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Mate choice: from sexual cues to cognitive adaptations.

Evolutionary psychologists have successfully combined sexual selection theory and empirical research to compile lists of sexual attractiveness cues used in human mate choice. But a list of inputs is not the same as a normative or descriptive model of a psychological adaptation. We need to shift from cataloguing sexual cues to modelling cognitive adaptations for mate choice. This theoretical chapter addresses how to make this transition in three parts. The introduction discusses four general problems with cue cataloguing as an evolutionary psychology research strategy: animals' promiscuous flexibility of cue use; cue use being marginal to cognition; cue use being marginal to the hard game-theoretical aspects of mate choice; and cue use being uninformative about the exact adaptive functions of mate choice. The middle section develops six critiques of current mate choice research: the obsession with sex difference; the over-emphasis on physical rather than behavioural cues; the assumption of weighted linear models of cue integration; the avoidance of game-theoretical problems of mutual choice and assortative mating; the neglect of co-evolution between mate choice heuristics and the cues that they select; and the failure to understand that mate choice is only worth doing if potential mates show significant genetic variance. The conclusion outlines a new normative and descriptive framework for mate choice, centred on the use of brutally efficient search heuristics that exploit the informational structure of human genotypes, phenotypes and populations to make good mate choices.

Adaptation, Psychological↗

Characterization of courtship sounds of species of the subgroup fasciola (Diptera, Drosophilidae, Drosophila repleta group): interspecific and interpopulational analyses.

The aim of this work was to characterize the male courtship song pattern of various species of the fasciola subgroup and to determine the level of variation both within and among species. The parameters analyzed were intrapulse interval (PI), interpulse interval (IPI), and intrapulse frequency (IF). Six different species were analyzed: D. coroica (three populations), D. ellisoni, D. fascioloides, D. moju, D. onca, and D. rosinae (one population each). There were significant differences among the six species for these three courtship song parameters. The IPI was the most variable parameter among these species, suggesting that this parameter is important for female discrimination. Four different hypotheses could explain this variation: 1. different selection pressures with absence of flow gene; 2. intraspecific sexual selection; 3. sympatric effects on song evolution; and 4. genetic drift. The PI was the only parameter that was significantly different among the three population of D. coroica. Low variability among populations within the same species was already observed for other subgroups and could be explained by the following hypotheses: strong selection acting on the song parameters, gene flow, or recent colonization from a common source. Additional studies of the courtship song of other species of the fasciola subgroup, as well as for other subgroups of the repleta group, and studies, using molecular makers, that focus on the genetic basis of the differences among these species in courtship song would allow us to evaluate the association of courtship song and sexual isolation in these species, and would also help us to understand the evolution of these behavioural differences.

Animals↗

Immune suppression and the cost of reproduction in the ground cricket, Allonemobius socius.

One of the most common life history trade-offs in animals is the reduction in survivorship with increasing reproductive effort. Despite the prevalence of this pattern, its underlying physiological mechanisms are not well understood. Here we test the hypothesis that immune suppression mediates this phenotypic trade-off by manipulating reproductive effort and measuring immune function and mortality rates in the striped ground cricket, Allonemobius socius. Because A. socius males provide females with a hemolymph-based nuptial gift during copulation, and many structural components of immunity reside in the hemolymph, we also predicted that sexual selection may differentially affect how disease resistance evolves in males and females. We found that an increased mating effort resulted in a reduced immune ability, coupled with an increased rate in age-specific mortality for both sexes. Thus, immune suppression appears to be a link between reproductive effort and cost in this system. In addition, males and females appeared to differentially invest in several aspects of immunity prior to mating, with males exhibiting a higher concentration of circulating hemocytes and a superior bacterial defense capability. This pattern may be the result of previously established positive selection on gift size due to its affect on female fecundity. In short, female choice for larger gifts may lead to a sexually dimorphic immune ability.

Age Factors↗

Potential fitness benefits from mate selection in the Atlantic cod (Gadus morhua).

