Genetic Variation For Fitness Research Articles

Mapping the Genetic Basis of Symbiotic Variation in Legume-Rhizobium Interactions inMedicago truncatula

Mutualisms are known to be genetically variable, where the genotypes differ in the fitness benefits they gain from the interaction. To date, little is known about the loci that underlie such genetic variation in fitness or whether the loci influencing fitness are partner specific, and depend on the genotype of the interaction partner. In the legume-rhizobium mutualism, one set of potential candidate genes that may influence the fitness benefits of the symbiosis are the plant genes involved in the initiation of the signaling pathway between the two partners. Here we performed quantitative trait loci (QTL) mapping in Medicago truncatula in two different rhizobium strain treatments to locate regions of the genome influencing plant traits, assess whether such regions are dependent on the genotype of the rhizobial mutualist (QTL × rhizobium strain), and evaluate the contribution of sequence variation at known symbiosis signaling genes. Two of the symbiotic signaling genes, NFP and DMI3, colocalized with two QTL affecting average fruit weight and leaf number, suggesting that natural variation in nodulation genes may potentially influence plant fitness. In both rhizobium strain treatments, there were QTL that influenced multiple traits, indicative of either tight linkage between loci or pleiotropy, including one QTL with opposing effects on growth and reproduction. There was no evidence for QTL × rhizobium strain or genotype × genotype interactions, suggesting either that such interactions are due to small-effect loci or that more genotype-genotype combinations need to be tested in future mapping studies.

The effects of reproductive specialization on energy costs and fitness genetic variances in cyclical and obligate parthenogenetic aphids.

Organisms with coexisting sexual and asexual populations are ideal models for studying the consequences of either reproductive mode on the quantitative genetic architecture of life-history traits. In the aphid Rhopalosiphum padi, lineages differing in their sex investment coexist but all share a common parthenogenetic phase. Here, we studied multiple genotypes of R. padi specialized either for sexual and asexual reproduction and compared their genetic variation in fitness during the parthenogenetic phase. Specifically, we estimated maintenance costs as standard metabolic rate (SMR), together with fitness (measured as the intrinsic rate of increase and the net reproductive rate). We found that genetic variation (in terms of broad-sense heritability) in fitness was higher in asexual genotypes compared with sexual genotypes. Also, we found that asexual genotypes exhibited several positive genetic correlations indicating that body mass, whole-animal SMR, and apterous individuals production are contributing to fitness. Hence, it appears that in asexual genotypes, energy is fully allocated to maximize the production of parthenogenetic individuals, the simplest possible form of aphid repertoire of life-histories strategies.

FISHER'S GEOMETRICAL MODEL OF FITNESS LANDSCAPE AND VARIANCE IN FITNESS WITHIN A CHANGING ENVIRONMENT

The fitness of an individual can be simply defined as the number of its offspring in the next generation. However, it is not well understood how selection on the phenotype determines fitness. In accordance with Fisher's fundamental theorem, fitness should have no or very little genetic variance, whereas empirical data suggest that is not the case. To bridge these knowledge gaps, we follow Fisher's geometrical model and assume that fitness is determined by multivariate stabilizing selection toward an optimum that may vary among generations. We assume random mating, free recombination, additive genes, and uncorrelated stabilizing selection and mutational effects on traits. In a constant environment, we find that genetic variance in fitness under mutation-selection balance is a U-shaped function of the number of traits (i.e., of the so-called "organismal complexity"). Because the variance can be high if the organism is of either low or high complexity, this suggests that complexity has little direct costs. Under a temporally varying optimum, genetic variance increases relative to a constant optimum and increasingly so when the mutation rate is small. Therefore, mutation and changing environment together can maintain high genetic variance. These results therefore lend support to Fisher's geometric model of a fitness landscape.

