Polyandrous females avoid costs of inbreeding

Why do females typically mate with more than one male? Female mating patterns have broad implications for sexual selection, speciation and conflicts of interest between the sexes, and yet they are poorly understood. Matings inevitably have costs, and for females, the benefits of taking more than one mate are rarely obvious. One possible explanation is that females gain benefits because they can avoid using sperm from genetically incompatible males, or invest less in the offspring of such males. It has been shown that mating with more than one male can increase offspring viability, but we present the first clear demonstration that this occurs because females with several mates avoid the negative effects of genetic incompatibility. We show that in crickets, the eggs of females that mate only with siblings have decreased hatching success. However, if females mate with both a sibling and a non-sibling they avoid altogether the low egg viability associated with sibling matings. If similar effects occur in other species, inbreeding avoidance may be important in understanding the prevalence of multiple mating

Experimental evolution reveals trade-offs between mating and immunity

This is the final version of the article. Available from the publisher via the DOI in this record.Immune system maintenance and upregulation is costly. Sexual selection intensity, which increases male investment into reproductive traits, is expected to create trade-offs with immune function. We assayed phenoloxidase (PO) and lytic activity of individuals from populations of the Indian meal moth, Plodia interpunctella, which had been evolving under different intensities of sexual selection. We found significant divergence among populations, with males from female-biased populations having lower PO activity than males from balanced sex ratio or male-biased populations. There was no divergence in anti-bacterial lytic activity. Our data suggest that it is the increased male mating demands in female-biased populations that trades-off against immunity, and not the increased investment in sperm transfer per mating that characterizes male-biased populations.The study was supported by the Australian Research Council (K.B.M. and L.W.S.) and a Royal Society Wolfson Research Merit Award (N.W.). We thank Sheridan Willis and Michelle Hares for assistance

Experimental evolution reveals divergence in female genital teeth morphology in response to sexual conflict intensity in a moth

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this record The rapid evolutionary divergence of male genital structures under sexual selection is well documented. However, variation in female genital traits and the potential for sexual conflict to drive the coevolution between male and female traits has only recently received attention. In many lepidopterans females possess genital teeth (collectively, signa). Comparative studies suggest these teeth, involved in the deflation of spermatophores, may have coevolved with male spermatophore thickness via sexually antagonistic coevolution in a contest over the rate of deflation of spermatophores within the reproductive tract. We tested the hypothesis that sexual conflict should generate coevolution between genital teeth and spermatophore morphology by examining these traits under experimental manipulation of sexual conflict intensity. Using micro-CT scanning, we examined spermatophore and teeth morphology in populations of the Indian moth, Plodia interpunctella, which had been evolving for 110 generations under different adult sex-ratio biases. We found divergence in female signa morphology in response to sexual conflict: females from female-biased populations (reduced sexual conflict) developed wider signa. However, we found no evidence of coevolution between signa traits and spermatophore thickness as reported from comparative studies. This article is protected by copyright. All rights reserved.Australian Research CouncilRoyal Societ

Good Genes and Sexual Selection in Dung Beetles (Onthophagus taurus): Genetic Variance in Egg-to-Adult and Adult Viability

Whether species exhibit significant heritable variation in fitness is central for sexual selection. According to good genes models there must be genetic variation in males leading to variation in offspring fitness if females are to obtain genetic benefits from exercising mate preferences, or by mating multiply. However, sexual selection based on genetic benefits is controversial, and there is limited unambiguous support for the notion that choosy or polyandrous females can increase the chances of producing offspring with high viability. Here we examine the levels of additive genetic variance in two fitness components in the dung beetle Onthophagus taurus. We found significant sire effects on egg-to-adult viability and on son, but not daughter, survival to sexual maturity, as well as moderate coefficients of additive variance in these traits. Moreover, we do not find evidence for sexual antagonism influencing genetic variation for fitness. Our results are consistent with good genes sexual selection, and suggest that both pre- and postcopulatory mate choice, and male competition could provide indirect benefits to females

Sexual conflict over remating interval is modulated by the sex peptide pathway

Sexual conflict, in which the evolutionary interests of males and females diverge, shapes the evolution of reproductive systems across diverse taxa. Here we used the fruit fly to study sexual conflict in natural, three-way interactions comprising a female, her current and previous mates. We manipulated the potential for sexual conflict by using sex peptide receptor (SPR) null females and by varying remating from 3 to 48h, a period during which natural rematings frequently occur. SPR-lacking females do not respond to sex peptide transferred during mating and maintain virgin levels of high receptivity and low fecundity. In the absence of SPR there was a convergence of fitness interests, with all individuals gaining highest productivity at 5h remating. This suggests that the expression of sexual conflict was reduced. We observed an unexpected second male-specific advantage to early remating, resulting from an increase in the efficiency of second male sperm use. This early window of opportunity for exploitation by second males depended on the presence of SPR. The results suggest that the sex peptide pathway can modulate the expression of sexual conflict in this system, and show how variation in the selective forces that shape conflict and co-operation can be maintained