Little evidence of benefits from female mate choice has been found when males provide no parental care or resources. Yet, good genes models of sexual selection suggest that elaborated male sexual characters are reliable signals of mate quality and that the offspring of males with elaborate sexual ornaments will perform better than those of males with less elaborate ornaments. We used cod (Gadus morhua L.), an externally fertilizing species where males provide nothing but sperm, to examine the potential of optimal mate selection with respect to offspring survival. By applying in vitro fertilizations, we crossed eight females with nine males in all possible combinations and reared each of the 72 sib groups. We found that offspring survival was dependent on which female was mated with which male and that optimal mate selection has the potential to increase mean offspring survival from 31.9 to 55.6% (a 74% increase). However, the size of the male sexual ornaments and sperm quality (i.e. sperm velocity and sperm density) could not predict offspring survival. Thus, even if there may be large fitness benefits of mate selection, we might not yet have identified the male characteristics generating high offspring survival.

Animals↗

Studies of human physique and sexual attractiveness: sexual preferences of men and women in China.

Men and women at Northwest University (n = 631), Xi'an, China, were asked to rate the attractiveness of male or female figures manipulated to vary somatotype, waist-to-hip ratio (WHR), secondary sexual traits, and other features. In study 1, women rated the average masculine somatotype as most attractive, followed by the mesomorphic (muscular), ectomorphic (slim), and endomorphic (heavily built) somatotypes, in descending order of preference. In study 2, the amount and distribution of masculine trunk (chest and abdominal) hair were altered progressively in a series of front-posed figures. Women rated figures with no or little trunk hair as most attractive. Study 3 assessed the attractiveness of front-posed male figures which varied only in length of their nonerect penis. Numerical ratings for this trait were low, but moderate lengthening of the penis (22% or 33% above average) resulted in a significant increase in scores for attractiveness. In study 4, Chinese men rated the attractiveness of back-posed female images varying in waist-to-hip ratio (WHR from 0.5-1.0). The 0.6 WHR figure was most preferred, followed by 0.7, while figures with higher ratios (0.9 or 1.0) were significantly less attractive. Study 5 rated the attractiveness of female skin color: men expressed a marked preference for images which were lighter in color, as compared to images of average or darker skin colors. These results, the first of their kind reported for a Chinese population, support the view that sexual selection has influenced the evolution of human physique and sexual attractiveness in men and women.

Asian People↗

Physiological suppression of sexual function of subordinate males: a subtle form of intrasexual competition among male sifakas (Propithecus verreauxi)?

In contrast to most anthropoid primates, sifakas (Propithecus verreauxi), like many group-living lemurs, exhibit a number of features that deviate from predictions of sexual selection theory. Despite a promiscuous mating system, they lack sexual dimorphism, suggesting that physical combat plays only a minor role in intrasexual competition for receptive females. In this study, we investigated the hypothesis that socioendocrinological mechanisms contribute to suppression of reproductive function of subordinate males. For that purpose, 10 male sifakas from five social groups were observed for 669 focal animal hours for 4 months, including the mating season, in Western Madagascar. Concomitantly 315 fecal samples of these animals were collected and the concentration of immunoreactive testosterone was quantified hy enzymeimmunoassay procedures. Clear dominance relationships existed among coresident males. Testosterone levels of dominant males were significantly higher than those of subordinates during, as well as outside, the mating season. Additionally, the increase in testosterone levels prior to the mating season was more pronounced for dominant than for subordinate males. These findings are in accordance with the hypothesis of suppression of sexual function of subordinate males, probably providing dominant males with ani advantage in sperm competition. If reproductive success is mainly determined by this nonagonistic form of intrasexual competition, the results of this study contribute an important piece to the puzzle of lacking sexual dimorphism in P. verreauxi.

Adaptation, Physiological↗

Sexual swellings advertise female quality in wild baboons.

