreased in a subdivided population relative to an undivided one, depending on the dominance coefficient FST and whether selection is hard or soft. Realistic levels of population structure tend to reduce the mean frequency of deleterious alleles. The mutation load tends to be decreased in a subdivided population for recessive alleles, as does the expected inbreeding depression. The magnitude of the effects of population subdivision tends to be greatest in species with hard selection rather than soft selection. Population structure can play an important role in determining the mean fitness of populations at equilibrium between mutation and selection.

scholarly journals The mutation load under tetrasomic inheritance and its consequences for the evolution of the selfing rate in autotetraploid species

1999 ◽  
Vol 74 (1) ◽  
pp. 31-42 ◽  
Author(s):  
J. RONFORT
Keyword(s):  
Inbreeding Depression ◽  
Deleterious Mutation ◽  
Stable State ◽  
Approximate Solutions ◽  
Balance Model ◽  
Deleterious Mutations ◽  
Selfing Rate ◽  
Mutation Load ◽  
Mean Fitness ◽  
The Mean

Single-locus equilibrium frequencies of a partially recessive deleterious mutation under the mutation–selection balance model are derived for partially selfing autotetraploid populations. Assuming multiplicative fitness interactions among loci, approximate solutions for the mean fitness and inbreeding depression values are also derived for the multiple locus case and compared with expectations for the diploid model. As in diploids, purging of deleterious mutations through consanguineous matings occurs in autotetraploid populations, i.e. the equilibrium mutation load is a decreasing function of the selfing rate. However, the variation of inbreeding depression with the selfing rate depends strongly on the dominance coefficients associated with the three heterozygous genotypes. Inbreeding depression can either increase or decrease with the selfing rate, and does not always vary monotonically. Expected issues for the evolution of the selfing rate consequently differ depending on the dominance coefficients. In some cases, expectations for the evolution of the selfing rate resemble expectations in diploids; but particular sets of dominance coefficients can be found that lead to either complete selfing or intermediate selfing rates as unique evolutionary stable state.

scholarly journals Multilocus models of inbreeding depression with synergistic selection and partial self-fertilization

1991 ◽  
Vol 57 (2) ◽  
pp. 177-194 ◽  
Author(s):  
B. Charlesworth ◽  
M. T. Morgan ◽  
D. Charlesworth
Keyword(s):  
Inbreeding Depression ◽  
Mutation Rate ◽  
Approximate Expression ◽  
Heterozygote Advantage ◽  
Selfing Rate ◽  
Deleterious Alleles ◽  
Plausible Model ◽  
Computer Calculations ◽  
Mean Fitness ◽  
The Mean

SummaryMean fitness and inbreeding depression values in multi-locus models of the control of fitness were studied, using both a model of mutation to deleterious alleles, and a model of heterozygote advantage. Synergistic fitness interactions between loci were assumed, to find out if this more biologically plausible model altered the conclusions we obtained previously using a model of multiplicative interactions. Systems of unlinked loci were assumed. We used deterministic computer calculations, and approximations based on normal or Poisson theory. These approximations gave good agreement with the exact results for some regions of the parameter space. In the mutational model, we found that the effect of synergism was to lower the number of mutant alleles per individual, and thus to increase the mean fitness, compared with the multiplicative case. Inbreeding depression, however, was increased. Similar effects on mean fitness and inbreeding depression were found for the case of heterozygote advantage. For that model, the results were qualitatively similar to those previously obtained assuming multiplicativity. With the mutational load model, however, the mean fitness sometimes decreased, and the inbreeding depression increased, at high selfing rates, after declining as the selfing rate increased from zero. We also studied the behaviour of modifier alleles that changed the selfing rate, introduced into equilibrium populations. In general, the results were similar to those with the multiplicative model, but in some cases an ESS selfing rate, with selfing slightly below one, existed. Finally, we derive an approximate expression for the inbreeding depression in completely selfing populations. This depends only on the mutation rate and the dominance coefficient and can therefore be used to obtain estimates of the mutation rate to mildly deleterious alleles for plant species.

scholarly journals Lethals in subdivided populations

2005 ◽  
Vol 86 (1) ◽  
pp. 41-51 ◽  
Author(s):  
SYLVAIN GLÉMIN
Keyword(s):  
Population Structure ◽  
Population Size ◽  
Inbreeding Depression ◽  
Deleterious Mutations ◽  
Mutation Load ◽  
General Picture ◽  
Analytical Approximations ◽  
Size Population ◽  
Subdivided Populations ◽  
Numerical Approaches

The fate of lethal alleles in populations is of interest in evolutionary and conservation biology for several reasons. For instance, lethals may contribute substantially to inbreeding depression. The frequency of lethal alleles depends on population size, but it is not clear how it is affected by population structure. By analysing the case of the infinite island model by numerical approaches and analytical approximations it is shown that, like population size, population structure affects the fate of lethal alleles if dominance levels are low. Inbreeding depression caused by such alleles is also affected by the population structure, whereas the mutation load is only weakly affected. Heterosis also depends on population structure, but it always remains low, of the order of the mutation rate or less. These patterns are compared with those caused by mildly deleterious mutations to give a general picture of the effect of population structure on inbreeding depression, heterosis, and the mutation load.