Effect of Host Species on the Distribution of Mutational Fitness Effects for an RNA Virus

Knowledge about the distribution of mutational fitness effects (DMFE) is essential for many evolutionary models. In recent years, the properties of the DMFE have been carefully described for some microorganisms. In most cases, however, this information has been obtained only for a single environment, and very few studies have explored the effect that environmental variation may have on the DMFE. Environmental effects are particularly relevant for the evolution of multi-host parasites and thus for the emergence of new pathogens. Here we characterize the DMFE for a collection of twenty single-nucleotide substitution mutants of Tobacco etch potyvirus (TEV) across a set of eight host environments. Five of these host species were naturally infected by TEV, all belonging to family Solanaceae, whereas the other three were partially susceptible hosts belonging to three other plant families. First, we found a significant virus genotype-by-host species interaction, which was sustained by differences in genetic variance for fitness and the pleiotropic effect of mutations among hosts. Second, we found that the DMFEs were markedly different between Solanaceae and non-Solanaceae hosts. Exposure of TEV genotypes to non-Solanaceae hosts led to a large reduction of mean viral fitness, while the variance remained constant and skewness increased towards the right tail. Within the Solanaceae hosts, the distribution contained an excess of deleterious mutations, whereas for the non-Solanaceae the fraction of beneficial mutations was significantly larger. All together, this result suggests that TEV may easily broaden its host range and improve fitness in new hosts, and that knowledge about the DMFE in the natural host does not allow for making predictions about its properties in an alternative host.

Clinical Study on the Efficacy of Fluvoxamine for Psychological Distress in Gynecologic Cancer Patients

Diagnosis of cancer causes psychological distress. The present study investigated the safety and efficacy of fluvoxamine therapy in gynecologic cancer patients with either adjustment disorder or major depression after cancer was diagnosed. Screening with the Hospital Anxiety and Depression Scale (HADS) was conducted at least 2 weeks after notification of the diagnosis of cancer in 214 gynecologic cancer patients hospitalized between January 2007 and December 2008. The HADS cutoff score was set at 11 points or greater. Informed consent to the study was obtained from 10 patients, and fluvoxamine was administered for 8 weeks. As primary end points, the safety and efficacy of fluvoxamine were evaluated using the HADS and the SF-36. As a secondary end point, the Clinical Global Impression was determined. The total HADS score, the anxiety score, and the depression score were significantly reduced after 6, 4, and 6 weeks of treatment, respectively. The SF-36 revealed significant improvement in vitality, mental health, and role (emotional) after 8 weeks of treatment. In the 5 patients with adjustment disorder, only the HADS anxiety score was significantly reduced after 4 weeks. In the 5 patients with major depression, the total HADS score, the anxiety score, and the depression score were significantly reduced after 6, 8, and 6 weeks, respectively. According to the SF-36, the adjustment-disorder groups showed significant improvement in mental health after 8 weeks of treatment, whereas the major-depression group showed significant improvement in vitality and role (emotional) after 8 weeks. No adverse events occurred in any subject. Assessment of the Clinical Global Impression suggested that fluvoxamine improved psychological distress in all 10 subjects. The present findings suggest that fluvoxamine is useful for alleviating psychological distress, including adjustment disorder and major depression, in gynecologic cancer patients. Management of psychological distress after diagnosis of cancer is important.

EFFECTS OF FOUNDING GENETIC VARIATION ON ADAPTATION TO A NOVEL RESOURCE

Population genetic theory predicts that adaptation in novel environments is enhanced by genetic variation for fitness. However, theory also predicts that under strong selection, demographic stochasticity can drive populations to extinction before they can adapt. We exposed wheat-adapted populations of the flour beetle (Tribolium castaneum) to a novel suboptimal corn resource, to test the effects of founding genetic variation on population decline and subsequent extinction or adaptation. As previously reported, genetically diverse populations were less likely to go extinct. Here, we show that among surviving populations, genetically diverse groups recovered faster after the initial population decline. Within two years, surviving populations significantly increased their fitness on corn via increased fecundity, increased egg survival, faster larval development, and higher rate of egg cannibalism. However, founding genetic variation only enhanced the increase in fecundity, despite existing genetic variation-and apparent lack of trade-offs-for egg survival and larval development time. Thus, during adaptation to novel habitats the positive impact of genetic variation may be restricted to only a few traits, although change in many life-history traits may be necessary to avoid extinction. Despite severe initial maladaptation and low population size, genetic diversity can thus overcome the predicted high extinction risk in new habitats.

