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Sexual selection and the evolution of evolvability.

Here we show that sexual selection can have an effect on the rate of mutation. We simulated the fate of a genetic modifier of the mutation rate in a sexual population with and without sexual selection (modelled using a female choice mechanism). Female choice for 'good genes' should reduce variability among male subjects, leaving insufficient differences to maintain female preferences. However, female choice can actually increase genetic variability by supporting a higher mutation rate in sexually selected traits. Increasing the mutation rate will be selected against because of the resulting decline in mean fitness. However, it also increases the probability of rare beneficial mutations arising, and mating skew caused by female preferences for male subjects carrying those beneficials with few deleterious mutations ('good genes') can lead to a mutation rate above that expected under natural selection. A choice of two male subjects was sufficient for there to be a twofold increase in the mutation rate as opposed to a decrease found under random mating.

Animals↗

Sexual selection and fertility.

Genetic models are analyzed in which sexual selection is combined with fertility selection. In these models, the sexual selection acts on males, the fertility selection on either males, females or both sexes. The phenotypes thus selected may be determined either by dominant and recessive alleles or by each homozygous and heterozygous genotype. Polymorphisms of dominant and recessive phenotypes can be maintained in equilibrium by a balance between sexual and fertility selection. Generally fertility selection has a greater effect than viability selection in determining the point of equilibrium. The dominant phenotype is maintained at a lower frequency when at a fertility disadvantage than when at a viability disadvantage. When about 20% or more of the females mate preferentially, the models show that equilibria will be established at very different frequencies depending on whether fertility selection acts on males, females or both sexes. These results, applied to data of preferential mating of melanic two-spot ladybirds, predict differences in fertility which can be use to test the models. Symmetric models of preferences for each genotype also give rise to polymorphisms if the heterozygotes obtain an overall advantage.

Animals↗

The Bateman gradient and the cause of sexual selection in a sex-role-reversed pipefish.

As a conspicuous evolutionary mechanism, sexual selection has received much attention from theorists and empiricists. Although the importance of the mating system to sexual selection has long been appreciated, the precise relationship remains obscure. In a classic experimental study based on parentage assessment using visible genetic markers, more than 50 years ago A. J. Bateman proposed that the cause of sexual selection in Drosophila is 'the stronger correlation, in males (relative to females), between number of mates and fertility (number of progeny)'. Half a century later, molecular genetic techniques for assigning parentage now permit mirror-image experimental tests of the 'Bateman gradient' using sex-role-reversed species. Here we show that, in the male-pregnant pipefish Syngnathus typhle, females exhibit a stronger positive association between number of mates and fertility than do males and that this relationship responds in the predicted fashion to changes in the adult sex ratio. These findings give empirical support to the idea that the relationship between mating success and number of progeny, as characterized by the Bateman gradient, is a central feature of the genetic mating system affecting the strength and direction of sexual selection.

Animals↗

Males' evolutionary responses to experimental removal of sexual selection.

We evaluated the influence of pre- and post-copulatory sexual selection upon male reproductive traits in a naturally promiscuous species, Drosophila melanogaster. Sexual selection was removed in two replicate populations through enforced monogamous mating with random mate assignment or retained in polyandrous controls. Monogamous mating eliminates all opportunities for mate competition, mate discrimination, sperm competition, cryptic female choice and, hence, sexual conflict. Levels of divergence between lines in sperm production and male fitness traits were quantified after 38-81 generations of selection. Three a priori predictions were tested: (i) male investment in spermatogenesis will be lower in monogamy-line males due to the absence of sperm competition selection, (ii) due to the evolution of increased male benevolence, the fitness of females paired with monogamy-line males will be higher than that of females paired with control-line males, and (iii) monogamy-line males will exhibit decreased competitive reproductive success relative to control-line males. The first two predictions were supported, whereas the third prediction was not. Monogamy males evolved a smaller body size and the size of their testes and the number of sperm within the testes were disproportionately further reduced. In contrast, the fitness of monogamous males (and their mates) was greater when reproducing in a non-competitive context: females mated once with monogamous males produced offspring at a faster rate and produced a greater total number of surviving progeny than did females mated to control males. The results indicate that sexual selection favours the production of increased numbers of sperm in D. melanogaster and that sexual selection favours some male traits conferring a direct cost to the fecundity of females.

Adaptation, Biological↗

Sexually selected signals are not similar to sports handicaps.

The handicap principle is a simple but powerful metaphor that has had a major impact on how biologists study and understand sexual selection. Here, I show that its application to signalling in sexual selection is not a valid generalization from its roots in economics. Although some signalling systems, with additive costs and benefits, have solutions that resemble sports handicaps, the signalling in sexual selection has multiplicative costs and benefits, and solutions that do not resemble sports handicaps. The sports analogy is technically incorrect, metaphorically misleading and a poor guide for empirical research on the signalling in sexual selection. The evolution of sexually selected signals is not a missing piece of Darwin's puzzle; it is an integral piece of the process of evolution by natural selection, and it should be approached with the same tools that we bring to bear on the evolution of other correlated traits involved in social interactions.

