Conditioning and sexual behavior: a review

Sexual behavior is directed by a sophisticated interplay between steroid hormone actions in the brain that give rise to sexual arousability and experience with sexual reward that gives rise to expectations of competent sexual activity, sexual desire, arousal, and performance. Sexual experience allows animals to form instrumental associations between internal or external stimuli and behaviors that lead to different sexual rewards. Furthermore, Pavlovian associations between internal and external stimuli allow animals to predict sexual outcomes. These two types of learning build upon instinctual mechanisms to create distinctive, and seemingly "automated," patterns of sexual response. This article reviews the literature on conditioning and sexual behavior with a particular emphasis on incentive sequences of sexual behavior that move animals from distal to proximal with regard to sexual stimuli during appetitive phases of behavior and ultimately result in copulatory interaction and mating during consummatory phases of behavior. Accordingly, the role of learning in sexual excitement, in behaviors that bring about the opportunity to mate, in courtship and solicitation displays, in sexual arousal and copulatory behaviors, in sexual partner preferences, and the short- and long-term influence of copulatory experience on sexual and reproductive function is examined. Although hormone actions set the stage for sexual activity by generating the ability of animals to become sexually excited and aroused, it is each animal's unique experience with sexual behavior and sexual reward that molds the strength of responses made toward sexual incentives.     

The association of sexual interest and sexual behaviors among adolescent women: a daily diary perspective

Theoretical and empirical linkages of adult women's sexual interest and sexual behaviors are relatively well-established, but few data address similar issues in adolescent women. This paper reviews data from published reports of associations of adolescent women's sexual interest and various sexual behaviors. All of the papers reported data collected from a single longitudinal cohort of young women. The primary source of data collection was daily diaries, allowing close temporal pairing of sexual interest with sexual behaviors. Young women's sexual interest on a given day was consistently and independently associated with sexual activity on that day, whether the behavior was first lifetime coitus, coitus, fellatio, cunnilingus, anal intercourse, or coitus during menses. We also found no evidence of influence of hormonal contraceptives on young women's sexual interest. Taken together, these data demonstrate the relevance of sexual interest as a key factor in young women's sexuality and sexual behavior.     

性选择与性冲突理论在植物繁殖生态学中的应用与进展

性选择与性冲突是植物繁殖性状多样性及性系统演化的重要动力, 二者密切相关却又有所区别, 理解它们的作用机制及其影响对于植物繁殖生态学的研究具有重要意义。当前, 性选择与性冲突理论在植物繁殖生态学中的运用已取得长足进展, 但国内相关研究较少, 对该领域关注不够。因此, 该文对该领域的基本理论和研究进展进行了综述。首先, 阐述性选择与性冲突理论在植物研究中的发展及其运用基础; 其次, 分别从授粉前和授粉后两个阶段详细介绍性选择与性冲突在有花植物繁殖过程中的作用机制及其影响, 并指出环境因素对它们所产生的影响; 最后, 对当前研究存在的不足及该领域未来的研究方向进行总结和展望。希望以此增强人们对性选择和性冲突理论的认识, 促进其在植物繁殖生态学中的运用与发展。     

Evolution of cyclic sexual reproduction under host-parasite interactions

Alternate changes of sexual and asexual generations was studied theoretically by numerical analyses, from the viewpoint of host-parasite infective interactions. The host species was considered a diploid organism, characterized by two loci determining parasite resistance type and sexual strategy, between which a linkage exists to a certain degree. The sexual reproductive strategy was assumed to be determined by three alleles: asexual, complete sexual and cyclic sexual alleles. The effect of the parasites was represented by decreasing fitness functions of each host-type frequency. Numerical analyses of various linkages between genes and frequency dependence of host fitness revealed that the dynamics varied depending on whether the cyclic sexual allele was dominant or recessive against the complete sexual allele. When the cyclic sexual allele was dominant, the cyclic sexual strain persisted under intermediate frequency dependence of host fitness. On the other hand, when the cyclic sexual allele was recessive, it always tended to spread and to be maintained in the population. In such situations, both complete and cyclic sexual strains coexist, but the asexual strain is lost.     