scholarly journals Mutation load decreases with haplotype age in wild Soay sheep

2021 ◽  
Author(s):  
M.A. Stoffel ◽  
S.E. Johnston ◽  
J.G. Pilkington ◽  
J.M Pemberton
Keyword(s):  
Inbreeding Depression ◽  
Purifying Selection ◽  
Small Population ◽  
First Year ◽  
Mutation Load ◽  
Soay Sheep ◽  
Recessive Mutations ◽  
Deleterious Alleles ◽  
Lower Population

AbstractRuns of homozygosity (ROH) are pervasive in diploid genomes and expose the effects of deleterious recessive mutations, but how exactly these regions contribute to variation in fitness remains unclear. Here, we combined empirical analyses and simulations to explore the deleterious effects of ROH with varying genetic map lengths in wild Soay sheep. Using a long-term dataset of 4,592 individuals genotyped at 417K SNPs, we found that inbreeding depression increases with ROH length. A 1% genomic increase in long ROH (>12.5cM) reduced the odds of first-year survival by 12%, compared to only 7% for medium ROH (1.56-12.5cM), while short ROH (<1.56cM) had no effect on survival. We show by forward genetic simulations that this is predicted: compared with shorter ROH, long ROH will have higher densities of deleterious alleles, with larger average effects on fitness and lower population frequencies. Taken together, our results are consistent with the idea that the mutation load decreases in older haplotypes underlying shorter ROH, where purifying selection has had more time to purge deleterious mutations. Finally, our study demonstrates that strong inbreeding depression can persist despite ongoing purging in a historically small population.

scholarly journals Selection responses of means and inbreeding depression for female fecundity in Drosophila melanogaster suggest contributions from intermediate-frequency alleles to quantitative trait variation

2007 ◽  
Vol 89 (2) ◽  
pp. 85-91 ◽  
Author(s):  
BRIAN CHARLESWORTH ◽  
TAKAHIRO MIYO ◽  
HELEN BORTHWICK
Keyword(s):  
Drosophila Melanogaster ◽  
Inbreeding Depression ◽  
Quantitative Trait ◽  
Balancing Selection ◽  
Evolutionary Genetics ◽  
Intermediate Frequency ◽  
Female Fecundity ◽  
Deleterious Alleles ◽  
The Mean ◽  
Change In Mean

SummaryThe extent to which quantitative trait variability is caused by rare alleles maintained by mutation, versus intermediate-frequency alleles maintained by balancing selection, is an unsolved problem of evolutionary genetics. We describe the results of an experiment to examine the effects of selection on the mean and extent of inbreeding depression for early female fecundity in Drosophila melanogaster. Theory predicts that rare, partially recessive deleterious alleles should cause a much larger change in the effect of inbreeding than in the mean of the outbred population, with the change in inbreeding effect having an opposite sign to the change in mean. The present experiment fails to support this prediction, suggesting that intermediate-frequency alleles contribute substantially to genetic variation in early fecundity.

scholarly journals A sampling theory of selectively neutral alleles in a subdivided population.

Genetics ◽  
1988 ◽  
Vol 119 (3) ◽  
pp. 721-729
Author(s):  
E R Tillier ◽  
G B Golding
Keyword(s):  
Population Structure ◽  
Population Subdivision ◽  
Continuous Approximation ◽  
Sample Distribution ◽  
Migration Rates ◽  
Practical Possibility ◽  
Structured Population ◽  
Subdivided Population ◽  
Subdivided Populations ◽  
Actual Size

Abstract Ewens' sampling distribution is investigated for a structured population. Samples are assumed to be taken from a single subpopulation that exchanges migrants with other subpopulations. A complete description of the probability distribution for such samples is not a practical possibility but an equilibrium approximation can be found. This approximation extracts the information necessary for constructing a continuous approximation to the complete distribution using known values of the distribution and its derivatives in randomly mating populations. It is shown that this approximation is as complete a description of a single biologically realistic subpopulation as is possible given standard uncertainties about the actual size of the migration rates, relative sizes of each of the subpopulations and other factors that might affect the genetic structure of a subpopulation. Any further information must be gained at the expense of generality. This approximation is used to investigate the effect of population subdivision on Watterson's test of neutrality. It is known that the infinite allele, sample distribution is independent of mutation rate when made conditional on the number of alleles in the sample. It is shown that the conditional, infinite allele, sample distribution from this approximation is also independent of population structure and hence Watterson's test is still approximately valid for subdivided populations.

scholarly journals Genomic evidence for inbreeding depression and purging of deleterious genetic variation in Indian tigers

2021 ◽  
Vol 118 (49) ◽  
pp. e2023018118
Author(s):  
Anubhab Khan ◽  
Kaushalkumar Patel ◽  
Harsh Shukla ◽  
Ashwin Viswanathan ◽  
Tom van der Valk ◽  
...  
Keyword(s):  
Inbreeding Depression ◽  
Purifying Selection ◽  
Isolated Population ◽  
Loss Of Function ◽  
Mutation Load ◽  
Genetic Rescue ◽  
Frequency Distributions ◽  
Recessive Mutations ◽  
Deleterious Alleles ◽  
Large Populations

