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Inheritance of a secondary sexual character in Drosophila silvestris.

Reciprocal crosses were carried out between laboratory stock specimens obtained from two races of Drosophila silvestris from the island of Hawaii that differ in a quantitative secondary sexual character. The race from the Hilo side of the island has a novel attribute, consisting of an extra row of cilia on the tibia of males, which is used during courtship. With regard to this character, sex-linked genes contribute about 30% of the difference, and the remaining 70% of the difference between the races is produced by genes on at least two autosomes. The novel character appears to have been the outcome of altered sexual selection in the Hilo-side race. In an altered genetic environment, resulting from a founder event or random genetic drift, sexual selection may take a new direction. Such a shift may serve as a model for incipient speciation.

Animals↗

Trade-offs between immune investment and sexual signaling in male mallards.

Allocation trade-offs between the immune system and sexual traits are central to current sexual selection hypotheses but remain contentious. Such trade-offs could be brought about by the dual action of testosterone that stimulates sexual signals but also suppresses immune functions and/or by competition for carotenoids that can be deposited in ornaments or used as antioxidants in support of immune functions. We investigated the trade-off between investment in immunity and maintenance of testosterone, carotenoids, and sexually selected, carotenoid-based bill color in male mallards. Following a nonpathogenic immune challenge, facultative immune investment resulted in a syndrome of changes in allocation. Plasma carotenoids disappeared from circulation proportional to antibody production. In addition, the reflectance spectrum of the bill was affected; greater antibody production was associated with an increase in relative UV reflectance. Although changes in bill reflectance and plasma carotenoids were related, the relationship appeared more complex than direct competition with immunity. Finally, maintenance of testosterone was affected by immune investment: testosterone levels declined substantially when males produced more antibodies. Because males with high testosterone are preferred by females, the decline in testosterone, in addition to carotenoid depletion and effects on bill reflectance, could constitute a significant cost of immune investment.

Adaptation, Physiological↗

The Hamilton-Zuk theory and initial test: an examination of some parasitological criticisms.

It has been supposed repeatedly that the Hamilton & Zuk paper of 1982 was parasitologically misconceived. These criticisms are shown to be inconsistent or mistaken. Various assumptions deemed unjustifiable by critics were either not made in the original paper, or have been made commonly, reasonably and pragmatically, by parasitologists themselves. The requirement by the theory for pathogenicity is examined, as is the need for the individuals to encounter the relevant parasites. The possible roles of parasite aggregation, parasites other than those directly involved, and costs of resistance, in making sosigonic selection less feasible are put into perspective. The rationale, interpretation and value of the preliminary comparative test with haematozoa are discussed. The advisability of incorporating host age, the location and timing of any sampling for parasites, and the actual ability of successful mates to resist disease is acknowledged. Attention is directed towards several specific observations. (1) It is a little recognised fact that Hamilton and Zuk assessed prevalence-showiness associations within surveys in restricted localities. (2) Many parasitological studies have not properly addressed the need to control for sampling effort when using prevalence data. (3) Contrary to some claims, the notion that haematozoa are sufficiently pathogenic is not unrealistic. (4) That non-resistant as well as resistant individuals might be free of patent or even latent infection at the time of mate choice was one of the possibilities mentioned originally. (5) In 1984, Eshel & Hamilton emphasised that environmental variation (whatever its source) has the potential to make mate selection for heritable characters based on perceptible variation less likely to evolve. (6) Hamilton & Zuk concluded that their findings "hint" that parasitism could be one agitator required for "good genes" sexual selection, a stance which is not immoderate. Lastly, the potential importance in evolutionary biology of parasite-mediated sexual selection as a form of co-evolution is considered briefly.

Host-Parasite Interactions↗

Variation in mate choice and mating preferences: a review of causes and consequences.