Signal trait sexual dimorphism and mutual sexual selection in Drosophila serrata

Abstract The evolution of sexual dimorphism may occur when natural and sexual selection result in different optimum trait values for males and females. Perhaps the most prominent examples of sexual dimorphism occur in sexually selected traits, for which males usually display exaggerated trait levels, while females may show reduced expression of the trait. In some species, females also exhibit secondary sexual traits that may either be a consequence of a correlated response to sexual selection on males or direct sexual selection for female secondary sexual traits. In this experiment, we simultaneously measure the intersex genetic correlations and the relative strength of sexual selection on males and females for a set of cuticular hydrocarbons in Drosophila serrata . There was significant directional sexual selection on both male and female cuticular hydrocarbons: the strength of sexual selection did not differ among the sexes but males and females preferred different cuticular hydrocarbons. In contrast with many previous studies of sexual dimorphism, intersex genetic correlations were low. The evolution of sexual dimorphism in D. serrata appears to have been achieved by sex-limited expression of traits controlled by genes on the X chromosome and is likely to be in its final stages.     

Sexual selection is positively associated with ecological generalism among agamid lizards

Natural and sexual selection shape the evolution of species but the interplay between them is poorly understood. Two phylogenetic studies on birds have suggested that species with greater sexual dichromatism have a broader habitat use. We show that in agamid lizards, species with more elaborate secondary sexual traits are also ecologically more opportunistic. Species with greater dimorphism in head size and ornamentation have greater altitudinal range and broader habitat use, respectively, and species with greater sexual dichromatism have wider microhabitat use. Body size was positively associated with sexual and ecological generalism, but associations between ecological and sexual traits remained after accounting for body size. We suggest that sexual and natural selection may be linked either because sexual selection can promote generalism at the population level by favouring 'good genes', or because higher population densities may be associated with both stronger sexual selection and broader resource use.     

Prevalence of sexual dysfunctions and other sexual disorders in adults with attention-deficit/hyperactivity disorder compared to the general population

Attention-deficit/hyperactivity disorder (ADHD) is characterized by symptoms of inattention and/or hyperactivity and impulsivity that lead to dysfunctioning in daily life. One of the affected areas of life that has so far not been studied in ADHD is sexual functioning. The goal of this study was to assess prevalence of sexual dysfunctions and other sexual disorders among adults with ADHD. A total of n = 136 adult patients treated in a Dutch outpatient ADHD clinic filled out two questionnaires to screen for sexual dysfunctions and other sexual disorders. We compared the prevalence of sexual dysfunctions and other sexual disorders in our ADHD patient population to results from two large surveys among the general Dutch population. We found that 39% of the male and 43% of the female ADHD patients had symptoms of a sexual dysfunction, and 17% of the male and 5% of the female ADHD patients had symptoms of any other sexual disorder. Only one male patient had received a diagnosis of a sexual disorder at this clinic prior to study participation. In conclusion, sexual dysfunctions and other sexual disorders are highly prevalent in adults with ADHD. Screening for sexual disorders should be therefore standard procedure during diagnostic assessment.     

The association between testosterone, sexual arousal, and selective attention for erotic stimuli in men

Twenty-six, eugonadal men between the ages of 18 and 27 participated in this investigation of the relationship between sexual arousal, testosterone (T) levels, and the processing of sexual information. At each of the two test sessions, subjects gave a blood sample, listened to an erotic or neutral priming audiotape, and completed a dichotic listening task designed to assess selective attention for sexual stimuli. Subjective levels of sexual arousal to the audiotape and sexual attitudes and sexual experience were assessed by self-report measures. Contrary to our hypothesis, there was no relationship between levels of free T and the strength of the selective attention bias for sexual stimuli. However, men who were more distracted by the sexual material in the task reported higher levels of sexual arousal to erotic imagery than men who were less distracted by the sexual material in the task (P less than 0.01). Moreover, men who were more sexually aroused by the erotic audiotape made significantly less shadowing errors in the erotic prime condition then they did during the neutral prime condition (P less than 0.05). There was a negative association between T and shadowing errors in the erotic prime condition (P less than 0.05). These results suggest that lower thresholds for sexual arousal are associated with a greater bias to attend to sexual information and that T may have effects on cognitive-motivational aspects of sexual behavior by enhancing attention to relevant stimuli.     

