scholarly journals An experimental test of the mutation-selection balance model for the maintenance of genetic variance in fitness components

2017 ◽  
Author(s):  
Nathaniel P. Sharp ◽  
Aneil F. Agrawal
Keyword(s):  
Genetic Variance ◽  
Balancing Selection ◽  
Statistical Tests ◽  
Balance Model ◽  
Male Mating ◽  
Deleterious Mutations ◽  
Fitness Component ◽  
Fitness Components ◽  
Female Fecundity ◽  
Mutational Variance

ABSTRACTDespite decades of research, the factors that maintain genetic variation for fitness are poorly understood. Mutation selection balance will always contribute to standing variance, but it is unclear what fraction of the variance in a typical fitness component can be explained by mutation-selection balance and whether fitness components differ in this respect. In theory, the level of standing variance in fitness due to mutation-selection balance can be predicted using the rate of fitness decline under mutation accumulation, and this prediction can be directly compared to the actual standing variance observed. This approach allows for controlled statistical tests of the sufficiency of the mutation-selection balance model, and could be used to identify traits or populations where genetic variance is maintained by factors beyond mutation-selection balance. For example, some traits may be influenced by sexually antagonistic balancing selection, resulting in an excess of standing variance beyond that generated by deleterious mutations. To encourage the application of this approach, we describe the underlying theory and use it to test the mutation selection balance model for three traits in Drosophila melanogaster. We find some evidence for non-mutational variance in male mating success and female fecundity relative to larval viability, which is consistent with balancing selection on sexual fitness components. Finally, we discuss the theoretical and practical limitations to this approach, and discuss how to apply it successfully.

scholarly journals An experimental test of the mutation-selection balance model for the maintenance of genetic variance in fitness components

2018 ◽  
Vol 285 (1890) ◽  
pp. 20181864 ◽  
Author(s):  
Nathaniel P. Sharp ◽  
Aneil F. Agrawal
Keyword(s):  
Genetic Variance ◽  
Balancing Selection ◽  
Statistical Tests ◽  
Balance Model ◽  
Male Mating ◽  
Male Mating Success ◽  
Deleterious Mutations ◽  
Fitness Component ◽  
Fitness Components ◽  
Female Fecundity

Despite decades of research, the factors that maintain genetic variation for fitness are poorly understood. It is unclear what fraction of the variance in a typical fitness component can be explained by mutation-selection balance (MSB) and whether fitness components differ in this respect. In theory, the level of standing variance in fitness due to MSB can be predicted using the rate of fitness decline under mutation accumulation, and this prediction can be directly compared to the standing variance observed. This approach allows for controlled statistical tests of the sufficiency of the MSB model, and could be used to identify traits or populations where genetic variance is maintained by other factors. For example, some traits may be influenced by sexually antagonistic balancing selection, resulting in an excess of standing variance beyond that generated by deleterious mutations. We describe the underlying theory and use it to test the MSB model for three traits in Drosophila melanogaster . We find evidence for differences among traits, with MSB being sufficient to explain genetic variance in larval viability but not male mating success or female fecundity. Our results are consistent with balancing selection on sexual fitness components, and demonstrate the feasibility of rigorous statistical tests of the MSB model.

scholarly journals Causes of natural variation in fitness: Evidence from studies of Drosophila populations

2015 ◽  
Vol 112 (6) ◽  
pp. 1662-1669 ◽  
Author(s):  
Brian Charlesworth
Keyword(s):  
Population Genetics ◽  
Genetic Variation ◽  
Balancing Selection ◽  
Large Contribution ◽  
Deleterious Mutations ◽  
Weak Selection ◽  
Fitness Components ◽  
Genetic Studies ◽  
Standing Variation ◽  
Available Information