Animal Communication↗

Sympatric speciation by sexual selection alone is unlikely.

According to Darwin, sympatric speciation is driven by disruptive, frequency-dependent natural selection caused by competition for diverse resources. Recently, several authors have argued that disruptive sexual selection can also cause sympatric speciation. Here, we use hypergeometric phenotypic and individual-based genotypic models to explore sympatric speciation by sexual selection under a broad range of conditions. If variabilities of preference and display traits are each caused by more than one or two polymorphic loci, sympatric speciation requires rather strong sexual selection when females exert preferences for extreme male phenotypes. Under this kind of mate choice, speciation can occur only if initial distributions of preference and display are close to symmetric. Otherwise, the population rapidly loses variability. Thus, unless allele replacements at very few loci are enough for reproductive isolation, female preferences for extreme male displays are unlikely to drive sympatric speciation. By contrast, similarity-based female preferences that do not cause sexual selection are less destabilizing to the maintenance of genetic variability and may result in sympatric speciation across a broader range of initial conditions. Certain groups of African cichlids have served as the exclusive motivation for the hypothesis of sympatric speciation by sexual selection. Mate choice in these fishes appears to be driven by female preferences for extreme male phenotypes rather than similarity-based preferences, and the evolution of premating reproductive isolation commonly involves at least several genes. Therefore, differences in female preferences and male display in cichlids and other species of sympatric origin are more likely to have evolved as isolating mechanisms under disruptive natural selection.

Animals↗

Direct versus indirect sexual selection: genetic basis of colour, size and recruitment in a wild bird.

Indirect and direct models of sexual selection make different predictions regarding the quantitative genetic relationships between sexual ornaments and fitness. Indirect models predict that ornaments should have a high heritability and that strong positive genetic covariance should exist between fitness and the ornament. Direct models, on the other hand, make no such assumptions about the level of genetic variance in fitness and the ornament, and are therefore likely to be more important when environmental sources of variation are large. Here we test these predictions in a wild population of the blue tit (Parus caeruleus), a species in which plumage coloration has been shown to be under sexual selection. Using 3 years of cross-fostering data from over 250 breeding attempts, we partition the covariance between parental coloration and aspects of nestling fitness into a genetic and environmental component. Contrary to indirect models of sexual selection, but in agreement with direct models, we show that variation in coloration is only weakly heritable h2<0.11, and that two components of offspring fitness-nestling size and fledgling recruitment-are strongly dependent on parental effects, rather than genetic effects. Furthermore, there was no evidence of significant positive genetic covariation between parental colour and offspring traits. Contrary to direct benefit models, however, we find little evidence that variation in colour reliably indicates the level of parental care provided by either males or females. Taken together, these results indicate that the assumptions of indirect models of sexual selection are not supported by the genetic basis of the traits reported on here.

Animals↗

Mating system and demographic constraints on the opportunity for sexual selection.

In monogamous systems the fitness difference between males due to competition for mates is limited to one female. This constraint presumably impedes the action of sexual selection relative to polygynous systems. In this paper, we use formal selection theory to show how population size and the adult sex ratio constrain the force of sexual selection and phenotypic evolution under monogamy and polygyny. The force of sexual selection is ultimately constrained by the number of males in a population and the theoretical limit to the rate of male phenotypic evolution is realized if a single male mates with one or many females. These results imply that the force of sexual selection is not strictly constrained by monogamy. The constraint on female phenotypic evolution is typically higher than the constraint on males under polygyny and similar to selection on males in monogamous systems. The sexual asymmetry in the force of selection under polygyny--not necessarily weak sexual selection on males of monogamous systems--may explain the prominence of sexual dimorphism in polygynous systems.

Animals↗

Sexual selection in relation to pest-management strategies.

The application of principles derived from the sexual selection literature can assist attempts to subvert the normal mating behavior of pests. Sexual selection encompasses both intermale competition for access to females and female choice of mates. It can operate during long-range attraction and short-range courtship, as well as after copulation. We review the major aspects of sexual selection and illustrate their application to pheromonal and SIT pest-management programs. Pheromones are important both in long-range attraction and in close-range mate choice; parapheromones may be very useful in pest management because of their influence on male mating success. Sexual selection theory provides a scheme for studying the normal mating behavior of a pest species and thus determining which attributes of the mass-reared sterile males are critical to their success with wild females. We hope that our review will suggest novel ways of attacking pests as well as encourage behavioral ecologists to study pest species.

