Sexual Conflict Research Articles

Remodeling male coercion and the evolution of sexual autonomy by mate choice.

Models of sexual conflict over mating, including conflict over indirect benefits of mate choice, have generally presumed that female resistance to male coercion must involve direct confrontation, which can lead to sexually antagonistic coevolutionary arms-races. We built a quantitative model examining the largely ignored possibility that females may evolve new, additional mate preferences for new male traits that undermine male capacity to coerce. Thus, females may "remodel" the coercive capacity of the male phenotype in order to enhance their own sexual autonomy-a novel alternative mechanism by which females may avoid arms-races. We demonstrate that evolutionary "remodeling" is possible, in spite of costs to males, because females that prefer males with protective, autonomy-enhancing traits (traits correlated with lower coercion effectiveness) are likelier to gain indirect benefits of having attractive mates. Our analysis reveals new possibilities for the evolution of systems of sexual conflict over indirect benefits, showing that autonomy-enhancing male traits can act as a "public good," benefiting all females regardless of mating preferences, leading to oscillatory dynamics; and that preferences for more protective male traits will often be favored relative to preferences for less protective traits, potentially leading to an evolutionary "snowball" of expanding sexual autonomy.

Sex-specific estimation of cis and trans regulation of gene expression in heads and gonads of Drosophila melanogaster.

The regulatory architecture of gene expression is known to differ substantially between sexes in Drosophila, but most studies performed so far used whole-body data and only single crosses, which may have limited their scope to detect patterns that are robust across tissues and biological replicates. Here, we use allele-specific gene expression of parental and reciprocal hybrid crosses between 6 Drosophila melanogaster inbred lines to quantify cis- and trans-regulatory variation in heads and gonads of both sexes separately across 3 replicate crosses. Our results suggest that female and male heads, as well as ovaries, have a similar regulatory architecture. On the other hand, testes display more and substantially different cis-regulatory effects, suggesting that sex differences in the regulatory architecture that have been previously observed may largely derive from testis-specific effects. We also examine the difference in cis-regulatory variation of genes across different levels of sex bias in gonads and heads. Consistent with the idea that intersex correlations constrain expression and can lead to sexual antagonism, we find more cis variation in unbiased and moderately biased genes in heads. In ovaries, reduced cis variation is observed for male-biased genes, suggesting that cis variants acting on these genes in males do not lead to changes in ovary expression. Finally, we examine the dominance patterns of gene expression and find that sex- and tissue-specific patterns of inheritance as well as trans-regulatory variation are highly variable across biological crosses, although these were performed in highly controlled experimental conditions. This highlights the importance of using various genetic backgrounds to infer generalizable patterns.

Evolution of reproductive isolation in a long-term evolution experiment with Drosophila melanogaster: 30 years of divergent life-history selection.

We ask if three decades and over 1,500 generations of divergent life-history selection on age at reproduction has resulted in the evolution of reproductive isolation (RI) between laboratory populations of Drosophila melanogaster. We tested for premating, postmating-prezygotic, and postzygotic reproductive isolation between three replicate population pairs. Large, evolved differences in body size between selection treatments suggested the potential for prezygotic barriers driven by sexual selection or physical incompatibilities between the sexes. Although a simple prediction would be preference for larger size, creating directional isolation, our results from individual mate choice trials indicate that populations from both selection treatments show a marked bias towards homotypic mate choice; indicative of prezygotic RI driven by sexual selection or sexual conflict. Hybridization between the focal populations resulted in the production of viable adult flies with intermediate size and developmental traits. We observed a suggestive but statistically nonsignificant trend of fitness decline in the F2 generation of hybrids, but no significant evidence suggesting the evolution of postmating-prezygotic or postzygotic RI. Our findings are in accord with extant literature that posits that premating RI evolves before postmating forms of RI.

Thermal phenotypic plasticity of pre- and post-copulatory male harm buffers sexual conflict in wild Drosophila melanogaster.