DNA sequencing has revealed high levels of variability within most species. Statistical methods based on population genetics theory have been applied to the resulting data and suggest that most mutations affecting functionally important sequences are deleterious but subject to very weak selection. Quantitative genetic studies have provided information on the extent of genetic variation within populations in traits related to fitness and the rate at which variability in these traits arises by mutation. This paper attempts to combine the available information from applications of the two approaches to populations of the fruitfly Drosophila in order to estimate some important parameters of genetic variation, using a simple population genetics model of mutational effects on fitness components. Analyses based on this model suggest the existence of a class of mutations with much larger fitness effects than those inferred from sequence variability and that contribute most of the standing variation in fitness within a population caused by the input of mildly deleterious mutations. However, deleterious mutations explain only part of this standing variation, and other processes such as balancing selection appear to make a large contribution to genetic variation in fitness components in Drosophila.

scholarly journals Fitness of allozyme variants in Drosophila pseudoobscura: II. Selection at the Est-5, Odh and Mdh-2 loci

1974 ◽  
Vol 24 (2) ◽  
pp. 137-149 ◽  
Author(s):  
Dragoslav Marinković ◽  
Francisco J. Ayala
Keyword(s):  
Allelic Variation ◽  
Balancing Selection ◽  
Natural Populations ◽  
Superior Performance ◽  
Adult Survival ◽  
Drosophila Pseudoobscura ◽  
Male Mating ◽  
Female Fecundity ◽  
Single Genotype ◽  
Fitness Parameters

SUMMARYWe have studied the effects on fitness of allelic variation at three gene loci (Est-5, Odh, and Mdh-2)coding for enzymes in Drosophila pseudoobscura. Genotype has a significant effect on fitness for all six parameters measured (female fecundity, male mating capacity, egg-to-adult survival under near-optimal and under competitive conditions, and rate of development under near-optimal and under competitive conditions). No single genotype is best for all six fitness parameters; rather, genotypes with superior performance during a certain stage of the life-cycle may have low fitness at some other stage, or in different environmental conditions. Heterozygotes are sometimes best when all fitness parameters are considered. There are significant interactions between loci. The various forms of balancing selection uncovered in our experiments may account for the polymorphisms occurring in natural populations of D. pseudoobscura at the three loci studied.

scholarly journals The genetic basis of inbreeding depression

1999 ◽  
Vol 74 (3) ◽  
pp. 329-340 ◽  
Author(s):  
BRIAN CHARLESWORTH ◽  
DEBORAH CHARLESWORTH
Keyword(s):  
Inbreeding Depression ◽  
Genetic Basis ◽  
Significant Contribution ◽  
Balancing Selection ◽  
Genetic Data ◽  
Deleterious Mutations ◽  
Fitness Components ◽  
Mutation Pressure ◽  
Quantitative Genetic ◽  
Experimental Approaches

Data on the effects of inbreeding on fitness components are reviewed in the light of population genetic models of the possible genetic causes of inbreeding depression. Deleterious mutations probably play a major role in causing inbreeding depression. Putting together the different kinds of quantitative genetic data, it is difficult to account for the very large effects of inbreeding on fitness in Drosophila and outcrossing plants without a significant contribution from variability maintained by selection. Overdominant effects of alleles on fitness components seem not to be important in most cases. Recessive or partially recessive deleterious effects of alleles, some maintained by mutation pressure and some by balancing selection, thus seem to be the most important source of inbreeding depression. Possible experimental approaches to resolving outstanding questions are discussed.

scholarly journals SELECTION BY FERTILITY IN DROSOPHILA PSEUDOOBSCURA

Genetics ◽  
1974 ◽  
Vol 77 (3) ◽  
pp. 559-564
Author(s):  
Wyatt W Anderson ◽  
Takao K Watanabe
Keyword(s):  
Balancing Selection ◽  
Low Frequency ◽  
Chromosomal Polymorphism ◽  
Intermediate Frequency ◽  
Male Mating ◽  
Female Fecundity ◽  
Differential Fertility ◽  
High Frequencies ◽  
Differential Viability ◽  
Males And Females