The aim of this review is to consider variation in mating preferences among females. We define mating preferences as the sensory and behavioural properties that influence the propensity of individuals to mate with certain phenotypes. Two properties of mating preferences can be distinguished: (1) "preference functions'-the order with which an individual ranks prospective mates and (2) "choosiness'-the effort an individual is prepared to invest in mate assessment. Patterns of mate choices can be altered by changing the costs of choosiness without altering the preference function. We discuss why it is important to study variation in female mating behaviour and identify five main areas of interest: Variation in mating preferences and costs of choosiness could (1) influence the rate and direction of evolution by sexual selection, (2) provide information about the evolutionary history of female preferences, (3) help explain inter-specific differences in the evolution of secondary sexual characteristics, (4) provide information about the level of benefits gained from mate choice, (5) provide information about the underlying mechanisms of mate choice. Variation in mate choice could be due to variability in preference functions, degree of choosiness, or both, and may arise due to genetic differences, developmental trajectories or proximate environmental factors. We review the evidence for genetic variation from genetic studies of heritability and also from data on the repeatability of mate-choice decisions (which can provide information about the upper limits to heritability). There can be problems in interpreting patterns of mate choice in terms of variation in mating preferences and we illustrate two main points. First, some factors can lead to mate choice patterns that mimic heritable variation in preferences and secondly other factors may obscure heritable preferences. These factors are divided into three overlapping classes, environmental, social and the effect of the female phenotype. The environmental factors discussed include predation risk and the costs of sampling; the social factors discussed include the effect of male-male interactions as well as female competition. We review the literature which presents data on how females sample males and discuss the number of cues females use. We conclude that sexual-selection studies have paid far less attention to variation among females than to variation among males, and that there is still much to learn about how females choose males and why different females make different choices. We suggest a number of possible lines for future research.

Animal Communication↗

Female copying increases the variance in male mating success.

Theoretical models of sexual selection assume that females choose males independently of the actions and choice of other individual females. Variance in male mating success in promiscuous species is thus interpreted as a result of phenotypic differences among males which females perceive and to which they respond. Here we show that, if some females copy the behavior of other females in choosing mates, the variance in male mating success and therefore the opportunity for sexual selection is greatly increased. Copying behavior is most likely in non-resource-based harem and lek mating systems but may occur in polygynous, territorial systems as well. It can be shown that copying behavior by females is an adaptive alternative to random choice whenever there is a cost to mate choice. We develop a statistical means of estimating the degree of female copying in natural populations where it occurs.

Animals↗

Repeatability of extra-pair mating in tree swallows.

Models of sexual selection assume that female mating preferences are heritable and, thus, repeatable for individual females across multiple mating episodes. Previous studies of the repeatability of female preference have examined individuals in captivity and focused presumably on social mate choice. However, extra-pair mating is widespread and can also influence sexual selection. We examined the repeatability of extra-pair mating in a wild population of tree swallows (Tachycineta bicolor) by experimentally inducing females to lay two clutches in rapid succession within the same season. We found that the proportion of extra-pair young and the number of extra-pair sires were highly repeatable for individual females. However, the repeatability of specific extra-pair sires was low. We suggest that this unusual pattern of mating may be due to females maximizing the heterozygosity of their offspring.

Animals↗

Convergent evolution of sexual shape dimorphism in Diptera.

Several patterns of sexual shape dimorphism, such as male body elongation, eye stalks, or extensions of the exoskeleton, have evolved repeatedly in the true flies (Diptera). Although these dimorphisms may have evolved in response to sexual selection on male body shape, conserved genetic factors may have contributed to this convergent evolution, resulting in stronger phenotypic convergence than might be expected from functional requirements alone. I compared phenotypic variation in body shape in two distantly related species exhibiting sexually dimorphic body elongation: Prochyliza xanthostoma (Piophilidae) and Telostylinus angusticollis (Neriidae). Although sexual selection appears to act differently on male body shape in these species, they exhibited strikingly similar patterns of sexual dimorphism. Likewise, patterns of within-sex shape variation were similar in the two species, particularly in males: relative elongation of the male head capsule, antenna, and legs was associated with reduced head capsule width and wing length, but was nearly independent of variation in thorax length. However, the two species presented contrasting patterns of static allometry: male sexual traits exhibited elevated allometric slopes in T. angusticollis, but not in P. xanthostoma. These results suggest that a shared pattern of covariation among traits may have channeled the evolution of sexually dimorphic body elongation in these species. Nonetheless, static allometries may have been shaped by species-specific selection pressures or genetic architectures.

