Sexual arousal in men: a review and conceptual analysis

Sexual arousal is an emotional/motivational state that can be triggered by internal and external stimuli and that can be inferred from central (including verbal), peripheral (including genital), and behavioral (including action tendencies and motor preparation) responses. This article, while focusing on sexual arousal in men, provides a conceptual analysis of this construct, reviews models of sexual arousal, and discusses the usefulness of perspectives derived from motivation and emotion research in improving our understanding of its determinants and behavioral correlates. In this, it considers the role of genital feedback in men's subjective sexual arousal and the connections between sexual arousal and sexual desire. Future research and definitions may increasingly focus on its central integrative functions (as opposed to its input and output characteristics). Yet, the study of sexual arousal can be expected to continue to benefit from the measurement of its genital, verbal, and behavioral components. Instances of discordance between response components suggest that they are, at least in part, under the control of different mechanisms, and it is proposed that a better understanding of sexual arousal will prove contingent on a better understanding of such mechanisms and the conditions under which they converge and diverge.     

Sexual selection and mate choice in evolutionary psychology

The importance of mate choice and sexual selection has been emphasized by the majority of evolutionary psychologists. This paper assesses three cases of work on mate choice and sexual selection in evolutionary psychology: David Buss on cross-cultural human mate preferences, Randy Thornhill and Steve Gangestad on the link between mate preferences and fluctuating asymmetry, and Geoffrey Miller on the role of Fisher’s runaway process in human evolution. A mixture of conceptual and empirical problems in each case highlights the general weakness of work in evolutionary psychology on these issues.

Sperm competition can drive a male-biased mutation rate

A pattern of male-biased mutation has been found in a wide range of species. The standard explanation for this bias is that there are greater numbers of mitotic cell divisions in the history of the average sperm, compared to the average egg, and that mutations typically result from errors made during replication. However, this fails to provide an ultimate evolutionary explanation for why the male germline would tolerate more mutations that are typically deleterious. One possibility is that if there is a tradeoff between producing large numbers of sperm and expending energetic resources in maintaining a lower mutation rate, sperm competition would select for males that produce larger numbers of sperm despite a higher resulting mutation rate. Here I describe a model that jointly considers the fitness consequences of deleterious mutation and mating success in the face of sperm competition. I show that a moderate level of sperm competition can account for the observation that the male germline tolerates a higher mutation rate than the female germline.     

Evolutionary morphology of the male reproductive system,spermatozoa and seminal fluid of spiders (Araneae,Arachnida) – Current knowledge and future directions

The male reproductive system and spermatozoa of spiders are known for their high structural diversity. Spider spermatozoa are flagellate and males transfer them to females in a coiled and encapsulated state using their modified pedipalps. Here, we provide a detailed overview of the present state of knowledge of the primary male reproductive system, sperm morphology and the structural diversity of seminal fluids with a focus on functional and evolutionary implications. Secondly, we conceptualized characters for the male genital system, spermiogenesis and spermatozoa for the first time based on published and new data. In total, we scored 40 characters for 129 species from 56 families representing all main spider clades. We obtained synapomorphies for several taxa including Opisthothelae, Araneomorphae, Dysderoidea, Scytodoidea, Telemidae, Linyphioidea, Mimetidae, Synotaxidae and the Divided Cribellum Clade. Furthermore, we recovered synspermia as a synapomorphy for ecribellate Haplogynae and thus propose Synspermiata as new name for this clade. We hope that these data will not only contribute to future phylogenetic studies but will also stimulate much needed evolutionary studies of reproductive systems in spiders.     

Mating games: cultural evolution and sexual selection

In this paper, we argue that mating games, a concept that denotes cultural practices characterized by a competitive element and an ornamental character, are essential drivers behind the emergence and maintenance of human cultural practices. In order to substantiate this claim, we sketch out the essential role of the game’s players and audience, as well as the ways in which games can mature and turn into relatively stable cultural practices. After outlining the life phase of mating games – their emergence, rise, maturation, and possible eventual decline – we go on to argue that participation in these games (in each phase) does make sense from an adaptationist point of view. The strong version of our theory which proposes that all cultural practices are, or once were, mating games, allows us to derive a set of testable predictions for the fields of archaeology, economics, and psychology.     