Strong sexual selection frequently leads to sexual conflict and ensuing male harm, whereby males increase their reproductive success at the expense of harming females. Male harm is a widespread evolutionary phenomenon with a strong bearing on population viability. Thus, understanding how it unfolds in the wild is a current priority. Here, we sampled a wild Drosophila melanogaster population and studied male harm across the normal range of temperatures under which it reproduces optimally in nature by comparing female lifetime reproductive success and underlying male harm mechanisms under monogamy (i.e. low male competition/harm) vs. polyandry (i.e. high male competition/harm). While females had equal lifetime reproductive success across temperatures under monogamy, polyandry resulted in a maximum decrease of female fitness at 24°C (35%), reducing its impact at both 20°C (22%), and 28°C (10%). Furthermore, female fitness components and pre- (i.e. harassment) and post-copulatory (i.e. ejaculate toxicity) mechanisms of male harm were asymmetrically affected by temperature. At 20°C, male harassment of females was reduced, and polyandry accelerated female actuarial aging. In contrast, the effect of mating on female receptivity (a component of ejaculate toxicity) was affected at 28°C, where the mating costs for females decreased and polyandry mostly resulted in accelerated reproductive aging. We thus show that, across a natural thermal range, sexual conflict processes and their effects on female fitness components are plastic and complex. As a result, the net effect of male harm on overall population viability is likely to be lower than previously surmised. We discuss how such plasticity may affect selection, adaptation and, ultimately, evolutionary rescue under a warming climate.

Students’ Perceptions of Guilt, Responsibility, and Just Punishment When Accounts of Campus Sexual Assault Conflict

Students’ Perceptions of Guilt, Responsibility, and Just Punishment When Accounts of Campus Sexual Assault Conflict

Searching for intralocus sexual conflicts in the three-spined stickleback (Gasterosteus aculeatus) genome.

Differences between sexes in trait fitness optima can generate intralocus sexual conflicts that have the potential to maintain genetic diversity through balancing selection. However, these differences are unlikely to be associated with strong selective coefficients and are challenging to detect. Additionally, recent studies have highlighted that duplications on sexual chromosomes can create artifactual signals of intralocus sexual conflicts. Thus, testing the relationship between intralocus sexual conflicts and balancing selection requires stringent filtering of duplicated regions, and dedicated methods to detect loci with low levels of intersex differentiation. In this study, we investigated intralocus sexual conflicts in the three-spined stickleback using whole-genome sequencing (mean coverage = 12×) of 50 females and 49 males from an anadromous population in the St. Lawrence River, Québec, Canada. After stringent filtering of duplications from the sex chromosomes, we compared three methods to detect intralocus sexual conflicts. We found only two genomic regions under potential intralocus sexual conflict that also showed signals of balancing selection. Overall, our results suggest that most intralocus sexual conflicts do not drive long-term balancing selection and are most likely transient.

Functional morphology of the praying mantis male genitalia (Insecta: Mantodea)

Male genitalia in praying mantids are highly complex, but we know little of how they function. I combined the micro-computed tomography of a copulating pair of the European mantis (Mantis religiosa) with public videos of copulation in various species of Mantodea and an analysis of literature. The function of each major element is reviewed. Copulation is divided into three phases: opening, anchoring and deposition. The opening is achieved by pulling the female subgenital plate with the male apical process. Multiple cases of female cooperation or resistance were observed and one case of coercion by the male. In species with the reduced apical process, female cooperation is mandatory. The male subgenital plate may participate in the opening as an integral part of the genitalia. After the opening, the conformation of the genitalia drastically changes, revealing activity of the genital papilla. Tight grasp on female genitalia is maintained solely by the clamp on the right phallomere, despite the overall complexity and predictions of sexual conflict theory. Other prominent elements show rhythmic motions, but their functions are not entirely clear and evidently involve spermatophore deposition, female stimulation or rival sperm removal. The opening and anchoring are similar in Mantodea and Blattodea, but achieved with non-homologous elements.

Increased male-induced harm in response to female-limited selection: interactive effects between intra- and interlocus sexual conflict?

Interlocus sexual conflict (IRSC) occurs because of shared interactions that have opposite effects on male and female fitness. Typically, it is assumed that loci involved in IRSC have sex-limited expression and are thus not directly affected by selective pressures acting on the other sex. However, if loci involved in IRSC have pleiotropic effects in the other sex, intersexual selection can shape the evolutionary dynamics of conflict escalation and resolution, as well as the evolution of reproductive traits linked to IRSC loci, and vice versa. Here we used an artificial selection approach in Japanese quail (Coturnix japonica) to test if female-limited selection on reproductive investment affects the amount of harm caused by males during mating. We found that males originating from lines selected for high female reproductive investment caused more oxidative damage in the female reproductive tract than males originating from lines selected for low female reproductive investment. This male-induced damage was specific to the oviduct and not found in other female tissues, suggesting that it was ejaculate-mediated. Our results suggest that intersexual selection shapes the evolution of IRSC and that male-induced harm may contribute to the maintenance of variation in female reproductive investment.

