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 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 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 Effect of partial selfing and polygenic selection on establishment in a new habitat

2019 ◽  
Author(s):  
Himani Sachdeva
Keyword(s):  
Inbreeding Depression ◽  
Evolutionary Dynamics ◽  
High Rate ◽  
Source Population ◽  
Selfing Rate ◽  
Deleterious Alleles ◽  
Wide Range ◽  
Mate Limitation ◽  
Polygenic Selection

AbstractThis paper analyzes how partial selfing in a large source population influences its ability to colonize a new habitat via the introduction of a few founder individuals. Founders experience inbreeding depression due to partially recessive deleterious alleles as well as maladaptation to the new environment due to selection on a large number of additive loci. I first introduce a simplified version of the Inbreeding History Model (Kelly, 2007) in order to characterize mutation-selection balance in a large, partially selfing source population under selection involving multiple non-identical loci. I then use individual-based simulations to study the eco-evolutionary dynamics of founders establishing in the new habitat under a model of hard selection. The study explores how selfing rate shapes establishment probabilities of founders via effects on both inbreeding depression and adaptability to the new environment, and also distinguishes the effects of selfing on the initial fitness of founders from its effects on the long-term adaptive response of the populations they found. A high rate of (but not complete) selfing is found to aid establishment over a wide range of parameters, even in the absence of mate limitation. The sensitivity of the results to assumptions about the nature of polygenic selection are discussed.

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 The effect of linkage and population size on inbreeding depression due to mutational load

1992 ◽  
Vol 59 (1) ◽  
pp. 49-61 ◽  
Author(s):  
D. Charlesworth ◽  
M. T. Morgan ◽  
B. Charlesworth
Keyword(s):  
Population Size ◽  
Inbreeding Depression ◽  
Finite Population ◽  
Low Frequency ◽  
Mutational Load ◽  
Selfing Rate ◽  
Effective Population ◽  
Modifier Locus ◽  
Mean Fitness ◽  
The Times

SummaryUsing a stochastic model of a finite population in which there is mutation to partially recessive detrimental alleles at many loci, we study the effects of population size and linkage between the loci on the population mean fitness and inbreeding depression values. Although linkage between the selected loci decreases the amount of inbreeding depression, neither population size nor recombination rate have strong effects on these quantities, unless extremely small values are assumed. We also investigate how partial linkage between the loci that determine fitness affects the invasion of populations by alleles at a modifier locus that controls the selfing rate. In most of the cases studied, the direction of selection on modifiers was consistent with that found in our previous deterministic calculations. However, there was some evidence that linkage between the modifier locus and the selected loci makes outcrossing less likely to evolve; more losses of alleles promoting outcrossing occurred in runs with linkage than in runs with free recombination. We also studied the fate of neutral alleles introduced into populations carrying detrimental mutations. The times to loss of neutral alleles introduced at low frequency were shorter than those predicted for alleles in the absence of selected loci, taking into account the reduction of the effective population size due to inbreeding. Previous studies have been confined to outbreeding populations, and to alleles at frequencies close to one-half, and have found an effect in the opposite direction. It therefore appears that associations between neutral and selected loci may produce effects that differ according to the initial frequencies of the neutral alleles.

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.

scholarly journals The apparent selection on neutral marker loci in partially inbreeding populations

1991 ◽  
Vol 57 (2) ◽  
pp. 159-175 ◽  
Author(s):  
D. Charlesworth
Keyword(s):  
Inbreeding Depression ◽  
Relative Fitness ◽  
Heterozygote Advantage ◽  
Neutral Locus ◽  
Fitness Value ◽  
Marker Loci ◽  
Neutral Marker ◽  
Computer Calculations

SummaryDeterministic computer calculations were used to investigate the effects on the fitnesses of genotypes at neutral loci that are caused by associations with several linked or unlinked selected loci, in partially self fertilizing populations. Both mutation to partially recessive alleles and heterozygote advantage at the selected loci were studied. In the heterozygote advantage models, either arbitrary linkage between all loci was modelled, with a single neutral locus, or many unlinked selected and neutral loci were modelled. Large apparent overdominance could be generated in all types of model studied. As has previously been suggested, these types of effect can explain the observed associations between fitness and heterozygosity in partially inbreeding populations. There were also apparent fitness differences between the genotypes at the neutral locus among the progeny produced by selfing, especially with linkage between the neutral and selected loci. There is thus no genotype-independent fitness value for these progeny. Marker based methods for estimating the relative fitness of selfed and outcrossed progeny assume equality of these fitnesses, and will therefore be inaccurate (with in most cases a bias towards overestimating the degree of inbreeding depression) when there is linkage between the neutral marker loci and loci determining fitness.