Live for the moment—Adaptations in the male genital system of a sexually cannibalistic spider (Theridiidae, Araneae)

Monogyny in spiders culminates in extreme traits, like dramatic male self-sacrifice and emasculation of the male by the female during copulation. Here we show that monogynous males can be highly adapted for this fatal sexual behaviour. Dwarf males of the one-palped theridiid spider Tidarren argo, which are cannibalised immediately after the insertion of their single copulatory organ, stop spermiogenesis when reaching adulthood. Their testes atrophy, which might economise the energy expenditures of these males. We also found that the amount of seminal fluid produced is stored in an enlarged seminal vesicle until the single sperm induction takes place. The volume of the seminal vesicle is similar to the sperm droplet taken up into the copulatory organ (palpal organ). Sperm uptake takes much longer than in related species most likely due to the large amount of seminal fluid. As shown by histological observations males are able to fill one of the paired female sperm storage organs during copulation thereby presumably impeding subsequent charging by rival males.     

Adaptive Evolution of Gamete-Recognition Proteins in Birds

Gamete-recognition proteins have been shown to evolve by positive selection in diverse organism groups, such as marine invertebrates and mammals, although underlying evolutionary mechanisms driving this rapid divergence are poorly understood. However, several hypotheses have been put forward to explain the observed pattern, including different forms of sexual conflict and sperm competition. Because female gametes require more energy to produce than male gametes, female organisms suffer more when fertilisation goes wrong. One process that results in a failed mammalian fertilisation is polyspermy, when >1 sperm fertilises the egg. However in birds, there is no such sexual conflict because multiple sperm typically bind and fuse with the egg. If sexual conflict driven by polyspermy avoidance is important for the evolution of gamete-recognition proteins in vertebrates, we expect to find positive selection in the genes to be less pronounced in birds. We therefore sequenced six genes (ZP1, ZP2, ZP4, ZPAX, CD9, and Acrosin) encoding gamete-recognition proteins in several bird species to test for positive selection. For comparison, we also analysed ortologous sequences in a set of mammalian species. We found no major differences in the occurrence of adaptive evolution and the strength of selection between bird and mammal orthologs. From this we conclude that polyspermy avoidance does not act as the main underlying evolutionary force shaping the rate of evolution in these genes. We discuss other possible processes that could explain positive selection of gamete-recognition proteins in birds and mammals, such as hybridisation avoidance, cryptic female choice, and postcopulatory sperm competition.     

Sexual selection affects the sizes of the mammalian prostate gland and seminal vesicles

The accessory reproductive glands of male mammals contribute the bulk of the secretions in which spermatozoa are transported to the female tract during copulation. Despite their morphological diversity,and the chemical complexity of their products,little is known about the possible effects of sexual selection upon these glands in mammals. Here we consider the seminal vesicles and prostate glands in a sample of 89 species and 60 genera representing 8 Orders of mammals. The sizes of the accessory glands are analysed in relation to body weight and testes weight. Both the seminal vesicles size and prostate size (corrected for body weight) correlate positively with relative testes size in this sample; this finding remains highly significant after application of procedures to correct for possible phylogenetic biases in the data set. The accessory reproductive glands are also significantly larger in those mammals which have large relative testes sizes,and in which the likelihood of sperm competition is greatest. These results support the hypothesis that sexual selection has played an important role in the evolution of the mammalian prostate gland and seminal vesicles.     

Copulation patterns in the golden orb-web spider Nephila madagascariensis

A consequence of multiple mating by females can be that the sperm of two or more males directly compete for the fertilisation of ova inside the female reproductive tract. Selection through sperm-competition favours males that protect their sperm against that of rivals and strategically allocate their sperm, e.g., according to the mating status of the female and the morphology of the spermatheca. In the majority of spiders, we encounter the otherwise unusual situation that females possess two independent insemination ducts, both ending in their own sperm storage organ, the spermatheca. Males have paired mating organs, but generally can only fill one spermatheca at a time. We investigated whether males of the African golden orb-web spider Nephila madagascariensis can prevent rival males from mating into the same spermatheca and whether the mating status of the female and/or the spermatheca causes differences in male mating behaviour. There was no significant difference in the duration of copulations into unused spermathecae of virgin and mated females. We found that copulations into previously inseminated spermathecae were generally possible, but shorter than copulations into the unused side of mated females or with virgins. Thus, male N. madagascariensis may have an advantage when they mate with virgins, but cannot prevent future males from mating. However, in rare instances, parts of the male genitals can completely obstruct a female genital opening.     

Degree of protandry reflects level of extrapair paternity in migratory songbirds

Males of most migratory organisms, including many birds, precede female conspecifics on their journey to the breeding areas. Several hypotheses have been proposed to explain the evolution of protandrous migration, yet they have rarely been tested at the interspecific level. Here, we provide correlational support for the “mate opportunity” hypothesis, which assumes that selection favours protandry in polygynous species where males gain significant fitness benefits from arriving earlier than females. Drawing on phenological data collected at two northern European stopover sites, we show that the time-lag in spring passage between males and females of five Palearctic migratory songbird species is positively associated with levels of extrapair paternity available from the literature. This suggests that males arrive relatively more in advance of females in species with high sperm competition where sexual selection through female choice is intense. Thus, protandry may arise from selection on the relative arrival timing of males and females rather than from selection within one of the sexes.     