ASSORTATIVE MATING BY FITNESS AND SEXUALLY ANTAGONISTIC GENETIC VARIATION

Recent documentations of sexually antagonistic genetic variation in fitness have spurred an interest in the mechanisms that may act to maintain such variation in natural populations. Using individual-based simulations, I show that positive assortative mating by fitness increases the amount of sexually antagonistic genetic variance in fitness, primarily by elevating the equilibrium frequency of heterozygotes, over most of the range of sex-specific selection and dominance. Further, although the effects of assortative mating by fitness on the protection conditions of polymorphism in sexually antagonistic loci were relatively minor, it widens the protection conditions under most reasonable scenarios (e.g., under heterozygote superiority when fitness is averaged across the sexes) but can also somewhat narrow the protection conditions under other circumstances. The near-ubiquity of assortative mating in nature suggests that it may contribute to upholding standing sexually antagonistic genetic variation in fitness.

Additive Genetic Variance, Heritability, and Inbreeding Depression in Male Extra-Pair Reproductive Success

The hypothesis that female extra-pair reproduction in socially monogamous animals reflects indirect genetic benefits requires that there be additive and/or nonadditive genetic variance in fitness. However, the specific hypotheses that male extra-pair reproductive success (EPRS) shows additive genetic variance (V(A)), heritability (h2), or inbreeding depression, and hence that females could acquire indirect genetic benefits through increased EPRS of sons, have not been explicitly tested. We used comprehensive genetic pedigree data from song sparrows (Melospiza melodia) to estimate V(A), h2, and inbreeding depression in the number of extra-pair offspring a male sired per year and the probability that a male would sire any extra-pair offspring per year. Inbreeding depression was substantial: more inbred males sired fewer extra-pair offspring and were less likely to sire any extra-pair offspring. In contrast, estimates of V(A) and h2 were close to 0, although 95% credible intervals were relatively wide. These data suggest that females could accrue indirect genetic benefits, in terms of increased EPRS of outbred sons, by mating with unrelated social or extra-pair mates. In contrast, any indirect benefit of extra-pair reproduction in terms of producing sons with high additive genetic value for EPRS is most likely to be small.

The effect of sex on the mean and variance of fitness in facultatively sexual rotifers

The evolution of sex is a classic problem in evolutionary biology. While this topic has been the focus of much theoretical work, there is a serious dearth of empirical data. A simple yet fundamental question is how sex affects the mean and variance in fitness. Despite its importance to the theory, this type of data is available for only a handful of taxa. Here, we report two experiments in which we measure the effect of sex on the mean and variance in fitness in the monogonont rotifer, Brachionus calyciflorus. Compared to asexually derived offspring, we find that sexual offspring have lower mean fitness and less genetic variance in fitness. These results indicate that, at least in the laboratory, there are both short- and long-term disadvantages associated with sexual reproduction. We briefly review the other available data and highlight the need for future work.

Combining capture–recapture data and pedigree information to assess heritability of demographic parameters in the wild

Quantitative genetic analyses have been increasingly used to estimate the genetic basis of life-history traits in natural populations. Imperfect detection of individuals is inherent to studies that monitor populations in the wild, yet it is seldom accounted for by quantitative genetic studies, perhaps leading to flawed inference. To facilitate the inclusion of imperfect detection of individuals in such studies, we develop a method to estimate additive genetic variance and assess heritability for binary traits such as survival, using capture-recapture (CR) data. Our approach combines mixed-effects CR models with a threshold model to incorporate discrete data in a standard 'animal model' approach. We employ Markov chain Monte Carlo sampling in a Bayesian framework to estimate model parameters. We illustrate our approach using data from a wild population of blue tits (Cyanistes caeruleus) and present the first estimate of heritability of adult survival in the wild. In agreement with the prediction that selection should deplete additive genetic variance in fitness, we found that survival had low heritability. Because the detection process is incorporated, capture-recapture animal models (CRAM) provide unbiased quantitative genetics analyses of longitudinal data collected in the wild.

Intraspecific genetic variation and competition interact to influence niche expansion.

Theory and empirical evidence show that intraspecific competition can drive selection favouring the use of novel resources (i.e. niche expansion). The evolutionary response to such selection depends on genetic variation for resource use. However, while genetic variation might facilitate niche expansion, genetically diverse groups may also experience weaker competition, reducing density-dependent selection on resource use. Therefore, genetic variation for fitness on different resources could directly facilitate, or indirectly retard, niche expansion. To test these alternatives, we factorially manipulated both the degree of genetic variation and population density in flour beetles (Tribolium castaneum) exposed to both novel and familiar food resources. Using stable carbon isotope analysis, we measured temporal change and individual variation in beetle diet across eight generations. Intraspecific competition and genetic variation acted on different components of niche evolution: competition facilitated niche expansion, while genetic variation increased individual variation in niche use. In addition, genetic variation and competition together facilitated niche expansion, but all these impacts were temporally variable. Thus, we show that the interaction between genetic variation and competition can also determine niche evolution at different time scales.

