scholarly journals An estimator of the Opportunity for Selection that is independent of mean fitness

Evolution ◽  
2020 ◽  
Vol 74 (9) ◽  
pp. 1942-1953 ◽  
Author(s):  
Robin S. Waples

2018 ◽  
Author(s):  
Joshua Franklin ◽  
Thomas LaBar ◽  
Christoph Adami

AbstractBackgroundPopulations exposed to a high mutation rate harbor abundant deleterious genetic variation, leading to depressed mean fitness. This reduction in mean fitness presents an opportunity for selection to restore adaptation through the evolution of mutational robustness. In extreme cases, selection for mutational robustness can lead to “flat” genotypes (with low fitness but high robustness) out-competing “fit” genotypes with high fitness but low robustness—a phenomenon known as “survival of the flattest”. While this effect was previously explored using the digital evolution system Avida, a complete analysis of the local fitness landscapes of “fit” and “flat” genotypes has been lacking, leading to uncertainty about the genetic basis of the survival of the flattest effect.ResultsHere, we repeated the survival of the flattest study and analyzed the mutational neighborhoods of fit and flat genotypes. We found that flat genotypes, compared to the fit genotypes, had a reduced likelihood of deleterious mutations as well as an increased likelihood of neutral and, surprisingly, of lethal mutations. This trend holds for mutants one to four substitutions away from the wild-type sequence. We also found that flat genotypes have, on average, no epistasis between mutations, while fit genotypes have, on average, positive epistasis.ConclusionsOur results demonstrate that the genetic causes of mutational robustness on complex fitness landscapes are multifaceted. While the traditional idea of the survival of the flattest effect emphasized the evolution of increased neutrality, others have argued for increased mutational sensitivity in response to strong mutational loads. Our results show that both increased neutrality and increased lethality can lead to the evolution of mutational robustness. Furthermore, strong negative epistasis is not required for mutational sensitivity to lead to mutational robustness. Overall, these results suggest that mutational robustness is achieved by minimizing heritable deleterious variation.



2019 ◽  
Vol 25 (3) ◽  
pp. 250-262 ◽  
Author(s):  
Joshua Franklin ◽  
Thomas LaBar ◽  
Christoph Adami

Populations exposed to a high mutation rate harbor abundant deleterious genetic variation, leading to depressed mean fitness. This reduction in mean fitness presents an opportunity for selection to restore fitness through the evolution of mutational robustness. In extreme cases, selection for mutational robustness can lead to flat genotypes (with low fitness but high robustness) outcompeting fit genotypes (with high fitness but low robustness)—a phenomenon known as survival of the flattest. While this effect was previously explored using the digital evolution system Avida, a complete analysis of the local fitness landscapes of fit and flat genotypes has been lacking, leading to uncertainty about the genetic basis of the survival-of-the-flattest effect. Here, we repeated the survival-of-the-flattest study and analyzed the mutational neighborhoods of fit and flat genotypes. We found that the flat genotypes, compared to the fit genotypes, had a reduced likelihood of deleterious mutations as well as an increased likelihood of neutral and, surprisingly, of lethal mutations. This trend holds for mutants one to four substitutions away from the wild-type sequence. We also found that flat genotypes have, on average, no epistasis between mutations, while fit genotypes have, on average, positive epistasis. Our results demonstrate that the genetic causes of mutational robustness on complex fitness landscapes are multifaceted. While the traditional idea of the survival of the flattest emphasized the evolution of increased neutrality, others have argued for increased mutational sensitivity in response to strong mutational loads. Our results show that both increased neutrality and increased lethality can lead to the evolution of mutational robustness. Furthermore, strong negative epistasis is not required for mutational sensitivity to lead to mutational robustness. Overall, these results suggest that mutational robustness is achieved by minimizing heritable deleterious variation.



2019 ◽  
Author(s):  
Ivain Martinossi-Allibert ◽  
Johanna Liljestrand Rönn ◽  
Elina Immonen

AbstractEnvironmental and physiological conditions affect how individual variation is expressed and translated into variance in fitness, the opportunity for natural selection. Competition for limiting resources can magnify variance in fitness and therefore selection, while abundance of resources should reduce it. But even in a common environment the strength of selection can be expected to differ across the sexes, as their fitness is often limited by different resources. Indeed most taxa show a greater opportunity for selection in males than in females, a bias often ascribed to intense competition among males for access to mating partners. This sex-bias could reverberate on many aspects of evolution, from speed of adaptation to genome evolution. It is unclear however, whether the sex-bias in opportunity for selection is robust to variations in environment or physiological condition that limit sex-specific resources. Here we test this in the model species C. maculatus by comparing female and male variance in relative fitness (opportunity for selection) under mate competition (i) with and without limitation of quality oviposition sites, and (ii) under delayed age at oviposition. Decreasing the abundance of the resource key to females or increasing their reproductive age was indeed challenging as shown by a reduction in mean fitness, however variance in fitness remained male-biased across the three treatments, with even an increased male-bias when females were limited by oviposition sites. This suggests that males remain the more variable sex independent of context, and that the opportunity for selection through males is indirectly affected by female-specific resource limitation.