The role of sexual and natural selection in shaping patterns of sexual dichromatism in the largest family of songbirds (Aves: Thraupidae)

Males and females can be under different evolutionary pressures if sexual and natural selection is differentially operating in each sex. As a result, many species have evolved sexual dichromatism, or differences in coloration between sexes. Although sexual dichromatism is often used as an index of the magnitude of sexual selection, sexual dichromatism is a composite trait. Here, we examine the evolution of sexual dichromatism in one of the largest and most ecologically diverse families of birds, the tanagers, using the avian visual perspective and a species‐level phylogeny. Our results demonstrate that the evolutionary decreases of sexual dichromatism are more often associated with larger and more frequent changes in male plumage coloration, and evolutionary increases are not more often associated with larger changes in either sex. Furthermore, we show that the crown and ventral plumage regions are correlated with sexual dichromatism in males, and that only male plumage complexity is positively correlated with sexual dichromatism. Finally, we demonstrate that light environment is important in shaping both plumage brilliance and complexity. By conducting a multilevel analysis of plumage evolution in males and females, we show that sexual dichromatism evolves via a mosaic of sexual and natural selection in both sexes.     

Sexual behavior and sexual orientation of the female rat after hormonal treatment during various stages of development

The amount of circulating sex steroids during Postnatal Days 30-90 was varied in normally developed and in androgenized female rats. The influence of these manipulations on sexual behavior and sexual orientation was investigated. Normally developed or neonatally androgenized females were ovariectomized and implanted with estradiol through Postnatal Days 30-90 or sham-implanted. The remaining subjects were left intact during that period. The hormonal condition during Postnatal Days 30-90 significantly affected the behavior of normally developed females, but affected the behavior of neonatally androgenized females only to minor extent. Estrogen implants in normally developed females enhanced masculine sexual responses and induced a female-directed sexual orientation. Feminine sexual responses were unaffected by this treatment. Sham-implanted, normally developed females showed a male-directed sexual orientation and fewer masculine sexual responses than subjects which were left intact during Postnatal Days 30-90. Neonatal androgen treatment in general resulted in elevated levels of masculine Neonatal androgen treatment in general resulted in elevated levels of masculine sexual responses, inhibited feminine sexual behavior, and facilitated a female-directed sexual orientation.     

The evolution of condition-dependent sexual dimorphism

Theory suggests that the net benefit of allocating resources to a sexual trait depends both on the strength of sexual selection on that trait and on individual condition. This predicts a tight coevolution between sexual dimorphism and condition dependence and suggests that these patterns of within-sex and between-sex variation may share a common genetic and developmental basis. Although condition-dependent expression of sexual traits is widely documented, the extent of covariation between condition dependence and sexual dimorphism remains poorly known. I investigated the effects of condition (larval diet quality) on multivariate sexual dimorphism in the fly Telostylinus angusticollis (Neriidae). Condition determined the direction of sexual size dimorphism and modulated sexual shape dimorphism by affecting allometric slopes and/or intercepts of sexually homologous traits in both sexes. Although the greatest responses to condition manipulation were observed in male sexual traits, both sexual and nonsexual traits exhibited substantial variation in the nature and magnitude of condition effects. Nonetheless, condition dependence and sexual dimorphism were remarkably congruent: variation in the strength of condition effects on male traits explained more than 90% of the variation in the magnitude of sexual dimorphism, whether quantified in terms of trait size or allometric slope. The genetic mechanisms that give rise to multivariate sexual dimorphism in body shape thus function in a strongly condition-dependent manner in this species, suggesting a common genetic basis for body shape variation within and between sexes.     