ABSTRACT Fertility, the component of selection due to female fecundity and male mating success, differed significantly among the ST/ST, ST/AR, and AR/AR karyotypes in experimental populations and varied with karyotypic frequency. In relation to ST/AR, ST/ST females and males had higher fertilities at low frequency; AR/AR males and females were at a significant fertility disadvantage at intermediate frequency, while at low and at high frequencies their fertilities matched or exceeded that of the heterokaryotype. These fertility differences were comparable in size to viability differences previously reported for D. pseudoobscura karyotypes. Differential fertility seems likely to be an important element, perhaps just as important as differential viability, in the balancing selection that maintains the chromosomal polymorphism in this species.

scholarly journals Characterization of Deleterious Mutations in Outcrossing Populations

Genetics ◽  
1998 ◽  
Vol 150 (2) ◽  
pp. 945-956 ◽  
Author(s):  
Hong-Wen Deng
Keyword(s):  
Genetic Variation ◽  
Genetic Variance ◽  
Estimation Bias ◽  
Deleterious Mutations ◽  
Environmental Variance ◽  
Higher Power ◽  
Sib Mating ◽  
The Mean ◽  
Simulation Results

Abstract Deng and Lynch recently proposed estimating the rate and effects of deleterious genomic mutations from changes in the mean and genetic variance of fitness upon selfing/outcrossing in outcrossing/highly selfing populations. The utility of our original estimation approach is limited in outcrossing populations, since selfing may not always be feasible. Here we extend the approach to any form of inbreeding in outcrossing populations. By simulations, the statistical properties of the estimation under a common form of inbreeding (sib mating) are investigated under a range of biologically plausible situations. The efficiencies of different degrees of inbreeding and two different experimental designs of estimation are also investigated. We found that estimation using the total genetic variation in the inbred generation is generally more efficient than employing the genetic variation among the mean of inbred families, and that higher degree of inbreeding employed in experiments yields higher power for estimation. The simulation results of the magnitude and direction of estimation bias under variable or epistatic mutation effects may provide a basis for accurate inferences of deleterious mutations. Simulations accounting for environmental variance of fitness suggest that, under full-sib mating, our extension can achieve reasonably well an estimation with sample sizes of only ∼2000-3000.

Comparing Mutational Variabilities

Genetics ◽  
1996 ◽  
Vol 143 (3) ◽  
pp. 1467-1483 ◽  
Author(s):  
David Houle ◽  
Bob Morikawa ◽  
Michael Lynch
Keyword(s):  
Life History ◽  
Morphological Traits ◽  
Life History Traits ◽  
Genetic Variance ◽  
Balance Model ◽  
Phenotypic Traits ◽  
Infinite Population ◽  
Persistence Time ◽  
Coefficients Of Variation ◽  
Balance Hypothesis

Abstract We have reviewed the available data on VM, the amount of genetic variation in phenotypic traits produced each generation by mutation. We use these data to make several qualitative tests of the mutation-selection balance hypothesis for the maintenance of genetic variance (MSB). To compare VM values, we use three dimensionless quantities: mutational heritability, the mutational coefficient of variation, CVM; and the ratio of the standing genetic variance to VM, VG/VM. Since genetic coefficients of variation for life history traits are larger than those for morphological traits, we predict that under MSB, life history traits should also have larger CVM. This is confirmed; life history traits have a median CVM value more than six times higher than that for morphological traits. VG/VM approximates the persistence time of mutations under MSB in an infinite population. In order for MSB to hold, VG/VM must be small, substantially less than 1000, and life history traits should have smaller values than morphological traits. VG/VM averages about 50 generations for life history traits and 100 generations for morphological traits. These observations are all consistent with the predictions of a mutation-selection balance model.

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.