Sex wars: a female genital spine forces male damselflies to shorten copulation duration.

In some species, males use weapons to harm females, increasing their short-term fitness. Here we show that females can use genital adaptations against males. Females of the damselfly Enallagma cyathigerum have a conspicuous vulvar spine on the eighth abdominal segment, which contacts with the male during copulation. We tested 3 hypotheses for its function: it (a) inflicts damage to the male during copulation; (b) facilitates endophytic oviposition; and (c) stimulates males during copulation to increase their investment. We found that males mated on average for 54 min with control females, but increased copulation to 99 min with females without spine. There was no evidence of physical harm of the spine on the male's seminal vesicle, which shows 8- to 18-folds, exactly where the spine contacts during copulation. Females with and without spine exhibited the same egg-laying rates and showed similar fecundity and fertility. Longevity was also similar in males mated to control and spineless females. In contrast to many species where females resist male harassment by behavioral responses, the morphological adaptation observed in E. cyathigerum appears to act as a sexual weapon, allowing females to control copulation duration. We suggest that the spine has evolved because of sexual conflict over mating duration.

Post-Trauma Stress Disorder in Sexual Harassment: A Case Report

Background: Violence or sexual harassment have been increasingly reported in recent years, especially among women. Not only at the global level, but also nationally and locally. Violence in the form of sexual harassment and rape can cause deep trauma to the victim, depression, and other mental disorders. Methods: A case study of sexual harassment that causes post-traumatic stress disorder. Results and discussion: an 18-year-old woman, still in high school, complained of fear after experiencing several times of violence or sexual harassment by both close people and strangers. With a multiaxial approach, the victim is diagnosed with Post-Traumatic Stress Disorder. It was confirmed that the victim had severe depression using the Hamilton Depression Rating Scale or extreme depression using the Beck Depression Inventory. In this case, several factors were identified such as: being a victim or previous sexual act, conflict, and violence in the family, emotionally unsupportive family environment, poor parent-child relationships, and poverty. Comprehensive management involves a multidisciplinary approach. Pharmacologically, Risperidone was given 1-milligram tablet every 24 hours orally in the morning, and Sertraline 25 milligrams tablet every 24 hours orally at night. Victims also receive psychotherapy, education, and social support. Furthermore, the victim is monitored regularly. Conclusion: Sexual violence or harassment is prone to cause Post-Traumatic Stress Disorder, so it requires a multidisciplinary approach and comprehensive management. Starting from taking a history to confirm the diagnosis, treatment, and monitoring.

Overruled by nature: A plastic response to environmental change disconnects a gene and its trait.

In Atlantic salmon, age at maturation is a life history trait governed by a sex-specific trade-off between reproductive success and survival. Following environmental changes across large areas of the Northeast Atlantic, many populations currently display smaller size at age and higher age at maturation. However, whether these changes reflect rapid evolution or plasticity is unknown. Approximately 1500 historical and contemporary salmon from the river Etne in Western Norway, genotyped at 50,000 SNPs, revealed three loci associated with age at maturation. These included vgll3 and six6 which collectively explained 36%-50% of the age at maturation variation in the 1983-1984 period. These two loci also displayed sex-specific epistasis, as the effect of six6 was only detected in males bearing two copies of the late maturation allele for vgll3. Strikingly, despite allelic frequencies at vgll3 remaining unchanged, the combined influence of these genes was nearly absent in all samples from 2013 to 2016, and genome-wide heritability strongly declined between the two time-points. The difference in age at maturation between males and females was upheld in the population despite the loss of effect from the candidate loci, which strongly points towards additional causative mechanisms resolving the sexual conflict. Finally, because admixture with farmed escaped salmon was excluded as the origin of the observed disconnection between gene(s) and maturation age, we conclude that the environmental changes observed in the North Atlantic during the past decades have led to bypassing of the influence of vgll3 and six6 on maturation through growth-driven plasticity.

Resolving sexual conflict in plants

Why is it that one plant species has separate sexes, while its sister species in the phylogeny remains hermaphroditic? This question lies at the core of Professor John Pannell’s research.

Domestication affects sex-biased gene expression evolution in the duck.