Estimation of Deleterious Genomic Mutation Parameters in Natural Populations by Accounting for Variable Mutation Effects Across Loci

Genetics ◽  
2002 ◽  
Vol 162 (3) ◽  
pp. 1487-1500 ◽  
Author(s):  
Hong-Wen Deng ◽  
Guimin Gao ◽  
Jin-Long Li
Keyword(s):  
Statistical Method ◽  
Mutation Rate ◽  
Genetic Variance ◽  
Natural Populations ◽  
Evolutionary Genetics ◽  
Simulation Studies ◽  
Fitness Effects ◽  
Mean Fitness ◽  
The Mean ◽  
Genomic Mutation

Abstract The genomes of all organisms are subject to continuous bombardment of deleterious genomic mutations (DGM). Our ability to accurately estimate various parameters of DGM has profound significance in population and evolutionary genetics. The Deng-Lynch method can estimate the parameters of DGM in natural selfing and outcrossing populations. This method assumes constant fitness effects of DGM and hence is biased under variable fitness effects of DGM. Here, we develop a statistical method to estimate DGM parameters by considering variable mutation effects across loci. Under variable mutation effects, the mean fitness and genetic variance for fitness of parental and progeny generations across selfing/outcrossing in outcrossing/selfing populations and the covariance between mean fitness of parents and that of their progeny are functions of DGM parameters: the genomic mutation rate U, average homozygous effect s, average dominance coefficient h, and covariance of selection and dominance coefficients cov(h, s). The DGM parameters can be estimated by the algorithms we developed herein, which may yield improved estimation of DGM parameters over the Deng-Lynch method as demonstrated by our simulation studies. Importantly, this method is the first one to characterize cov(h, s) for DGM.

scholarly journals Adapting the engine to the fuel: mutator populations can reduce the mutational load by reorganizing their genome structure

2019 ◽  
Vol 19 (1) ◽  
Author(s):  
Jacob Pieter Rutten ◽  
Paulien Hogeweg ◽  
Guillaume Beslon
Keyword(s):  
Mutation Rate ◽  
Genome Structure ◽  
Mutational Load ◽  
Bacterial Populations ◽  
Long Run ◽  
Stable Conditions ◽  
Long Time ◽  
Natural Isolates ◽  
Mean Fitness ◽  
The Mean

Abstract Background Mutators are common in bacterial populations, both in natural isolates and in the lab. The fate of these lineages, which mutation rate is increased up to 100 ×, has long been studied using population genetics models, showing that they can spread in a population following an environmental change. However in stable conditions, they suffer from the increased mutational load, hence being overcome by non-mutators. However, these results don’t take into account the fact that an elevated mutation rate can impact the genetic structure, hence changing the sensitivity of the population to mutations. Here we used Aevol, an in silico experimental evolution platform in which genomic structures are free to evolve, in order to study the fate of mutator populations evolving for a long time in constant conditions. Results Starting from wild-types that were pre-evolved for 300,000 generations, we let 100 mutator populations (point mutation rate ×100) evolve for 100,000 further generations in constant conditions. As expected all populations initially undergo a fitness loss. However, after that the mutator populations started to recover. Most populations ultimately recovered their ancestors fitness, and a significant fraction became even fitter than the non-mutator control clones that evolved in parallel. By analyzing the genomes of the mutators, we show that the fitness recovery is due to two mechanisms: i. an increase in robustness through compaction of the coding part of the mutator genomes, ii. an increase of the selection coefficient that decreases the mean-fitness of the population. Strikingly the latter is due to the accumulation of non-coding sequences in the mutators genomes. Conclusion Our results show that the mutational burden that is classically thought to be associated with mutator phenotype is escapable. On the long run mutators adapted their genomes and reshaped the distribution of mutation effects. Therewith the lineage is able to recover fitness even though the population still suffers the elevated mutation rate. Overall these results change our view of mutator dynamics: by being able to reduce the deleterious effect of the elevated mutation rate, mutator populations may be able to last for a very long time; A situation commonly observed in nature.

scholarly journals Epistasis, inbreeding depression and the evolution of self-fertilization

2019 ◽  
Author(s):  
Diala Abu Awad ◽  
Denis Roze
Keyword(s):  
Inbreeding Depression ◽  
Stabilizing Selection ◽  
Linkage Disequilibria ◽  
Deleterious Alleles ◽  
Mating System Evolution ◽  
Special Cases ◽  
Self Fertilization ◽  
Mean Fitness ◽  
Selection For ◽  
Individual Based Simulation

ABSTRACTInbreeding depression resulting from partially recessive deleterious alleles is thought to be the main genetic factor preventing self-fertilizing mutants from spreading in outcrossing hermaphroditic populations. However, deleterious alleles may also generate an advantage to selfers in terms of more efficient purging, while the effects of epistasis among those alleles on inbreeding depression and mating system evolution remain little explored. In this paper, we use a general model of selection to disentangle the effects of different forms of epistasis (additive-by-additive, additive-by-dominance and dominance-by-dominance) on inbreeding depression and on the strength of selection for selfing. Models with fixed epistasis across loci, and models of stabilizing selection acting on quantitative traits (generating distributions of epistasis) are considered as special cases. Besides its effects on inbreeding depression, epistasis may increase the purging advantage associated with selfing (when it is negative on average), while the variance in epistasis favors selfing through the generation of linkage disequilibria that increase mean fitness. Approximations for the strengths of these effects are derived, and compared with individual-based simulation results.

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