Evolutionary insight from a humble fly: Sperm competition and the yellow dungfly

This is the author accepted manuscript. The final version is available from the Royal Society via the DOI in this recordData accessibility: This article has no additional data.Studies of the yellow dungfly in the 1960s provided one of the first quantitative demonstrations of the costs and benefits associated with male and female reproductive behaviour. These studies advanced appreciation of sexual selection as a significant evolutionary mechanism and contributed to the 1970s paradigm shift toward individual selectionist thinking. Three behaviours in particular led to the realization that sexual selection can continue during and after mating: (i) female receptivity to remating, (ii) sperm displacement and (iii) post-copulatory mate guarding. These behaviours either generate, or are adaptations to sperm competition, cryptic female choice and sexual conflict. Here we review this body of work, and its contribution to the development of post-copulatory sexual selection theory. This article is part of the theme issue 'Fifty years of sperm competition'

On staying grounded and avoiding Quixotic dead ends

The 15 articles in this special issue on The Representation of Concepts illustrate the rich variety of theoretical positions and supporting research that characterize the area. Although much agreement exists among contributors, much disagreement exists as well, especially about the roles of grounding and abstraction in conceptual processing. I first review theoretical approaches raised in these articles that I believe are Quixotic dead ends, namely, approaches that are principled and inspired but likely to fail. In the process, I review various theories of amodal symbols, their distortions of grounded theories, and fallacies in the evidence used to support them. Incorporating further contributions across articles, I then sketch a theoretical approach that I believe is likely to be successful, which includes grounding, abstraction, flexibility, explaining classic conceptual phenomena, and making contact with real-world situations. This account further proposes that (1) a key element of grounding is neural reuse, (2) abstraction takes the forms of multimodal compression, distilled abstraction, and distributed linguistic representation (but not amodal symbols), and (3) flexible context-dependent representations are a hallmark of conceptual processing

Ejaculate Economics: Testing the Effects of Male Sexual History on the Trade-Off between Sperm and Immune Function in Australian Crickets

Trade-offs between investment into male sexual traits and immune function provide the foundation for some of the most prominent models of sexual selection. Post-copulatory sexual selection on the male ejaculate is intense, and therefore trade-offs should occur between investment into the ejaculate and the immune system. Examples of such trade-offs exist, including that between sperm quality and immunity in the Australian cricket, Teleogryllus oceanicus. Here, we explore the dynamics of this trade-off, examining the effects that increased levels of sexual interaction have on the viability of a male's sperm across time, and the concomitant effects on immune function. Males were assigned to a treatment, whereby they cohabited with females that were sexually immature, sexually mature but incapable of copulation, or sexually mature and capable of copulation. Sperm viability of each male was then assessed at two time points: six and 13 days into the treatment, and immune function at day 13. Sperm viability decreased across the time points, but only for males exposed to treatment classes involving sexually mature females. This decrease was similar in magnitude across both sexually mature classes, indicating that costs to the expression of high sperm viability are incurred largely through levels of pre-copulatory investment. Males exposed to immature females produced sperm of low viability at both time points. Although we confirmed a weak negative association between sperm viability and lytic activity (a measure of immune response to bacterial infection) at day 13, this relationship was not altered across the mating treatment. Our results highlight that sperm viability is a labile trait, costly to produce, and subject to strategic allocation in these crickets

Male-female interactions drive the (un)repeatability of copula duration in an insect.

Across the animal kingdom the duration of copulation varies enormously from a few seconds to several days. Functional explanations for this variation are largely embedded within sperm competition theory in which males modulate the duration of copula in order to optimise their fitness. However, copulation is the union of two protagonists which are likely to have separate and often conflicting reproductive interests, yet few experimental designs specifically assess the effect of male-female interactions on the duration of copulation. This can result in inexact assertions over which sex controls copulatory behaviour. Here we analyse the repeatability of copulatory behaviour in the seed beetle Callosobruchus maculatus to determine which sex exerts primary influence over copulation duration. In C. maculatus, copulation follows two distinct phases: an initial quiescent phase followed by a period of vigorous female kicking behaviour that culminates in the termination of copulation. When males or females copulated with several novel mates, copulatory behaviour was not significantly repeatable. By contrast, when males or females mated repeatedly with the same mate, copula duration was repeatable. These data suggest copulatory behaviour in C. maculatus to be largely the product of male-female interactions rather than the consistent, sex-specific modulation of copula duration of one protagonist in response to the phenotypic variation presented by the other protagonist

Sperm death and dumping in Drosophila

Mating with more than one male is the norm for females of many species. In addition to generating competition between the ejaculates of different males, multiple mating may allow females to bias sperm use. In Drosophila melanogaster, the last male to inseminate a female sires approximately 80% of subsequent progeny. Both sperm displacement, where resident sperm are removed from storage by the incoming ejaculate of the copulating male, and sperm incapacitation, where incoming seminal fluids supposedly interfere with resident sperm, have been implicated in this pattern of sperm use. But the idea of incapacitation is problematic because there are no known mechanisms by which an individual could damage rival sperm and not their own. Females also influence the process of sperm use, but exactly how is unclear. Here we show that seminal fluids do not kill rival sperm and that any 'incapacitation' is probably due to sperm ageing during sperm storage. We also show that females release stored sperm from the reproductive tract (sperm dumping) after copulation with a second male and that this requires neither incoming sperm nor seminal fluids. Instead, males may cause stored sperm to be dumped or females may differentially eject sperm from the previous mating