The females of many Old World primate species produce prominent and conspicuous swellings of the perineal skin around the time of ovulation. These sexual swellings have been proposed to increase competition among males for females or to increase the likelihood of a female getting fertilized, by signalling either a female's general reproductive status, or the timing of her ovulation. Here we show that sexual swellings in wild baboons reliably advertise a female's reproductive value over her lifetime, in accordance with a theoretical model of honest signalling. Females with larger swellings attained sexual maturity earlier, produced both more offspring and more surviving offspring per year than females with smaller swellings, and had a higher overall proportion of their offspring survive. Male baboons use the size of the sexual swelling to determine their mating effort, fighting more aggressively to consort females with larger swellings, and spending more time grooming these females. Our results document an unusual case of a sexually selected ornament in females, and show how males, by mating selectively on the basis of the size of the sexual swelling, increase their probability of mating with females more likely to produce surviving offspring.

Animals↗

The origins of sexual dimorphism in body size in ungulates.

Jarman (1974) proposed a series of relationships between habitat use, food dispersion, and social behavior and hypothesized a series of evolutionary steps leading to sexual dimorphism in body size through sexual selection in African antelope species. The hypothesis states that sexual size dimorphism evolved in a three-step process. Initially, ancestral monomorphic and monogamous ungulate species occupying closed habitats radiated into open grassland habitats. Polygynous mating systems then rapidly evolved in response to the aggregation of males and females, perhaps in relation to the clumped distribution of food resources in open habitats. Subsequently, size dimorphism evolved in those species occupying open habitats, but not in species that remained in closed habitats or retained monogamy. This hypothesis has played an important role in explaining the origins of sexual dimorphism in mammals. However, the temporal sequence of the events that Jarman proposed has never been demonstrated. Here we use a phylogeny of extant ungulate species, along with maximum-likelihood statistical techniques, to provide a test of Jarman's hypothesis.

Animals↗

Mating success of wild type and sepia mutants Drosophila melanogaster in different choice.

Mating behaviour of red-eyed (wt) and brown-eyed (sepia) Drosophila melanogaster was studied under light conditions. Mating success was directly observed in mating vials and techniques usually applied in the studies of sexual selection ("female choice" and "multiple choice"). The comparison of sexual activity of mutant and wild types clearly indicates that they are not equally successful in matings. Sepia eye colour mutation decreases sexual activity of Drosophila melanogaster males, influences the preference ability of females and decreases the number of progeny from homogamic mating of the se x se type, as well as from heterogamic copulations in which sepia females take part. Non-random mating of wild type males and sepia females (in "multiple-choice" situation), with genetically and phenotypically different individuals, could be another mechanism for conservation of genetic polymorphism in natural populations.

Animals↗

Rapid evolution of reproductive proteins in abalone and Drosophila.

Observations from different taxa, including plants, protozoa, insects and mammals, indicate that proteins involved in reproduction evolve rapidly. Several models of adaptive evolution have been proposed to explain this phenomenon, such as sexual conflict, sexual selection, self versus non-self recognition and pathogen resistance. Here we discuss the potential role of sexual conflict in the rapid evolution of reproductive genes in two different animal systems, abalone (Haliotis) and Drosophila. In abalone, we reveal how specific interacting sperm-egg proteins were identified and discuss this identification in the light of models for rapid protein evolution and speciation. For Drosophila, we describe the genomic approaches taken to identify male accessory gland proteins and female reproductive tract proteins. Patterns of protein evolution from both abalone and Drosophila support the predicted patterns of rapid protein evolution driven by sexual conflict. We stress however that other selective pressures may contribute to the rapid evolution that is observed. We conclude that the key to distinguishing between sexual conflict and other mechanisms of protein evolution will be an integration of genetic, experimental and theoretical data.

Animals↗

Effects of sexual dimorphism on facial attractiveness.