The Sexually Antagonistic Genes of Drosophila melanogaster

Author SummaryMales and females of many species are different: many of these differences are thought to have evolved because the sexes often have needs and strategies that do not coincide. For example, in fruit-flies, females may do best by concentrating their efforts in acquiring resources to be able to lay more eggs, while males would benefit most from increasing their mating and fertilization success. Such differences generate a sexual “conflict of interests”, and since as a general rule each behavioural, morphological or physiological characteristic is regulated by the same set of genes in the two sexes, this conflict takes place ultimately at the genetic level. In our study, we combined data on the reproductive success of different lines of fruit-flies with their gene expression profiles. We show that a large proportion of genes that contribute to male fertilization success are detrimental for female fecundity, and vice-versa. These results indicate that an optimal genotype for both sexes does not exist: many genes maintain different variants because they have opposite effects in males and females, perhaps helping to explain how genetic diversity is maintained in the face of selection.

Identifying Genes that Help One Sex but Harm the Other

Traits that help one sex can harm the other, resulting in conflicting evolutionary pressures on males and females. In zebra finches (Taeniopygia guttata), for instance, males with redder bills attract more mates but females with redder bills attract fewer mates. This battle of the sexes is thought to extend to the genetic level, with individual genes favoring one sex over the other. Some of the strongest evidence for these sexually antagonistic genes comes from studies showing that fruit fly lines with high reproductive success in one sex typically have low reproductive success in the other. Thus, if males in a particular line have many offspring, the females do not and vice versa. The genes underlying this sexual tug-of-war, however, have been difficult to find. Now, Paolo Innocenti and Edward Morrow reveal this conflict's genetic basis by linking the expression of sexually antagonistic genes in the fruit fly Drosophila melanogaster to the reproductive success of one sex at the expense of the other. The first step was identifying fruit fly lines where males and females had differential reproductive success or fitness. The researchers tested the fitness of males and females in 100 lines of fruit flies. For each line, test males and competitor males vied for mates. The test males had the normal red eyes, whereas the competitor males and females had brown eyes. Because red eyes are dominant, the proportion of red-eyed offspring yielded the relative fitness of males in each line. Similarly, females from each line and competitor females vied for food, with the number of offspring yielding the relative fitness of females in each line. As expected, lines with high-fitness males generally had low-fitness females, whereas lines with high-fitness females generally had low-fitness males. Next, Innocenti and Morrow compared gene expression in five lines with high male and low female fitness, five lines where the reverse held, and five lines where the fitness of the two sexes was average. RNA analysis of the males and females in these 15 lines revealed that although most gene expression was sex-biased, only a minority of transcripts were consistently associated with male or female fitness. While about 17,000 transcripts, or about 90% of the total, were expressed at higher levels in one sex or the other, only 867 were associated with male fitness and only 634 were associated with female fitness. To identify which genes were sexually antagonistic, the researchers determined which of these transcripts were associated with fitness in one sex at the expense of the other. They found that nearly 1,500 transcripts benefitted one sex over the other, with roughly half favoring males and half favoring females. These transcripts correspond to nearly 1,300 known genes, and these sexually antagonistic genes comprise the majority (about two-thirds) of the sex-specific fitness genes. This suggests that sexually antagonistic selection maintains genetic variation for fitness. The researchers were surprised that sexually antagonistic genes accounted for so few of the sex-biased transcripts, which were expressed at higher levels in one sex over the other, but were not necessarily associated with fitness. Sex-biased gene expression has been used as a proxy for sexual antagonism, based on the assumption that these differences in expression reflect conflict at the genetic level. However, this study questions that approach: sexually antagonistic genes accounted for a mere 8% of the total sex-biased transcripts. Innocenti and Morrow also tested—and confirmed—the key prediction that sexually antagonistic genes should accumulate on sex chromosomes. Because these chromosomes are inherited differently by males and females, this should help resolve the evolutionary conflict between the sexes. X chromosomes could have a recessive “male fitness” gene, for instance, which would benefit males with that gene on their single X chromosome. But this male benefit would not necessarily come at the expense of females: they would not be harmed if this recessive gene was on only one of their X chromosomes. As expected, the researchers found that sexually antagonistic genes were overwhelmingly concentrated on the X chromosome. Previous studies suggest that sexually antagonistic genes are widespread amongst animals, from insects to birds to mammals. By offering this initial glimpse of the genetic basis of such findings, Innocenti and Morrow lay the groundwork for pinpointing the locations and functions of these genes that pit males and females against each other. Innocenti P, Morrow EH (2010) The Sexually Antagonistic Genes of Drosophila melanogaster. doi:10.1371/journal.pbio.1000335 Males and females display the yin-yang duality represented in the Taijitu (above): opposing forces (sexually antagonistic selection) tend to drive them apart, but an ontogenic constraint—the shared genetic material—prevents their disjunction. ...

