Manmohan D. Sharma

Male attractiveness, fertility and susceptibility to oxidative stress are influenced by inbreeding in Drosophila simulans.

Inbreeding frequently leads to inbreeding depression, a reduction in the trait values of inbred individuals. Inbreeding depression has been documented in sexually selected characters in several taxa, and while there is correlational evidence that male fertility is especially susceptible to inbreeding depression, there have been few direct experimental examinations of this. Here, we assessed inbreeding depression in male fertility and a range of other male fitness correlates in Drosophila simulans. We found that male fertility and attractiveness were especially susceptible to inbreeding depression. Additionally, levels of testicular oxidative stress were significantly elevated in inbred males, although sperm viability did not differ between inbred and outbred males. Copulation duration, induction of oviposition, and the proportion of eggs hatching did not differ for females mated to inbred or outbred males. Nevertheless, our results clearly show that key male fitness components are impaired by inbreeding and provide evidence that aspects of male fertility are especially susceptible to inbreeding depression.

ANTAGONISTIC RESPONSES TO NATURAL AND SEXUAL SELECTION AND THE SEX-SPECIFIC EVOLUTION OF CUTICULAR HYDROCARBONS IN DROSOPHILA SIMULANS

Natural and sexual selection are classically thought to oppose one another, and although there is evidence for this, direct experimental demonstrations of this antagonism are largely lacking. Here, we assessed the effects of sexual and natural selection on the evolution of cuticular hydrocarbons (CHCs), a character subject to both modes of selection, in Drosophila simulans. Natural selection and sexual selection were manipulated in a fully factorial design, and after 27 generations of experimental evolution, the responses of male and female CHCs were assessed. The effects of natural and sexual selection differed greatly across the sexes. The responses of female CHCs were generally small, but CHCs evolved predominantly in the direction of natural selection. For males, profiles evolved via sexual and natural selection, as well as through the interaction between the two, with some male CHC components only evolving in the direction of natural selection when sexual selection was relaxed. These results indicate sex‐specific responses to selection, and that sexual and natural selection act antagonistically for at least some combinations of CHCs.

Female mate preferences in Drosophila simulans: evolution and costs.

Female mate preference is central to sexual selection, and all indirect benefit models require that there is genetic variation in female preference. This has rarely been tested however, with relatively few studies documenting heritable variation in female preference and even fewer that have directly selected on mate preference to unequivocally show that it can evolve. Additionally, costs of mate preference are poorly understood even though these have implications for preference evolution. We selected on female preference for ebony‐males in replicate Drosophila simulans lines, and generated a rapid evolutionary response in both replicates, with the proportion of females mating with ebony‐males increasing from approximately 5% to 30% after five generations of selection. This increase was independent of changes in ebony‐males as only females were included in our selection regime. We could detect no cost to mate preference itself other than that associated with the fitness consequences of mating with ebony males.

Sexual and Natural Selection Both Influence Male Genital Evolution

Rapid and divergent evolution of male genital morphology is a conspicuous and general pattern across internally fertilizing animals. Rapid genital evolution is thought to be the result of sexual selection, and the role of natural selection in genital evolution remains controversial. However, natural and sexual selection are believed to act antagonistically on male genital form. We conducted an experimental evolution study to investigate the combined effects of natural and sexual selection on the genital-arch lobes of male Drosophila simulans. Replicate populations were forced to evolve under lifetime monogamy (relaxed sexual selection) or lifetime polyandry (elevated sexual selection) and two temperature regimes, 25°C (relaxed natural selection) or 27°C (elevated natural selection) in a fully factorial design. We found that natural and sexual selection plus their interaction caused genital evolution. Natural selection caused some aspects of genital form to evolve away from their sexually selected shape, whereas natural and sexual selection operated in the same direction for other shape components. Additionally, sexual and natural selection tended to favour larger genitals. Thus we find that the underlying selection driving genital evolution is complex, does not only involve sexual selection, and that natural selection and sexual selection do not always act antagonistically.

Environmental heterogeneity, multivariate sexual selection and genetic constraints on cuticular hydrocarbons in Drosophila simulans

Sexual selection is responsible for the evolution of many elaborate traits, but sexual trait evolution could be influenced by opposing natural selection as well as genetic constraints. As such, the evolution of sexual traits could depend heavily on the environment if trait expression and attractiveness vary between environments. Here, male Drosophila simulans were reared across a range of diets and temperatures, and we examined differences between these environments in terms of (i) the expression of male cuticular hydrocarbons (CHCs) and (ii) which male CHC profiles were most attractive to females. Temperature had a strong effect on male CHC expression, whereas the effect of diet was weaker. Male CHCs were subject to complex patterns of directional, quadratic and correlational sexual selection, and we found differences between environments in the combination of male CHCs that were most attractive to females, with clearer differences between diets than between temperatures. We also show that genetic covariance between environments is likely to cause a constraint on independent CHC evolution between environments. Our results demonstrate that even across the narrow range of environmental variation studied here, predicting the outcome of sexual selection can be extremely complicated, suggesting that studies ignoring multiple traits or environments may provide an over‐simplified view of the evolution of sexual traits.