2020 ◽  
Author(s):  
Robin S. Waples

AbstractVariation among individuals in number of offspring (fitness, k) sets an upper limit to the evolutionary response to selection. This constraint is quantified by Crow’s Opportunity for Selection (I), which is the variance in relative fitness . Crow’s I has been widely used but remains controversial because it depends on mean offspring number in a sample . Here I used a generalized Wright-Fisher model that allows for unequal probabilities of producing offspring to evaluate behavior of Crow’s I and related indices under a wide range of sampling scenarios. Analytical and numerical results are congruent and show that rescaling the sample variance to its expected value at a fixed removes dependence of I on mean offspring number, but the result still depends on choice of . A new index is introduced, , which makes Î independent of sample without the need for variance rescaling. ΔI has a straightforward interpretation as the component of variance in relative fitness that exceeds that expected under a null model of random reproductive success. ΔI can be used to directly compare estimates of the Opportunity for Selection for samples from different studies, different sexes, and different life stages.



Genetics ◽  
1997 ◽  
Vol 146 (2) ◽  
pp. 471-479 ◽  
Author(s):  
Michael Travisano

The effect of environment on adaptation and divergence was examined in two sets of populations of Escherichia coli selected for 1000 generations in either maltose- or glucose-limited media. Twelve replicate populations selected in maltose-limited medium improved in fitness in the selected environment, by an average of 22.5%. Statistically significant among-population genetic variation for fitness was observed during the course of the propagation, but this variation was small relative to the fitness improvement. Mean fitness in a novel nutrient environment, glucose-limited medium, improved to the same extent as in the selected environment, with no statistically significant among-population genetic variation. In contrast, 12 replicate populations previously selected for 1000 generations in glucose-limited medium showed no improvement, as a group, in fitness in maltose-limited medium and substantial genetic variation. This asymmetric pattern of correlated responses suggests that small changes in the environment can have profound effects on adaptation and divergence.



Genetics ◽  
2003 ◽  
Vol 164 (3) ◽  
pp. 1119-1128 ◽  
Author(s):  
Elie S Dolgin ◽  
Sarah P Otto

AbstractThe segregation of alleles disrupts genetic associations at overdominant loci, causing a sexual population to experience a lower mean fitness compared to an asexual population. To investigate whether circumstances promoting increased sex exist within a population with heterozygote advantage, a model is constructed that monitors the frequency of alleles at a modifier locus that changes the relative allocation to sexual and asexual reproduction. The frequency of these modifier alleles changes over time as a correlated response to the dynamics at a fitness locus under overdominant selection. Increased sex can be favored in partially sexual populations that inbreed to some extent. This surprising finding results from the fact that inbred populations have an excess of homozygous individuals, for whom sex is always favorable. The conditions promoting increased levels of sex depend on the selection pressure against the homozygotes, the extent of sex and inbreeding in the population, and the dominance of the invading modifier allele.



Genetics ◽  
1974 ◽  
Vol 77 (3) ◽  
pp. 569-589
Author(s):  
Martin L Tracey ◽  
Francisco J Ayala

ABSTRACT Recent studies of genetically controlled enzyme variation lead to an estimation that at least 30 to 60% of the structural genes are polymorphic in natural populations of many vertebrate and invertebrate species. Some authors have argued that a substantial proportion of these polymorphisms cannot be maintained by natural selection because this would result in an unbearable genetic load. If many polymorphisms are maintained by heterotic natural selection, individuals with much greater than average proportion of homozygous loci should have very low fitness. We have measured in Drosophila melanogaster the fitness of flies homozygous for a complete chromosome relative to normal wild flies. A total of 37 chromosomes from a natural population have been tested using 92 experimental populations. The mean fitness of homozygous flies is 0.12 for second chromosomes, and 0.13 for third chromosomes. These estimates are compatible with the hypothesis that many (more than one thousand) loci are maintained by heterotic selection in natural populations of D. melanogaster.



1999 ◽  
Vol 74 (1) ◽  
pp. 31-42 ◽  
Author(s):  
J. RONFORT

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.



Genetics ◽  
2002 ◽  
Vol 160 (3) ◽  
pp. 1191-1202 ◽  
Author(s):  
Michael C Whitlock

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.



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