Journal Article↗

Contrasting mutual sexual selection on homologous signal traits in Drosophila serrata.

The nature of male mating preferences, and how they differ from female mating preferences in species with conventional sex roles, has received little attention in sexual selection studies. We estimated the form and strength of sexual selection as a consequence of male and female mating preferences in a laboratory-based population of Drosophila serrata. The differences between sexual selection on male and female signal traits (cuticular hydrocarbons [CHCs]) were evaluated within a formal framework of linear and nonlinear selection gradients. Females tended to exert linear sexual selection on male CHCs, whereas males preferred intermediate female CHC phenotypes leading to convex (stabilizing) selection gradients. Possible mechanisms determining the nonlinear nature of sexual selection on female CHCs are proposed.

Animals↗

Juvenile immune status affects the expression of a sexually selected trait in field crickets.

Parasite-mediated sexual selection theory presumes that variation in sexual traits reliably reflects variation in parasite resistance among available mates. One mechanism that may warrant signal honesty involves costs of immune system activation in the case of a parasitic infection. We investigated this hypothesis in male field crickets Gryllus campestris, whose attractiveness to females depends on characteristics of the sound-producing harp that are essentially fixed following adult eclosion. During the nymphal stage, males subjected to one of two feeding regimes were challenged with bacterial lipopolysaccharides (LPS) to investigate condition-dependent effects on harp development as compared to other adult traits. Nymphal nutritional status positively affected adult body size, condition, and harp size. However, nymphal immune status affected harp size only, with LPS-males having smaller harps than control-injected males. In addition, the harps of LPS-males showed a lesser degree of melanization, indicating an enhanced substrate use by the melanin-producing enzyme cascade of the immune system. Thus, past immune status is specifically mirrored in sexual traits, suggesting a key role for deployment costs of immunity in parasite-mediated sexual selection.

Animal Communication↗

Is sexual selection and species recognition a continuum? Mating behavior of the stalk-eyed fly Drosophila heteroneura.

If behavioral isolation between species can evolve as a consequence of sexual selection within a species, then traits that are both sexually selected and used as a criterion of species recognition by females should be identifiable. The broad male head of the Hawaiian picture-winged fly Drosophila heteroneura is a novel sexual dimorphism that may be sexually selected and involved in behavioral isolation from D. silvestris. We found that males with broad heads are more successful in sexual selection, both through female mate choice and through aggressive interactions. However, female D. heteroneura do not discriminate against hybrids on the basis of their head width. Thus, this novel trait is sexually selected but is not a major contributor to species recognition. Our methods should be applicable to other species in which behavioral isolation is a factor.

Animals↗

Sexual selection on multivariate phenotype in wild and mass-reared Ceratitis capitata (Diptera: Tephritidae).

Males with a larger thorax and narrower face were found to be favoured by directional sexual selection in a previous field cage experiment where Ceratitis capitata males from the Seib 6-96 genetic sexing strain competed with wild males from Alto Valle (Patagonia) for the possession of wild females. Targets of sexual selection, however, might differ between wild and laboratory females as a response to adaptation to mass-rearing conditions. To evaluate possible divergences on the targets of sexual selection as a by-product of adaptation to mass-rearing conditions, field cage tests were performed with both wild and laboratory females. To avoid possible bias due to correlation among the measured traits (eye length [EL], face width [FW], head width [HW], and thorax length [TL]), a multivariate analysis was applied. Consistent with the previous experiment, the results indicated that TL and FW are probable targets of directional sexual selection independently of female strain. However, laboratory females were less selective than wild ones. Additionally, correlational sexual selection was detected acting on the multivariate phenotype. The effects of correlational selection overlap with those of directional selection on each single trait. The analysis of mating pair characteristics showed patterns that do not match the expectations for a random mating system. The current analysis indicates that during mating pair formation two processes overlap. On the one hand, sexual selection favours males with larger size (TL) and narrower faces (FW). This effect occurs in both wild and laboratory females. In addition, assortative mating based on both phenotype and origin was also observed.

Animals↗

How cuckoldry can decrease the opportunity for sexual selection: data and theory from a genetic parentage analysis of the sand goby, Pomatoschistus minutus.

Alternative mating strategies are common in nature and are generally thought to increase the intensity of sexual selection. However, cuckoldry can theoretically decrease the opportunity for sexual selection, particularly in highly polygamous species. We address here the influence of sneaking (fertilization thievery) on the opportunity for sexual selection in the sand goby Pomatoschistus minutus, a marine fish species in which males build and defend nests. Our microsatellite-based analysis of the mating system in a natural sand goby population shows high rates of sneaking and multiple mating by males. Sneaker males had fertilized eggs in approximately 50% of the assayed nests, and multiple sneakers sometimes fertilized eggs from a single female. Successful males had received eggs from 2 to 6 females per nest (mean = 3.4). We developed a simple mathematical model showing that sneaking in this polygynous sand goby population almost certainly decreases the opportunity for sexual selection, an outcome that contrasts with the usual effects of cuckoldry in socially monogamous animals. These results highlight a more complex and interesting relationship between cuckoldry rates and the intensity of sexual selection than previously assumed in much of the literature on animal mating systems.