Supergene degeneration opposes polymorphism: The curious case of balanced lethals

2021 ◽  
Author(s):  
Emma Berdan ◽  
Alexandre Blanckaert ◽  
Roger K Butlin ◽  
Thomas Flatt ◽  
Tanja Slotte ◽  
...  
Keyword(s):  
Equilibrium State ◽  
Balancing Selection ◽  
Mutation Accumulation ◽  
Small Population ◽  
Quasi Equilibrium ◽  
Deleterious Mutations ◽  
Critical Factors ◽  
Critical Aspect ◽  
Allopatric Divergence

Supergenes offer some of the most spectacular examples of long-term balancing selection in nature but their origin and maintenance remain a mystery. A critical aspect of supergenes is reduced recombination between arrangements. Reduced recombination protects adaptive multi-trait phenotypes, but can also lead to degeneration through mutation accumulation. Mutation accumulation can stabilize the system through the emergence of associative overdominance (AOD), destabilize the system, or lead to new evolutionary outcomes. One such outcome is the formation of balanced lethal systems, a maladaptive system where both supergene arrangements have accumulated deleterious mutations to the extent that both homozygotes are inviable, leaving only heterozygotes to reproduce. Here, we perform a simulation study to understand the conditions under which these different outcomes occur, assuming a scenario of introgression after allopatric divergence. We found that AOD aids the invasion of a new supergene arrangement and the establishment of a polymorphism. However, this polymorphism is easily destabilized by further mutation accumulation. While degradation may strengthen AOD, thereby stabilizing the supergene polymorphism, it is often asymmetric, which is the key disrupter of the quasi-equilibrium state of the polymorphism. Furthermore, mechanisms that accelerate degeneration also tend to amplify asymmetric mutation accumulation between the supergene arrangements and vice versa. As the evolution of a balanced lethal system requires symmetric degradation of both arrangements, this leaves highly restricted conditions under which such a system could evolve. We show that small population size and low dominance coefficients are critical factors, as these reduce the efficacy of selection. The dichotomy between the persistence of a polymorphism and degradation of supergene arrangements likely underlies the rarity of balanced lethal systems in nature.

scholarly journals Do Downscaled General Circulation Models Reliably Simulate Historical Climatic Conditions?

2018 ◽  
Vol 22 (10) ◽  
pp. 1-22 ◽  
Author(s):  
Andrew R. Bock ◽  
Lauren E. Hay ◽  
Gregory J. McCabe ◽  
Steven L. Markstrom ◽  
R. Dwight Atkinson
Keyword(s):  
General Circulation ◽  
Statistical Tests ◽  
Circulation Model ◽  
General Circulation Models ◽  
Climatic Conditions ◽  
Low Flow ◽  
Balance Model ◽  
Monthly Precipitation ◽  
Significance Level ◽  
Ks Test

Abstract The accuracy of statistically downscaled (SD) general circulation model (GCM) simulations of monthly surface climate for historical conditions (1950–2005) was assessed for the conterminous United States (CONUS). The SD monthly precipitation (PPT) and temperature (TAVE) from 95 GCMs from phases 3 and 5 of the Coupled Model Intercomparison Project (CMIP3 and CMIP5) were used as inputs to a monthly water balance model (MWBM). Distributions of MWBM input (PPT and TAVE) and output [runoff (RUN)] variables derived from gridded station data (GSD) and historical SD climate were compared using the Kolmogorov–Smirnov (KS) test For all three variables considered, the KS test results showed that variables simulated using CMIP5 generally are more reliable than those derived from CMIP3, likely due to improvements in PPT simulations. At most locations across the CONUS, the largest differences between GSD and SD PPT and RUN occurred in the lowest part of the distributions (i.e., low-flow RUN and low-magnitude PPT). Results indicate that for the majority of the CONUS, there are downscaled GCMs that can reliably simulate historical climatic conditions. But, in some geographic locations, none of the SD GCMs replicated historical conditions for two of the three variables (PPT and RUN) based on the KS test, with a significance level of 0.05. In these locations, improved GCM simulations of PPT are needed to more reliably estimate components of the hydrologic cycle. Simple metrics and statistical tests, such as those described here, can provide an initial set of criteria to help simplify GCM selection.

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