Genes with sex-biased expression are thought to underlie sexually dimorphic phenotypes and are therefore subject to different selection pressures in males and females. Many authors have proposed that sexual conflict leads to the evolution of sex-biased expression, which allows males and females to reach separate phenotypic and fitness optima. The selection pressures associated with domestication may cause changes in population architectures and mating systems, which in turn can alter their direction and strength. We compared sex-biased expression and genetic signatures in wild and domestic ducks (Anas platyrhynchos), and observed changes of sexual selection and identified the genomic divergence affected by selection forces. The extent of sex-biased expression in both sexes is positively correlated with the level of both d N /d S and nucleotide diversity. This observed changing pattern may mainly be owing to relaxed genetic constraints. We also demonstrate a clear link between domestication and sex-biased evolutionary rate in a comparative framework. Decreased polymorphism and evolutionary rate in domesticated populations generally matched life-history phenotypes known to experience artificial selection. Taken together, our work suggests the important implications of domestication in sex-biased evolution and the roles of artificial selection and sexual selection for shaping the diversity and evolutionary rate of the genome.

Female genital concealment and a corresponding male clasping apparatus in Australian ripple bugs (Hemiptera: Veliidae)

Conflicts of interest over reproduction between males and females are widespread in sexually reproducing species. This is exemplified in water striders (Gerridae), where females vigorously resist costly mating attempts, and males and females often exhibit elaborate grasping and anti-grasping morphological traits. Like water striders, their sister-group, the ripple bugs (Veliidae), share similar life histories and are expected to face similar conflicts over mating. Veliids in the genus Nesidovelia exhibit elaborate sexual dimorphism, which is predicted to function in intersexual antagonistic struggles. This includes concealed genitalia in females, and elaborate pregenital abdominal modifications in males. By documenting mating behaviour in Nesidovelia peramoena and freezing pairs in copula, we show that males and females struggle prior to mating, and male abdominal modifications function to gain access to the female's concealed genitalia. This is consistent with, though not limited to, sexual conflict.

The roles of sexual selection and sexual conflict in shaping patterns of genome and transcriptome variation.

Sexual dimorphism is one of the most prevalent, and often the most extreme, examples of phenotypic variation within species, and arises primarily from genomic variation that is shared between females and males. Many sexual dimorphisms arise through sex differences in gene expression, and sex-biased expression is one way that a single, shared genome can generate multiple, distinct phenotypes. Although many sexual dimorphisms are expected to result from sexual selection, and many studies have invoked the possible role of sexual selection to explain sex-specific traits, the role of sexual selection in the evolution of sexually dimorphic gene expression remains difficult to differentiate from other forms of sex-specific selection. In this Review, we propose a holistic framework for the study of sex-specific selection and transcriptome evolution. We advocate for a comparative approach, across tissues, developmental stages and species, which incorporates an understanding of the molecular mechanisms, including genomic variation and structure, governing gene expression. Such an approach is expected to yield substantial insights into the evolution of genetic variation and have important applications in a variety of fields, including ecology, evolution and behaviour.

Sexually antagonistic selection maintains genetic variance when sexual dimorphism evolves.

Genetic variance (VG) in fitness related traits is often unexpectedly high, evoking the question how VG can be maintained in the face of selection. Sexually antagonistic (SA) selection favouring alternative alleles in the sexes is common and predicted to maintain VG, while directional selection should erode it. Both SA and sex-limited directional selection can lead to sex-specific adaptations but how each affect VG when sexual dimorphism evolves remain experimentally untested. Using replicated artificial selection on the seed beetle Callosobruchus maculatus body size we recently demonstrated an increase in size dimorphism under SA and male-limited (ML) selection by 50% and 32%, respectively. Here we test their consequences on genetic variation. We show that SA selection maintained significantly more ancestral, autosomal additive genetic variance than ML selection, while both eroded sex-linked additive variation equally. Ancestral female-specific dominance variance was completely lost under ML, while SA selection consistently sustained it. Further, both forms of selection preserved a high genetic correlation between the sexes (rm,f). These results demonstrate the potential for sexual antagonism to maintain more genetic variance while fuelling sex-specific adaptation in a short evolutionary time scale, and are in line with predicted importance of sex-specific dominance reducing sexual conflict over alternative alleles.

Genetic study reveals local differentiation persisting in the face of high connectivity and a genomic inversion likely linked with sexual antagonism in a common marine fish

Abstract Sustainable harvest of wild populations requires knowledge of the underlying population structure. The focus of this study is on goldsinny wrasse (Ctenolabrus rupestris), a small marine fish inhabiting coastal waters of the north-eastern Atlantic. This species is caught in large numbers to serve as cleaner fish in salmonid aquaculture. We genotyped 2073 goldsinny wrasse from 43 sites along the Scandinavian coastline with 143 SNPs. Seven of the SNPs were linked and likely reside within a large genomic inversion dominated by one haplotype. The heterokaryotype of the putative inversion displayed sex-specific growth patterns, potentially resolving sexual antagonism for this trait. The unlinked 134 SNPs showed modest isolation-by-distance with samples from the northernmost locations showing highest divergence, whereas sites farther south were much more interconnected. Genetic divergence (FST) was highly variable among sites within regions, suggesting a varying degree of connectivity and local divergence. We conclude that despite a high degree of gene-flow mediated through pelagic dispersal in early life stages, regional and some local population structure remains due to limited adult movement in addition to other unidentified factors. Consequently, the species might be more vulnerable to local disturbances than previously anticipated.