Paternity assessment in free-ranging wild boar (Sus scrofa) – Are littermates full-sibs?

Multiple paternity within litters occurs in various groups of mammals exhibiting different mating systems. Using seven genetic markers (i.e., microsatellites), we investigated the paternity of littermates in free-ranging wild boar (Sus scrofa) in a Mediterranean habitat. Using the software CERVUS 2.0 we estimated the probability of detecting multiple paternity across all loci (D), the probability of paternity (W) and a statistic Δ that allows the assignment of paternity to the most likely male with strict and relaxed levels of confidence. Multiple paternity was inferred for one of the nine analysed litters at the 80% confidence level. This suggests that a single male may control the access to receptive adult females and it shows that multiple paternity is not very common in the studied free-ranging wild boar population. Despite the possible occurrence of sperm competition and/or female cryptic choice, mate guarding seems to play a significant role in sexual selection. To better understand the wild boar's mating strategies further studies analysing the reproductive success of both sexes and under different environmental conditions should be conducted.     

性选择、配偶外父亲身份确认程度、遗传变异性和保护

岛屿动物中的性选择强度不高,其原因可能是由于岛屿种群的遗传变异性水平较低。本文作者检验了鸟类岛屿种群是否具有较低的遗传变异性、性选择强度大的种群是否具有较高的突变输入率（rate of mutational input),在鸟类岛屿种群中是否具有较低的性选择强度（可以根据配偶外父亲身份的频率来估计）。小卫星共有谱带系数（minisatellite band sharing coefficient)可确定无亲源关系个体之间的遗传变异性,对与遗传变异性有关的雄性个体的父亲（paternity)进行了成对比较以检验如下假说：在具有较多遗传变异的种群中,雌性个体更经常地进行配偶外交配。在小卫星谱带系数较低的鸟类种群中,配偶外父亲的频次较高。对岛屿和大陆鸟类进行的第二个比较分析表明：岛屿种群中的配偶外父亲频次较低,遗传变异性也较低,其部分原因在于突变输入（mutational input)减少。上述发现表明：（1）父亲确认程度（parternity)随遗传变异性的数量而增加;（2）在遗传变异性较大的种群中,突变率较高,性选择的程度更激烈;（3）岛屿种群中性选择的强度一般比大陆种群弱。这对于理解遗传变异性的空间变异、理解岛屿种群和其它隔离种群的保护问题有重要启示。     

Testosterone, testes size, and mating success in birds: a comparative study

Reproductive behaviors of vertebrates are often underpinned by temporal patterns of hormone secretion. We investigated interspecific patterns of circulating testosterone in male birds to test the hypothesis that testosterone plays a crucial role in sexual selection as determined by degree of polygyny and extra-pair paternity. We predicted that the evolution of increased levels of polygyny and extra-pair paternity would have resulted in the evolution of increased levels of testosterone to allow males more efficiently to compete for mates. This hypothesis was tested in comparative analyses of 116 species of birds using Generalized Least Squares Models. We assessed the importance of latitudinal distribution, because this can confound the relationship between testosterone and mating success. There were weak positive phylogenetic correlations between measures of testosterone and estimates of mating success at the social level, but this association appeared to be confounded by latitudinal distribution, a significant correlate of testosterone titers. However, we found a significantly positive relationship between peak and residual peak testosterone (which is the peak testosterone level that is controlled for the baseline level) and extra-pair paternity independent of latitude. These results suggest that selection pressures arising from social and sexual mating differently affected testosterone levels with the former being mediated by factors associated with latitudinal distribution. An analysis of residual testes size revealed a positive association between peak and residual testosterone and testes size relative to body size. In a path analysis, we show that relative testis size primarily evolved in association with intense sperm competition and thus high sperm production, and these mechanisms had a secondary impact on blood testosterone levels at a phylogenetic scale. Our results suggest that sperm competition has played an important role in the evolution of reproductive mechanisms in birds.     