Sexual conflict over mating in Gnatocerus cornutus? Females prefer lovers not fighters

Female mate choice and male–male competition are the typical mechanisms of sexual selection. However, these two mechanisms do not always favour the same males. Furthermore, it has recently become clear that female choice can sometimes benefit males that reduce female fitness. So whether male–male competition and female choice favour the same or different males, and whether or not females benefit from mate choice, remain open questions. In the horned beetle, Gnatocerus cornutus, males have enlarged mandibles used to fight rivals, and larger mandibles provide a mating advantage when there is direct male–male competition for mates. However, it is not clear whether females prefer these highly competitive males. Here, we show that female choice targets male courtship rather than mandible size, and these two characters are not phenotypically or genetically correlated. Mating with attractive, highly courting males provided indirect benefits to females but only via the heritability of male attractiveness. However, mating with attractive males avoids the indirect costs to daughters that are generated by mating with competitive males. Our results suggest that male–male competition may constrain female mate choice, possibly reducing female fitness and generating sexual conflict over mating.

The Genetics of Cuticular Hydrocarbon Profiles in the Fruit Fly Drosophila simulans

Female mate choice is one mechanism of sexual selection and, provided there is adequate genetic variation in the male traits that are the target of this selection, they will evolve via female choice. Cuticular hydrocarbons (CHCs) are important in Drosophila mate choice, but relatively little is known about the underlying genetic architecture of CHC profiles in Drosophila simulans. Here, we used gas chromatography-mass spectrometry to investigate patterns of genetic variation in the CHC profiles of male and female D. simulans using isofemale lines. We found substantial genetic variation for CHC profiles and individual CHC components, and individual CHCs were frequently strongly genetically correlated, with a tendency for negative covariance between long- and short-chain CHCs in males. Intersexual genetic covariances were often weak and frequently differed in sign. These findings are novel and significant, highlighting the previously unexplored genetic architecture of CHCs in D. simulans and suggest that this architecture may facilitate sex-specific CHC evolution.

Local adaptation of a bacterium is as important as its presence in structuring a natural microbial community

Local adaptation of a species can affect community composition, yet the importance of local adaptation compared with species presence per se is unknown. Here we determine how a compost bacterial community exposed to elevated temperature changes over 2 months as a result of the presence of a focal bacterium, Pseudomonas fluorescens SBW25, that had been pre-adapted or not to the compost for 48 days. The effect of local adaptation on community composition is as great as the effect of species presence per se, with these results robust to the presence of an additional strong selection pressure: an SBW25-specific virus. These findings suggest that evolution occurring over ecological time scales can be a key driver of the structure of natural microbial communities, particularly in situations where some species have an evolutionary head start following large perturbations, such as exposure to antibiotics or crop planting and harvesting.

Longevity, calling effort, and metabolic rate in two populations of cricket

Intraspecific variation in a resting metabolic rate (RMR) is likely to be an important determinant of energetic-resource use and may influence the resources subsequently available for allocation to traits not directly associated with somatic maintenance. The influence of RMR on resource availability could be especially important for condition-dependent sexual traits, such as cricket calls, that are themselves energetically costly to produce. RMR may also be associated with longevity, either negatively because individuals with a high RMR burn resources faster and die young, or positively as individuals with high RMR are more able to accrue resources to fuel survival. Additionally, the associations between RMR and other characters may vary across populations if differential selection or drift shapes these traits. Here we tested for differences in RMR, body mass, calling effort, and longevity in two populations of cricket Gryllodes sigillatus and then evaluated the potential influence of RMR on calling and longevity. We find that RMR, calling effort, and longevity varied across populations, but mass did not. Controlling for population and mass, RMR was not significantly associated with calling effort, but was negatively associated with longevity. These findings suggest that male crickets that live fast die young.

Sexual selection in flies: a comparison of Drosophila simulans and D. melanogaster

The traditional view of sexual selection via female mate choice is that female preference for certain males either has no net fitness cost or is beneficial to overall female fitness. A more contemporary view is that preferred males can at times reduce female fitness. This view has arisen from the realisation that conflict between the sexes is an inevitable feature of sexual reproduction, as each sex necessarily has a different agenda for maximizing fitness. Despite the hailing of sexual conflict as a paradigm shift and its prevalence in the recent sexual selection literature, compelling evidence that attractive males reduce female fitness remains taxonomically restricted. Here we review the findings of a series of investigations into the fitness consequences of female preference in the fly Drosophila simulans and compare them with its sibling species, D. melanogaster. We show that there are stark differences in the fitness consequences of mating with preferred males in the two species and discuss this contrast with reference to the current debates in the sexual selection literature.