Women's genital sexual arousal to oral versus penetrative heterosexual sex varies with menstrual cycle phase at first exposure

Reproductive-aged women show increased interest in sexual activity during the fertile phase of the menstrual cycle that can motivate sexual behavior and thereby increase the likelihood of conception. We examined whether women demonstrated greater sexual responses (subjective and genital sexual arousal) to penetrative versus oral sexual activities during the fertile versus non-fertile phases of their cycles, and whether women's arousal responses were influenced by the phase during which they were first exposed to these sexual stimuli (e.g., Slob et al., 1991; Wallen and Rupp, 2010). Twenty-two androphilic women completed two identical sexual arousal assessments in which genital responses were measured with a vaginal photoplethysmograph and their feelings of sexual arousal were recorded. Women viewed an array of 90 s films varying by couple type (female–female, male–male, female–male) and sexual activity type (oral or penetrative), during the fertile (follicular) and non-fertile (luteal) phases of their menstrual cycle, with the order of cycle phase at the first testing session counter-balanced. Women tested first in the fertile phase showed significantly greater genital arousal to female–male penetrative versus oral sex in both testing sessions, whereas self-reports of sexual arousal were not affected by cycle phase or testing order. These results contribute to a growing body of research suggesting that fertility status at first exposure to sexual stimuli has a significant effect on subsequent sexual responses to sexual stimuli, and that this effect may differ for subjective versus genital sexual arousal.     

Parallel evolution of sexual isolation in sticklebacks

Mechanisms of speciation are not well understood, despite decades of study. Recent work has focused on how natural and sexual selection cause sexual isolation. Here, we investigate the roles of divergent natural and sexual selection in the evolution of sexual isolation between sympatric species of threespine sticklebacks. We test the importance of morphological and behavioral traits in conferring sexual isolation and examine to what extent these traits have diverged in parallel between multiple, independently evolved species pairs. We use the patterns of evolution in ecological and mating traits to infer the likely nature of selection on sexual isolation. Strong parallel evolution implicates ecologically based divergent natural and/or sexual selection, whereas arbitrary directionality implicates nonecological sexual selection or drift. In multiple pairs we find that sexual isolation arises in the same way: assortative mating on body size and asymmetric isolation due to male nuptial color. Body size and color have diverged in a strongly parallel manner, similar to ecological traits. The data implicate ecologically based divergent natural and sexual selection as engines of speciation in this group.     

Sex Attracts: Investigating Individual Differences in Attentional Bias to Sexual Stimuli

We investigated the impact of sexual stimuli and the influence of sexual motivation on the performance in a dot-probe task and a line-orientation task in a large sample of males and females. All pictures (neutral, erotic) were rated on the dimensions of valence, arousal, disgust, and sexual arousal. Additionally, questionnaires measuring sexual interest/desire/motivation were employed. The ratings of the sexual stimuli point to a successful picture selection because sexual arousal did not differ between the sexes. The stimuli were equally arousing for men and women. Higher scores in the employed questionnaires measuring sexual interest/desire/motivation led to higher sexual arousal ratings of the sex pictures. Attentional bias towards sex pictures was observed in both experimental tasks. The attentional biases measured by the dot-probe and the line-orientation task were moderately intercorrelated suggesting attentional bias as a possible marker for a sex-attention trait. Finally, only the sexual sensation seeking score correlated with the attentional biases of the two tasks. Future research is needed to increase the predictive power of these indirect measures of sexual interest.     

Experimental evidence that sexual conflict influences the opportunity, form and intensity of sexual selection

Sexual interactions are often rife with conflict. Conflict between members of the same sex over opportunities to mate has long been understood to effect evolution via sexual selection. Although conflict between males and females is now understood to be widespread, such conflict is seldom considered in the same light as a general agent of sexual selection. Any interaction between males or females that generates variation in fitness, whether due to conflict, competition or mate choice, can potentially influence sexual selection acting on a range of male traits. Here we seek to address a lack of direct experimental evidence for how sexual conflict influences sexual selection more broadly. We manipulate a major source of sexual conflict in the black field cricket, Teleogryllus commodus, and quantify the resulting changes in the nature of sexual selection using formal selection analysis to statistically compare multivariate fitness surfaces. In T. commodus, sexual conflict occurs over the attachment time of an external spermatophore. By experimentally manipulating the ability of males and females to influence spermatophore attachment, we found that sexual conflict significantly influences the opportunity, form, and intensity of sexual selection on male courtship call and body size. When males were able to harass females, the opportunity for selection was smaller, the form of selection changed, and sexual selection was weaker. We discuss the broader evolutionary implications of these findings, including the contributions of sexual conflict to fluctuating sexual selection and the maintenance of additive genetic variation.     