Fisher’s fundamental theorem of inclusive fitness and the change in fitness due to natural selection when conspecifics interact

Competition and cooperation is fundamental to evolution by natural selection, both in animals and plants. Here, I investigate the consequences of such interactions for response in fitness due to natural selection. I provide quantitative genetic expressions for heritable variance and response in fitness due to natural selection when conspecifics interact. Results show that interactions among conspecifics generate extra heritable variance in fitness, and that interacting with kin is the key to evolutionary success because it translates the extra heritable variance into response in fitness. This work also unifies Fisher's fundamental theorem of natural selection (FTNS) and Hamilton's inclusive fitness (IF). The FTNS implies that natural selection maximizes fitness, whereas Hamilton proposed maximization of IF. This work shows that the FTNS describes the increase in IF, rather than direct fitness, at a rate equal to the additive genetic variance in fitness. Thus, Hamilton's IF and Fisher's FTNS both describe the maximization of IF.

The Good‐Genes and Compatible‐Genes Benefits of Mate Choice

Genetic benefits from mate choice could be attained by choosing mates with high heritable quality ("good genes") and that are genetically compatible ("compatible genes"). We clarify the conceptual and empirical framework for estimating genetic benefits of mate choice, stressing that benefits must be measured from offspring fitness because there are no unequivocal surrogates for genetic quality of individuals or for compatibility of parents. We detail the relationship between genetic benefits and additive and nonadditive genetic variance in fitness, showing that the benefits have been overestimated in previous verbal treatments. We point out that additive benefits readily arise from nonadditive gene action and that the idea of "heritable nonadditive benefits" is a misconception. We review the empirical evidence of the magnitude of benefits of good genes and compatible genes in animal populations, and we outline the most promising future directions for empirical research on the genetic benefits of mate choice.

Contribution of chromosomal polymorphisms to the G‐matrix ofMimulus guttatus

Evolution of genetic (co)variances (the G-matrix) fundamentally influences multitrait divergence. Here, we isolated the contribution of two chromosomal quantitative trait loci (QTLs), a meiotic drive locus and a polymorphic inversion, to the overall G-matrix for a suite of floral, phenological and male fitness traits in a population of Mimulus guttatus. This allowed us to predict the evolution of trait means and genetic (co)variances as a function of allele frequencies, and to evaluate theories about the maintenance of genetic variation in fitness. Individuals generated using a replicated F(2) breeding design were grown under common conditions, genotyped and measured for trait values. Significant additive genetic variance existed for all traits, and most genetic covariances were significantly nonzero. Both QTLs contribute to the additive genetic (co)variances of multiple traits. Pleiotropy was not generally consistent, either between QTLs or with the genetic background. Shifts in allele frequencies at either QTL are predicted to result in substantial changes in the G-matrix. Both QTLs contribute substantially to the genetic variation in pollen viability. The Drive QTL, and perhaps also the inversion, demonstrates the contribution of balancing selection to the maintenance of genetic variation in fitness.

EFFECTS OF TEMPERATURE ON THE FITNESS COST OF RESISTANCE TO BACTERIOPHAGE T4 INESCHERICHIA COLI.