Cryptic female choice during spermatophore transfer in Tribolium castaneum (Coleoptera: Tenebrionidae)

Sexual selection in both males and females promotes traits and behaviors that allow control over paternity when female mates with multiple males. Nonetheless, mechanisms of cryptic female choice have been consistently overlooked, due to traditional focus on sperm competition as well as difficulty in distinguishing male vs. female influence over processes occurring during and after mating. The first part of this study describes morphology and transformation of Tribolium castaneum spermatophores inferred from dissecting females immediately after normal or interrupted copulations. T. castaneum males are found to transfer spermatophores as an invaginated tube that everts inside the female bursa and which is filled with sperm during copulation. This sequence of events makes it feasible for females to control the sperm quantity transferred in each spermatophore. Through manipulation of the male phenotypic quality (by starvation) and manipulation of female control over sperm transfer (by killing a subset of females), the second part of this study examines whether females use control over transferred sperm quantity as a cryptic choice mechanism. Fed males transferred significantly more sperm per spermatophore than starved males but only when mating with live females. These results suggest an active differentiation by live females against starved males and provide an evidence for the proposed cryptic female choice mechanism.     

Contest competition and men's facial hair: beards may not provide advantages in combat

In contemporary human societies, where direct male-male competition is reduced compared to ancestral societies, sporting competitions remain an avenue for status acquisition via intra-sexual competition. Beards are the most visually salient and sexually dimorphic of men's secondary sexual traits and play a strong role in communicating masculinity, dominance and aggressiveness intra-sexually. Hypotheses have been advanced that beards provide advantages in intra-sexual combat, as protective organs and honest signals of fighting ability. Here we provide the first test of these hypotheses using data from professional mixed martial arts fighters competing in the Ultimate Fighting Championship. We explored whether secondary sexual traits (height, weight, beardedness), fighting stance (southpaw, orthodox), arm reach and past contest experiences impact on contest outcomes. If beards function as protective organs, bearded fighters should succumb to fewer knock-outs, and hence protection to injuries to the jaw, fewer abrasions and lacerations to the face and brain damage than clean-shaven fighters. Alternatively, if beards signal fighting ability then bearded fighters should win more fights. We found no evidence that beardedness was associated with fewer losses by knock-out or greater fighter ability. While fighters with longer reaches won more fights, neither stance nor past experience influenced fight outcomes. We suggest that beards represent dishonest signals of formidability that may serve to curtail the escalation of intra-sexual conflict through intimidation rather than providing advantages in direct combat.     

Size matters: genital allometry in an African mole-rat (Family: Bathyergidae)

Typically, sexually selected traits show positive allometry and high coefficients of variation (CV). To date, many studies on the allometry of genitalia have focused on insects. In addition, studies have largely ignored the potential for sexual selection on female genitalia, despite male and female structures presumably co-evolving. Insects tend to show negative allometry in both male and female genitalia, while in contrast, the few studies carried out in mammals (males only) show positive allometry. Reasons for these differences between the taxa still remain unclear. However, in mammals, three main mechanisms have been proposed for genital evolution, namely, sperm competition, female cryptic choice and sexual conflict. In the first such study that we are aware of, we examined intra-specific genital allometry in both males and females of a mammal, the subterranean solitary Cape dune mole-rat, Bathyergus suillus. We found positive allometry occurring in male genitalia, which is consistent with previous vertebrate studies. Similarly, we found that female genitalia also exhibited positive allometry further supporting the notion of co-evolution of male and female genitalia. Although it is difficult to distinguish between the forces or mechanisms determining this directional selection, we suggest that several reproductive advantages are incurred as a result of positive allometric relationship of the genitalia in B. suillus and such advantages are also likely in other subterranean mammals. Our study further highlights the differences in genital allometry across taxa.     

Conditioned sexual arousal in a nonhuman primate

Conditioning of sexual arousal has been demonstrated in several species from fish to humans but has not been demonstrated in nonhuman primates. Controversy exists over whether nonhuman primates produce pheromones that arouse sexual behavior. Although common marmosets copulate throughout the ovarian cycle and during pregnancy, males exhibit behavioral signs of arousal, demonstrate increased neural activation of anterior hypothalamus and medial preoptic area, and have an increase in serum testosterone after exposure to odors of novel ovulating females suggestive of a sexually arousing pheromone. Males also have increased androgens prior to their mate's ovulation. However, males presented with odors of ovulating females demonstrate activation of many other brain areas associated with motivation, memory, and decision making. In this study, we demonstrate that male marmosets can be conditioned to a novel, arbitrary odor (lemon) with observation of erections, and increased exploration of the location where they previously experienced a receptive female, and increased scratching in post-conditioning test without a female present. This conditioned response was demonstrated up to a week after the end of conditioning trials, a much longer lasting effect of conditioning than reported in studies of other species. These results further suggest that odors of ovulating females are not pheromones, strictly speaking and that marmoset males may learn specific characteristics of odors of females providing a possible basis for mate identification.