Animals↗

Females avoid manipulative males and live longer.

Female mate choice has been demonstrated in a wide variety of species and is now accepted as an important factor in sexual selection. One of the remaining questions, however, is why females prefer specific males. Do females or their offspring benefit from their choice? Or do females choose mates to minimize costs of mating? Here we show that, in the ovoviviparous cockroach Nauphoeta cinerea, where sexual selection has been well documented, females chose mates to avoid costly male manipulation. Females were partnered with preferred or nonpreferred mates, and fitness of the females measured. We found that females lived longer when they mated with preferred males. Female lifespan depended on the rate at which offspring developed from egg to parturition: slower development led to longer life. We manipulated the male pheromone and showed that the component of the pheromone blend that makes males attractive to females also delayed parturition. Thus, like other aspects of sexual conflict in this species, offspring development and thereby the mother's lifespan depended on exposure of females to specific components of the male pheromone. Males benefit from manipulating offspring development because females with accelerated parturition remained unreceptive whereas females with slower developing offspring readily remated after giving birth to their offspring. Our results suggest a hormone-like role for the male pheromone in N. cinerea and provide the first direct evidence of mate choice to avoid male manipulation. This study shows that dominant males may not be preferred males if they are manipulating females, why multiple components with contrasting effects can exist in a sexual signal, and emphasizes the complex fitness relationships that can arise in species with sexual conflict.

Animals↗

Experimental manipulation of male behaviour during copulation in Stenomacra marginella (Heteroptera: Largidae): effect on copula duration, female remating and oviposition.

Apparently stimulatory male copulatory behaviour (MCB) is widespread among arthropods and it could help males to increase their fitness by inducing favourable behavioural and physiological changes in females. The empirical study of female responses to MCB is hindered because its experimental manipulation is difficult. We have developed a technique for reducing, with minimal disturbance, the frequency of MCB in the true bug Stenomacra marginella. Here, we test the idea that, in a polygamous species like S. marginella, sexual selection favours males whose MCB induces females to increase copula duration (thereby increasing the amount of sperm and accessory substances transferred), reduce their sexual receptivity to additional males and increase their rate of oviposition. Males prevented from performing MCB increased their rate of attempts to perform MCB. Copulations with previously mated females were of longer duration than those with virgin females, probably as a male adaptation for sperm competition, and MCB could have played a role in inducing this effect. Partial or total experimental reduction of MCB frequency had no effect on remating rates, because most females accepted remating at the first opportunity (1 day after their first copula). The probability of egg laying was reduced in females whose first mate was partially prevented from performing copulatory courtship, but not in females whose first mate was completely prevented from performing copulatory courtship. This is an intriguing result and further experiments are needed to understand its causes. We hypothesize that MCB evolved as a result of sexual selection.

Animals↗

Female barn owls (Tyto alba) advertise good genes.