Resistance to predation, herbivory, or disease often comes at a cost such that resistant genotypes are competitively inferior to their sensitive counterparts in the absence of predators, herbivores, or pathogens. The effects of this trade-off on natural populations depend on its sensitivity to environmental changes. We used Escherichia coli and bacteriophage T4 as a model predator/prey system to study the effects of temperature on the cost of resistance. An array of independent T4-resistant mutants, derived from a single ancestral strain of E. coli B, had a mean reduction in competitive fitness that depended strongly on environmental temperature; the cost of resistance generally increased with temperature. Genetic variance for fitness among phage-resistant mutants also depended on temperature; however, genetic variance increased at high and low thermal extremes. These results suggest that temperature is likely to be an important determinant of the consequences of predation in natural communities. We also discuss the underlying mechanistic basis for the cost of resistance in this system and its interaction with temperature.

Mate choice for nonadditive genetic benefits and the maintenance of genetic diversity in song sparrows

The lek paradox asserts that strong directional selection via female choice should deplete additive genetic variation in fitness and consequently any benefit to females expressing the preference. Recently, we have provided a novel resolution to the paradox by showing that nonadditive genetic effects such as overdominance can be inherited from parent to offspring, and populations with females that express a mating preference for outbred males maintain higher genetic variation than populations with females that mate randomly. Here, we test our dynamic model using empirical data previously published from a small island population of song sparrows (Melospiza melodia). The model assumes that fitness and male trait expression display overdominance effects. The results demonstrate that female choice for outbred males mediated by directional selection on song repertoire size provides a heritable benefit to offspring through reduced inbreeding depression. Within the population, we estimate the heritability of the inbreeding coefficient to be 0.18 +/- 0.08 (SD). Furthermore, we show that mate choice for outbred males increases fitness-related genetic variation in the population by 12% and thereby reduces inbreeding depression by 1% per generation in typical years and upwards of 15% in severe years. Thus, mate choice may help to stave off population extinction in this and other small populations.

Unifying Life‐History Analyses for Inference of Fitness and Population Growth

The lifetime fitnesses of individuals comprising a population determine its numerical dynamics, and genetic variation in fitness results in evolutionary change. This dual importance of individual fitness is well understood, but empirical fitness records generally violate the assumptions of standard statistical approaches. This problem has undermined comprehensive study of fitness and impeded empirical synthesis of the numerical and genetic dynamics of populations. Recently developed aster models remedy this problem by explicitly modeling the dependence of later-expressed components of fitness (e.g., fecundity) on those expressed earlier (e.g., survival to reproduce). Moreover, aster models employ different sampling distributions for different components of fitness (e.g., binomial for survival over a given interval and Poisson for fecundity). Analysis is done by maximum likelihood, and the resulting distributions for lifetime fitness closely approximate observed data. We illustrate the breadth of aster models' utility with three examples demonstrating estimation of the finite rate of increase, comparison of mean fitness among genotypic groups, and analysis of phenotypic selection. Aster models offer a unified approach to addressing the breadth of questions in evolution and ecology for which life-history data are gathered.

Mate choice for non-additive genetic benefits: A resolution to the lek paradox

In promiscuous mating systems, females often show a consistent preference to mate with one or a few males, presumably to acquire heritable genetic benefits for their offspring. However, strong directional selection should deplete additive genetic variation in fitness and consequently any benefit to expressing the preference by females (referred to as the lek paradox). Here, we provide a novel resolution that examines non-additive genetic benefits, such as overdominance or inbreeding, as a source of genetic variation. Focusing on the inbreeding coefficient f and overdominance effects, we use dynamic models to show that (1) f can be inherited from sire to offspring, (2) populations with females that express a mating preferences for outbred males (low f) maintain higher genetic variation than populations with females that mate randomly, and (3) preference alleles for outbred males can invade populations even when the alleles are associated with a fecundity cost. We show that non-additive genetic variation due to overdominance can be converted to additive genetic variation and becomes “heritable” when the frequencies of alternative homozygous genotypes at fitness loci deviate from equality. Unlike previous models that assume an infinite population size, we now show that genetic drift in finite populations can lead to the necessary deviations in the frequencies of homozygous genotypes. We also show that the “heritability of f,” and hence the benefit to a mating preference for non-additive genetic benefits, is highest in small populations and populations in which a smaller number of loci contribute to fitness via overdominance. Our model contributes to the solution of the lek paradox.