Increasing habitat fragmentation leads to wild populations becoming small, isolated, and threatened by inbreeding depression. However, small populations may be able to purge recessive deleterious alleles as they become expressed in homozygotes, thus reducing inbreeding depression and increasing population viability. We used whole-genome sequences from 57 tigers to estimate individual inbreeding and mutation load in a small–isolated and two large–connected populations in India. As expected, the small–isolated population had substantially higher average genomic inbreeding (FROH = 0.57) than the large–connected (FROH = 0.35 and FROH = 0.46) populations. The small–isolated population had the lowest loss-of-function mutation load, likely due to purging of highly deleterious recessive mutations. The large populations had lower missense mutation loads than the small–isolated population, but were not identical, possibly due to different demographic histories. While the number of the loss-of-function alleles in the small–isolated population was lower, these alleles were at higher frequencies and homozygosity than in the large populations. Together, our data and analyses provide evidence of 1) high mutation load, 2) purging, and 3) the highest predicted inbreeding depression, despite purging, in the small–isolated population. Frequency distributions of damaging and neutral alleles uncover genomic evidence that purifying selection has removed part of the mutation load across Indian tiger populations. These results provide genomic evidence for purifying selection in both small and large populations, but also suggest that the remaining deleterious alleles may have inbreeding-associated fitness costs. We suggest that genetic rescue from sources selected based on genome-wide differentiation could offset any possible impacts of inbreeding depression.

scholarly journals Genomic evidence for inbreeding depression and purging of deleterious genetic variation in Indian tigers

2021 ◽  
Author(s):  
Anubhab Khan ◽  
Kaushalkumar Patel ◽  
Harsh Shukla ◽  
Ashwin Viswanathan ◽  
Tom van der Valk ◽  
...  
Keyword(s):  
Inbreeding Depression ◽  
Purifying Selection ◽  
Isolated Population ◽  
Loss Of Function ◽  
Mutation Load ◽  
Genetic Rescue ◽  
Frequency Distributions ◽  
Recessive Mutations ◽  
Deleterious Alleles ◽  
Large Populations

Increasing habitat fragmentation leads to wild populations becoming small, isolated, and threatened by inbreeding depression. However, small populations may be able to purge recessive deleterious alleles as they become expressed in homozygotes, thus reducing inbreeding depression and increasing population viability. We used genome sequencing of 57 tigers to estimate individual inbreeding and mutation loads in a small-isolated, and two large-connected populations in India. As expected, the small-isolated population had substantially higher average genomic inbreeding (FROH=0.57) than the large-connected (FROH=0.35 and FROH=0.46) populations. The small-isolated population had the lowest loss-of-function mutation load, likely due to purging of highly deleterious recessive mutations. The large populations had lower missense mutation loads than the small-isolated population, but were not identical, possibly due to different demographic histories. While the number of the loss-of-function alleles in the small-isolated population was lower, these alleles were at high frequencies and homozygosity than in the large populations. Together, our data and analyses provide evidence of (a) high mutation load; (b) purging and (c) the highest predicted inbreeding depression, despite purging, in the small-isolated population. Frequency distributions of damaging and neutral alleles uncover genomic evidence that purifying selection has removed part of the mutation load across Indian tiger populations. These results provide genomic evidence for purifying selection in both small and large populations, but also suggest that the remaining deleterious alleles may have inbreeding associated fitness costs. We suggest that genetic rescue from sources selected based on genome-wide differentiation should offset any possible impacts of inbreeding depression.

scholarly journals King's formula for the mutation load with epistasis.

Genetics ◽  
1988 ◽  
Vol 120 (3) ◽  
pp. 853-856
Author(s):  
A S Kondrashov ◽  
J F Crow
Keyword(s):  
Mutation Rate ◽  
Haploid Genome ◽  
Relative Fitness ◽  
Mutation Load ◽  
Asexual Population ◽  
Deleterious Alleles ◽  
Correct Formula ◽  
The Mean ◽  
Mutant Genes ◽  
New Mutations

Abstract A formula by J. L. King gives the equilibrium mutation load as L = 2 sigma ui(1 - qi)/z - x) in which ui is the mutation rate to deleterious alleles at the ith locus, qi is the frequency of mutant alleles at this locus, x is the mean number of such mutant genes per individual before selection, z is the mean number in individuals eliminated by selection, and the summation is over all relevant loci. We show that this rule is inaccurate for intense selection and that a correct formula is L = 2 sigma ui(1 - qi) w/(z - x) = 2U w/(z - x) = 2U/(z - x + 2U) in which U is the mean number of new mutations per haploid genome in the population and w is the mean relative fitness before selection. If w/(z - x) less than 1/2, the mutation load is less than the Haldane value (U less than or equal to L less than or equal to 2U) and can be considerably less. In a diploid asexual population, however, with independent occurrence of mutations, L = 1 - e-2U regardless of the mode of selection.

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