The good genes hypothesis of sexual selection postulates that ornamentation signals superior genetic quality to potential mates. Support for this hypothesis comes from studies on male ornamentation only, while it remains to be shown that female ornamentation may signal genetic quality as well. Female barn owls (Tyto alba) display more black spots on their plumage than males. The expression of this plumage trait has a genetic basis and it has been suggested that males prefer to mate with females displaying more black spots. Given the role of parasites in the evolution of sexually selected traits and of the immune system in parasite resistance, we hypothesize that the extent of female plumage 'spottiness' reflects immunological defence. We assessed the genetic variation in specific antibody production against a non-pathogenic antigen among cross-fostered nestlings and studied its covariation with the plumage spottiness of genetic parents. The magnitude of the antibody response was positively correlated with the plumage spottiness of the genetic mother but not of the genetic father. Our study thereby provides the first experimental support, to our knowledge, for the hypothesis that female ornamentation signals genetic quality.

Animals↗

Heightened condition dependence is not a general feature of male eyespan in stalk-eyed flies (Diptera: Diopsidae).

Stalk-eyed flies are exemplars of sexual selection leading to the evolution of exaggerated male ornaments (eyespan). In Sphyracephala beccarri, there is no evidence for female mate choice for exaggerated male eyespan and only minor sex differences in eyespan. We used S. beccarri to test whether heightened condition dependence only evolves when male eyespan becomes sexually exaggerated. Male eyespan showed heightened condition dependence under food stress compared with a control trait (wing length). However, female eyespan displayed a similar pattern and there was no sex difference in the degree of increased eyespan sensitivity. The finding that eyespan is a sensitive indicator of food stress, even in an unexaggerated state, suggests that this may have acted as a pre-adaptation to the role of eyespan in sexual signalling in other Diopsid species. These results are consistent with handicap theory and Fisher's view of how sexual selection is initiated.

Analysis of Variance↗

Unusually dynamic sex roles in a fish.

Sex roles are typically thought of as being fixed for a given species. In most animals males compete for females, whereas the females are more reluctant to mate. Therefore sexual selection usually acts most strongly on males. This is explained by males having a higher potential reproductive rate than females, leading to more males being sexually active (a male-biased operational sex ratio). However, what determines sex roles and the strength of sexual selection is a controversial and much debated question. In this large-scale field study, we show a striking temporal plasticity in the mating competition of a fish (two-spotted goby, Gobiusculus flavescens). Over the short breeding season fierce male-male competition and intensive courtship behaviour in males were replaced by female-female competition and actively courting females. Hence, sex role reversal occurred rapidly. This is the first time that a shift in sex roles has been shown in a vertebrate. The shift might be explained by a large decline in male abundance, strongly skewing the sex ratio towards females. Notably, the sex role reversal did not occur at an equal operational sex ratio, contrary to established sex role theory.

Animals↗

Fixation of advantageous alleles in partially self-fertilizing populations. The effect of different selection modes.

The expected fixation probability of an advantageous allele was examined in a partially self-fertilizing hermaphroditic plant species using the diffusion approximation. The selective advantage of the advantageous allele was assumed to be increased viability, increased fecundity, or an increase in male fitness. The mode of selection, as well as the selfing rate, the population size, and the dominance of the advantageous allele, affect the fixation probability of the allele. In general it was found that increases in selfing rate decrease the fixation probability under male sexual selection, increase fixation probability under fecundity selection, and increase when recessive and decrease when dominant under viability selection. In some cases the highest fixation probability of advantageous alleles under fecundity or under male sexual selection occurred at an intermediary selfing rate. The expected mean fixation times of the advantageous allele were also examined using the diffusion approximation.

Alleles↗

Incest: the psychological problem and the biological contradiction.

We have yet to map an "inbreeding avoider" gene or design a psychological test that can accurately diagnose incest transgressors. Until those events occur, sociobiology, nursing, and psychology must explain phenomena, such as the incest taboo, with a theoretical language of supposition. It is in this light that the etiology of taboo is analyzed. This article proposes that a propensity for sexual selection originates in the gene system, and what becomes taboo is acquired through the learning that accompanies the experiences of the individual and culture when sexual selection occurs. Influences of the genetic system and psychological culture on the existence of a taboo are contrasted. The conclusion posits an eclectic taboo hypothesis, derived from the congruencies found in the literature, and suggests how this hypothesis can guide nursing interventions with incestual families.