Testosterone-dependent secondary sexual characteristics in males may signal immunological competence and are sexually selected for in several species. In humans, oestrogen-dependent characteristics of the female body correlate with health and reproductive fitness and are found attractive. Enhancing the sexual dimorphism of human faces should raise attractiveness by enhancing sex-hormone-related cues to youth and fertility in females, and to dominance and immunocompetence in males. Here we report the results of asking subjects to choose the most attractive faces from continua that enhanced or diminished differences between the average shape of female and male faces. As predicted, subjects preferred feminized to average shapes of a female face. This preference applied across UK and Japanese populations but was stronger for within-population judgements, which indicates that attractiveness cues are learned. Subjects preferred feminized to average or masculinized shapes of a male face. Enhancing masculine facial characteristics increased both perceived dominance and negative attributions (for example, coldness or dishonesty) relevant to relationships and paternal investment. These results indicate a selection pressure that limits sexual dimorphism and encourages neoteny in humans.

Adult↗

Fisherian and Wrightian theories of speciation.

Fisher's theory of sexual selection, Wright's shifting-balance theory, and recent models based on them are reviewed as mechanisms of animal speciation. The joint evolution of mating preferences and secondary sexual characters can cause rapid nonadaptive phenotypic divergence and premating isolation between geographically separated populations, or along a cline. Extensive comparative data on Drosophila species support the suggestion of R. A. Fisher and T. Dobzhansky that the evolution of mating preferences can reinforce partial postmating isolation between sympatric populations. The interaction of natural selection and random genetic drift in local populations with a small effective size can produce a rapid transition between relatively stable phenotypes separated by an adaptive valley, or between chromosomal rearrangements with a heterozygote disadvantage. Large demographic fluctuations, such as frequent random local extinction and colonization, are required for the rapid spread of new adaptations (or karyotypes) when intermediate phenotypes (or rearrangement heterozygotes) are selected against.

Animals↗

Facial shape and judgements of female attractiveness.

The finding that photographic and digital composites (blends) of faces are considered to be attractive has led to the claim that attractiveness is averageness. This would encourage stabilizing selection, favouring phenotypes with an average facial structure. The 'averageness hypothesis' would account for the low distinctiveness of attractive faces but is difficult to reconcile with the finding that some facial measurements correlate with attractiveness. An average face shape is attractive but may not be optimally attractive. Human preferences may exert directional selection pressures, as with the phenomena of optimal outbreeding and sexual selection for extreme characteristics. Using composite faces, we show here that, contrary to the averageness hypothesis, the mean shape of a set of attractive faces is preferred to the mean shape of the sample from which the faces were selected. In addition, attractive composites can be made more attractive by exaggerating the shape differences from the sample mean. Japanese and caucasian observers showed the same direction of preferences for the same facial composites, suggesting that aesthetic judgements of face shape are similar across different cultural backgrounds. Our finding that highly attractive facial configurations are not average shows that preferences could exert a directional selection pressure on the evolution of human face shape.

Adult↗

Intralocus sexual conflict can drive the evolution of genomic imprinting.

Genomic imprinting is a phenomenon whereby the expression of an allele differs depending upon its parent of origin. There is an increasing number of examples of this form of epigenetic inheritance across a wide range of taxa, and imprinting errors have also been implicated in several human diseases. Various hypotheses have been put forward to explain the evolution of genomic imprinting, but there is not yet a widely accepted general hypothesis for the variety of imprinting patterns observed. Here a new evolutionary hypothesis, based on intralocus sexual conflict, is proposed. This hypothesis provides a potential explanation for much of the currently available empirical data, and it also makes new predictions about patterns of genomic imprinting that are expected to evolve but that have not, as of yet, been looked for in nature. This theory also provides a potential mechanism for the resolution of intralocus sexual conflict in sexually selected traits and a novel pathway for the evolution of sexual dimorphism.

Animals↗

Reconstructing the evolution of complex bird song in the oropendolas.