Animals↗

Sexual selection, germline mutation rate and sperm competition.

BACKGROUND: An important component of sexual selection arises because females obtain viability benefits for their offspring from their mate choice. Females choosing extra-pair fertilization generally favor males with exaggerated secondary sexual characters, and extra-pair paternity increases the variance in male reproductive success. Furthermore, females are assumed to benefit from 'good genes' from extra-pair sires. How additive genetic variance in such viability genes is maintained despite strong directional selection remains an evolutionary enigma. We propose that sexual selection is associated with elevated mutation rates, changing the balance between mutation and selection, thereby increasing variance in fitness and hence the benefits to be obtained from good genes sexual selection. Two hypotheses may account for such elevated mutation: (1) Increased sperm production associated with sperm competition may increase mutation rate. (2) Mutator alleles increase mutation rates that are revealed by the expression of condition-dependent secondary sexual characters used by choosy females during their mate choice. M Petrie has independently developed the idea that mutator alleles may account for the maintenance of genetic variation in viability despite strong directional selection. RESULTS: A comparative study of birds revealed a positive correlation between mutation rate at minisatellite loci and extra-pair paternity, but not between mutation rate and relative testes mass which is a measure of relative sperm production. Minisatellite mutation rates were not related to longevity, suggesting a meiotic rather than a mitotic origin of mutations. CONCLUSION: We found evidence of increased mutation rate in species with more intense sexual selection. Increased mutation was not associated with increased sperm production, and we suggest that species with intense sexual selection may maintain elevated mutation rates because sexual selection continuously benefits viability alleles expressed in condition-dependent characters. Sexual selection may increase mutational input, which in turn feeds back on sexual selection because of increased variance in viability traits.

Animals↗

Sexual selection and mate choice.

The past two decades have seen extensive growth of sexual selection research. Theoretical and empirical work has clarified many components of pre- and postcopulatory sexual selection, such as aggressive competition, mate choice, sperm utilization and sexual conflict. Genetic mechanisms of mate choice evolution have been less amenable to empirical testing, but molecular genetic analyses can now be used for incisive experimentation. Here, we highlight some of the currently debated areas in pre- and postcopulatory sexual selection. We identify where new techniques can help estimate the relative roles of the various selection mechanisms that might work together in the evolution of mating preferences and attractive traits, and in sperm-egg interactions.

Adaptation, Physiological↗

The dilemma of female mate selection in the brown bear, a species with sexually selected infanticide.

Because of differential investment in gametes between sexes, females tend to be the more selective sex. Based on this concept, we investigate mate selection in a large carnivore: the brown bear (Ursus arctos). We hypothesize that, in this species with sexually selected infanticide (SSI), females may be faced with a dilemma: either select a high-quality partner based on phenotypic criteria, as suggested by theories of mate choice, or rather mate with future potentially infanticidal males as a counter-strategy to SSI. We evaluated which male characteristics were important in paternity assignment. Among males available in the vicinity of the females, the largest, most heterozygous and less inbred and also the geographically closest males were more often the fathers of the female's next litter. We suggest that female brown bears may select the closest males as a counter-strategy to infanticide and exercise a post-copulatory cryptic choice, based on physical attributes, such as a large body size, reflecting male genetic quality. However, male-male competition either in the form of fighting before copulation or during the post-copulatory phase, in the form of sperm competition, cannot entirely be ruled out.

Animals↗

Models of sexual selection on a quantitative genetic trait when preference is acquired by sexual imprinting.

The evolution of a quantitative genetic trait under stabilizing viability selection and sexual selection is modeled for a polygynous species in which female mating preferences are acquired by sexual imprinting on the parents and by exposure to the surviving population at large. Stabilizing viability selection acts equally on both sexes in the case of a sexually monomorphic trait and on males only in the case of a dimorphic trait. A genetically fixed sensory or perceptual bias defines the origin of the scale on which the trait is measured, and the possibility is incorporated that female preferences may deviate asymmetrically from the familiar-either toward or away from this origin. When viability selection is strong relative to sexual selection, the models predict that the mean trait value will evolve to the viability optimum. With intermediate ratios of the strength of viability to sexual selection, a stable equilibrium can occur on either side of this viability optimum, depending on the direction of asymmetry in female preferences. When viability selection is relatively weak and certain other conditions are also satisfied, runaway selection is predicted.

Animals↗