Sex and context: hormones and primate sexual motivation

Gonadal hormones regulate the ability to copulate in most mammalian species, but not in primates because copulatory ability has been emancipated from hormonal control. Instead, gonadal hormones primarily influence sexual motivation. This separation of mating ability from hormonally modulated mating interest allows social experience and context to powerfully influence the expression of sexual behavior in nonhuman primates, both developmentally and in adulthood. For example, male rhesus monkeys mount males and females equally as juveniles, but mount females almost exclusively as adults. Having ejaculated with a female better predicted this transition to female mounting partners than did increased pubertal testosterone (T). It is proposed that increased pubertal T stimulates male sexual motivation, increasing the male's probability of sexual experience with females, ultimately producing a sexual preference for females. Eliminating T in adulthood reduces male sexual motivation in both humans and rhesus monkeys, but does not eliminate the capacity to engage in sex. In male rhesus monkeys the effects of reduced androgens on sexual behavior vary with social status and sexual experience. Human sexual behavior also varies with hormonal state, social context, and cultural conventions. Ovarian hormones influence female sexual desire, but the specific sexual behaviors engaged in are affected by perceived pregnancy risk, suggesting that cognition plays an important role in human sexual behavior. How the physical capacity to mate became emancipated from hormonal regulation in primates is not understood. This emancipation, however, increases the importance of motivational systems and results in primate sexual behavior being strongly influenced by social context.     

Context-dependent development of sexual ornamentation: implications for a trade-off between current and future breeding efforts

Allocation of resources into the development of sexual displays is determined by a trade-off between the competing demands of current reproduction and self-maintenance. When reproduction overlaps with acquisition of sexual ornamentation, such as in birds with a yearly post-breeding moult, such a trade-off can be expressed in elaboration of sexual traits used in subsequent matings. In turn, selection for elaboration of sexual ornaments should favour resolution of this trade-off through a modification of the ornaments' development, resulting in variable and life history-dependent development of sexual displays. Here we examined a novel hypothesis that the trade-off between current reproduction and development of sexual ornamentation in the house finch (Carpodacus mexicanus) can be mediated by the shared effects of prolactin - a pituitary hormone that regulates both parental care and moult in this species. We compared developmental variation in sexual ornamentation between breeding, nonbreeding, and juvenile males and examined the relative contribution of residual levels of prolactin and individual condition during moult to the acquisition of sexual ornamentation. Males that invested heavily in parental care entered post-breeding moult in lower condition and later in the season, but their higher plasma prolactin was associated with shorter and more intense moult ultimately resulting in equal or greater elaboration of sexual ornamentation compared with nonparental males. Elaboration of sexual ornamentation of nonparental males that entered moult in greater condition, but with lower prolactin, was produced by longer and earlier moult and by lesser overlap in moult between sexual ornaments. Ornamentation of juvenile males that acquire sexual ornamentation for the first time was closely associated with physiological condition during moult. We discuss the implications of such context-dependent ontogenies of sexual ornamentation and resulting differences in condition-dependence of sexual traits across life history stages on the evolution of female preference for elaborated sexual displays.     

EVOLUTION OF DIVERGENT FEMALE MATING PREFERENCE IN RESPONSE TO EXPERIMENTAL SEXUAL SELECTION

Sexual selection is predicted to drive the coevolution of mating signals and preferences (mating traits) within populations, and could play a role in speciation if sexual isolation arises due to mating trait divergence between populations. However, few studies have demonstrated that differences in mating traits between populations result from sexual selection alone. Experimental evolution is a promising approach to directly examine the action of sexual selection on mating trait divergence among populations. We manipulated the opportunity for sexual selection (low vs. high) in populations of Drosophila pseudoobscura. Previous studies on these experimental populations have shown that sexual selection manipulation resulted in the divergence between sexual selection treatments of several courtship song parameters, including interpulse interval (IPI) which markedly influences male mating success. Here, we measure female preference for IPI using a playback design to test for preference divergence between the sexual selection treatments after 130 generations of experimental sexual selection. The results suggest that female preference has coevolved with male signal, in opposite directions between the sexual selection treatments, providing direct evidence of the ability of sexual selection to drive the divergent coevolution of mating traits between populations. We discuss the implications in the context sexual selection and speciation.