Adult↗

Genetic covariance between indices of body condition and immunocompetence in a passerine bird.

BACKGROUND: Condition-dependence is a ubiquitous feature of animal life histories and has important implications for both natural and sexual selection. Mate choice, for instance, is typically based on condition-dependent signals. Theory predicts that one reason why condition-dependent signals may be special is that they allow females to scan for genes that confer high parasite resistance. Such explanations require a genetic link between immunocompetence and body condition, but existing evidence is limited to phenotypic associations. It remains unknown, therefore, whether females selecting males with good body condition simply obtain a healthy mate, or if they acquire genes for their offspring that confer high immunocompetence. RESULTS: Here we use a cross-foster experimental design to partition the phenotypic covariance in indices of body condition and immunocompetence into genetic, maternal and environmental effects in a passerine bird, the zebra finch Taeniopygia guttata. We show that there is significant positive additive genetic covariance between an index of body condition and an index of cell-mediated immune response. In this case, genetic variance in the index of immune response explained 56% of the additive genetic variance in the index of body condition. CONCLUSION: Our results suggest that, in the context of sexual selection, females that assess males on the basis of condition-dependent signals may gain genes that confer high immunocompetence for their offspring. More generally, a genetic correlation between indices of body condition and imuunocompetence supports the hypothesis that parasite resistance may be an important target of natural selection. Additional work is now required to test whether genetic covariance exists among other aspects of both condition and immunocompetence.

Animals↗

Positive selection in the egg receptor for abalone sperm lysin.

The mechanism of speciation is a central problem in evolutionary biology. In free-spawning animals with no complex mating behavior, prezygotic reproductive isolation (speciation) could result from the rapid divergence of genes coding for sperm and egg proteins that bind each other during fertilization. In abalone, sperm lysin evolves rapidly by positive Darwinian selection. The egg vitelline envelope receptor for lysin had previously been shown to evolve neutrally and be subjected to concerted evolution. Several mathematical simulations predict that both male and female reproductive proteins should evolve rapidly by positive selection. Here we report that the sequence diversity of the amino-terminal end of the egg vitelline envelope receptor for lysin has been promoted by positive Darwinian selection. These data provide molecular support for theoretical models showing that the two sexes are locked in a "coevolutionary chase" that could be driven by processes such as sexual selection, sexual conflict, or microbial attack (pathogen avoidance). The result of this continuous coevolution of the gamete recognition system could be the splitting of one population into two that are reproductively isolated (speciation).

Animals↗

Polyandrous females discriminate against previous mates.

In most animal species, particularly those in which females engage in polyandry, mate choice is a sequential process in which a female must choose to mate or not to mate with each male encountered. Although a number of theoretical and empirical investigations have examined the effects of sequential mate choice on the operation of sexual selection, how females respond to solicitation by previous mates has received little attention. Here, we report the results of a study carried out on the polyandrous pseudoscorpion, Cordylochernes scorpioides, that assessed the sexual receptivity of once-mated females presented after a lapse of 1.5 hr or 48 hr with either their first mate or a different male. Females exhibited a high level of receptivity to new males, irrespective of intermating interval. By contrast, time between matings exerted a strong effect on female receptivity to previous mates. After a lapse of 48 hr, females did not differ significantly in their receptivity toward previous mates and different males, whereas at 1.5 hr after first mating, females were almost invariably unreceptive to males from whom they had previously accepted sperm. This result could not be attributed to male size or mating experience or to male sexual receptivity. Indeed, males were as willing to transfer sperm to a previous mate as they were to a new female. This difference between males and females in their propensity to remate with the same individual may reflect a conflict between the sexes, with males seeking to minimize postcopulatory sexual selection and females actively keeping open the opportunity for sperm competition and female choice of sperm by discriminating against previous mates.

Animals↗