Females adopt sexual catalepsy to facilitate mating.

Theory predicts that males and females of dioecious species typically engage in an evolutionary sexual conflict over the frequency and choice of mating partner. Female sexual cannibalism, a particularly dramatic illustration of this conflict, is widespread in certain animal taxa including spiders. Nevertheless, females of some funnel weaving spiders that are generally aggressive to conspecifics enter a cataleptic state after male courtship, ensuring the males can mate without risk of attack. In this study, we demonstrated that the physical posture and duration, metabolites, and central neurotransmitters of females of Aterigena aculeata in sexual catalepsy closely resemble females in thanatosis but are distinct from those in anesthesia, indicating that the courted females feign death to eliminate the risk of potentially aggressive responses and thereby allow preferred males to mate. Unlike the taxonomically widespread thanatosis, which generally represents a deceptive visual signal that acts against the interest of the receivers, sexual catalepsy of females in the funnel weaving spiders may deliver a sexual-receptive signal to the courting males and thereby benefit both the signal senders and receivers. Therefore, sexual catalepsy in A. aculeata may not reflect a conflict but rather a confluence of interest between the sexes.

Fertility signalling games: should males obey the signal?

Game theory is frequently used to study conflicting interests between the two sexes. Males often benefit from a higher mating rate than females do. A temporal component of this conflict has rarely been modelled: females' interest in mating may depend on when females become fertile. This sets conditions for male-female coevolution, where females may develop fertility signals, and males may obey the signal, such that they only target signalling females. Modelling this temporal aspect to sexual conflict yields two equilibria: (i) a trivial equilibrium without signals and with males targeting all females, and (ii) a signalling equilibrium where all females signal before ovulation, and where either some, or all, males obey the signal. The 'all males obey the signal' equilibrium is more likely if we assume that discriminating males have an advantage in postcopulatory sperm competition, while in the absence of this benefit, we find the 'some males obey the signal' equilibrium. The history of game-theoretic models of sex differences often portrays one sex as the 'winner' and the opposite sex as the 'loser'. From early models emphasizing 'battle of the sexes'-style terminology, we recommend moving on to describe the situation as non-signalling equilibria having stronger unresolved sexual conflict than signalling equilibria. This article is part of the theme issue 'Half a century of evolutionary games: a synthesis of theory, application and future directions'.

Y chromosome-linked variation affects locomotor activity in male Drosophila melanogaster and is robust to differences in thermal environment

Although containing genes important for sex determination, genetic variation within the Y chromosome was traditionally predicted to contribute little to the expression of sexually dimorphic traits. This prediction was shaped by the assumption that the chromosome harbours few protein-coding genes, and that capacity for Y-linked variation to shape adaptation would be hindered by the chromosome’s lack of recombination and holandric inheritance. Consequently, most studies exploring the genotypic contributions to sexually dimorphic traits have focused on the autosomes and X chromosome. Yet, several studies have now demonstrated that the Y chromosome harbours variation affecting male fitness, moderating the expression of hundreds of genes across the nuclear genome. Furthermore, emerging results have shown that expression of this Y-linked variation may be sensitive to environmental heterogeneity, leading to the prediction that Y-mediated gene-by-environment interactions will shape the expression of sexually dimorphic phenotypes. We tested this prediction, investigating whether genetic variation across six distinct Y chromosome haplotypes affects the expression of locomotor activity, at each of two temperatures (20 and 28 °C) in male fruit flies (Drosophila melanogaster). Locomotor activity is a sexually dimorphic trait in this species, previously demonstrated to be under intralocus sexual conflict. We demonstrate Y haplotype effects on male locomotor activity, but the rank order and magnitude of these effects were unaltered by differences in temperature. Our study contributes to a growing number of studies demonstrating Y-linked effects moderating expression of traits evolving under sexually antagonistic selection, suggesting a role for the Y chromosome in shaping outcomes of sexual conflict.