The elaborate songs of songbirds are frequent models for investigating the evolution of animal signals. However, few previous studies have attempted to reconstruct historical changes in song evolution using a phylogenetic comparative approach. In particular, no comparative studies of bird song have used a large number of vocal characters and a well-supported, independently derived phylogeny. We identified 32 features in the complex vocal displays of male oropendolas (genera Psarocolius, Gymnostinops, and Ocyalus) that are relatively invariant within taxa and mapped these characters onto a robust molecular phylogeny of the group. Our analysis revealed that many aspects of oropendola song are surprisingly evolutionarily conservative and thus are potentially useful characters for reconstructing historical patterns. Of the characters that varied among taxa, nearly two thirds (19 of 29) showed no evidence of evolutionary convergence or reversal when mapped onto the tree, which was reflected in a high overall consistency index (CI = 0.78) and retention index (RI = 0.88). Some reconstructed patterns provided evidence of selection on these signals. For example, rapid divergence of the songs of the Montezuma oropendola, Gymnostinops montezuma, from those of closely related taxa suggests the recent influence of strong sexual selection. In general, our results provide insights into the mode of vocal evolution in songbirds and suggest that complex vocalizations can provide information about phylogeny. Based on this evidence, we use song characters to estimate the phylogenetic affinities of three oropendola taxa for which molecular data are not yet available.

Animals↗

Sexual attitudes and their relation to cervical intraepithelial neoplasia in Maiduguri, Nigeria.

Five hundred randomly selected sexually active women attending antenatal, postnatal, family planning and gynaecological clinics in the Department of Obstetrics and Gynaecology, University of Maiduguri Teaching Hospital, had questionnaires administered on their sexual attitudes/ behaviour and their 'pap'smears taken for cytological assessment. The aim was to identify any significant association between sexual attitudes/behaviour and cervical intraepithelial neoplasia (CIN) for the purpose of targeting the population most at risk. Ninety-one women had varying degrees of CIN, a prevalence rate of 18.2%. However, multiple sexual partners was the only factor incriminated to significantly increase the risk of occurrence of CIN. Targeting women with multiple sexual partners may help in directing limited screening resources to those more likely to be positive.

Journal Article↗

Why humans need type 5 phosphodiesterase inhibitors.

Erectile dysfunction (ED) is a widespread medical condition affecting millions of males at any age and requiring medical treatment. ED may simply reflect a limit of human physiology, yet ED equates to genetic death and the high prevalence of ED is a clear evolutionary paradox. Why is a condition which totally blocks reproduction so widespread? Epidemiology shows that impotence is a common symptom of almost all major diseases and male reproductive physiology is sensitive to general health and to environmental, psychological, and physical stressors. Moreover, erectile dysfunction is a predictor of myocardial infarction and stroke. Briefly, efficient erection is a marker of good health and good health prognosis. In the animal kingdom, mate choice is often based on extravagant and cumbersome physical traits (such as the peacock's tail). These traits, that are necessary for gene propagation, are at the same time efficient handicaps which reduce the survival of the carrier. What animal studies show, is that these handicaps can function as honest indicators of the individual's fitness. In other words, their expression is condition-dependent and only individuals with high phenotypic quality present the trait. By mating with males which express the trait, females indirectly select for individuals of superior fitness which will be inherited by the offspring. Erection is very clearly a condition-dependent trait in the human species. We suggest that the fragility of male sexual physiology is a sexually selected handicap which hampers the reproduction of individuals with lower phenotypic quality.

Biological Evolution↗

Sexually transmitted disease in birds: occurrence and evolutionary significance.

Sexually transmitted diseases (STDS) span two current areas of sexual selection theory, namely the roles of multiple mating in determining individual reproductive success, and of parasites in mate choice, yet have been relatively neglected in the ecological literature. I reviewed the occurrence of STDS in populations of commercially kept birds and found widespread evidence for the existence of pathogenic STDS in such populations. STDs may have important consequences for the evolution of behaviour, reproductive physiology and some secondary sexual characteristics. Where STDS are costly they are hypothesized to affect the evolution of mating systems, and, via selection for hostility in the female reproductive tract, to explain high levels of sperm mortality after insemination. The potential for coevolutionary cycling is large, as some STDS may coevolve with female and male reproductive physiology, which may themselves coevolve. Although little information currently exists concerning the occurrence of STDS in wild birds, techniques for their identification are well established. This study raises a number of testable predictions about the consequences of STDS for avian reproductive biology, and I suggest that STDS should be considered as a potentially powerful factor in future studies of mate choice and sperm competition.

Animals↗