How generalized CNS arousal strengthens sexual arousal (and vice versa)

Heightened states of generalized CNS arousal are proposed here to facilitate sexual arousal in both males and females. Genetic, pharmacologic and biophysical mechanisms by which this happens are reviewed. Moreover, stimulation of the genital epithelia, as triggers of sex behavior, is hypothesized to lead to a greater generalized arousal in a manner that intensifies sexual motivation. Finally, launched from histochemical studies intended to characterize cells in the genital epithelium, a surprising idea is proposed that links density of innervation with the efficiency of wound healing and with the capacity of that epithelium to stimulate generalized CNS arousal. Thus, bidirectional arousal-related mechanisms that foster sexual behaviors are envisioned as follows: from specific to generalized (as with genital stimulation) and from generalized to specific.     

Stimulus and hormonal determinants of flehmen behavior in cats

Felids are the main group of animals, other than ungulates, that display flehmen behavior during sociosexual interactions. In ungulates the behavior is evoked most readily by olfactory investigation of urine and vaginal secretions, and is believed to be involved in the transport of fluid-borne chemical stimuli, such as sex pheromones, from the oral cavity to the vomeronasal organ. In this study of cats flehmen was virtually always preceded by nasooral contact with the stimulus material, supporting the notion that flehmen in this species is also involved in the transport of fluid-borne stimuli. As in ungulates, flehmen in cats during heterosexual encounters was found to be displayed by males only. However, the sexual dimorphism was situation specific. In exploring a urine-marked room without another cat present, females also performed flehmen, albeit less frequently than males, and when urine was applied to the nasooral surface, flehmen was evoked equally reliably in females and males. Administration of testosterone propionate to spayed female cats paired with estrogen-treated females markedly increased their tendency to genitally inspect the female partner and subsequently perform flehmen. Thus the sexually dimorphic attributes of flehmen behavior are not only stimulus dependent, but also influenced by concurrent hormone stimulation.     

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.     

Grooming as a secondary behavior in the shrimp Macrobrachium rosenbergii (Crustacea,Decapoda, Caridea)

The giant freshwater prawn, Macrobrachium rosenbergii, is a large shrimp extensively used in aquaculture whose grooming behaviors were analyzed in this study. Macrobrachium rosenbergii exhibits three unique male morphotypes that differ in their behavior, morphology and physiology: small-clawed males (SM), orange-clawed males (OC) and blue-clawed males (BC). The largest and most dominant males, BC males, are predicted to have significantly different grooming behaviors compared to females and the other two male morphotypes. These BC males may be too large and bulky to efficiently groom and may dedicate more time to mating and agonistic interactions than grooming behaviors. Observations were conducted to look at the prevalence of grooming behaviors in the absence and presence of conspecifics and to determine if any differences in grooming behavior exist among the sexes and male morphotypes. Significant differences in the grooming behaviors of all individuals (females and male morphotypes) were found. BC males tended to have the highest grooming time budget (percent of time spent grooming) while SM males had a relatively low grooming time budget. The grooming behaviors of the male morphotypes differed, indicating while these males play distinct, separate roles in the social hierarchy, they also have different grooming priorities. The conditions in which Macrobrachium rosenbergii are cultured may result in increased body fouling, which may vary, depending on the grooming efficiencies and priorities of these male morphotypes. Overall, grooming behaviors were found to be a secondary behavior which only occurred when primary behaviors such as mating, feeding or fighting were not present.     

Disgust versus Lust: Exploring the Interactions of Disgust and Fear with Sexual Arousal in Women

Sexual arousal is a motivational state that moves humans toward situations that inherently pose a risk of disease transmission. Disgust is an emotion that adaptively moves humans away from such situations. Incongruent is the fact that sexual activity is elementary to human fitness yet involves strong disgust elicitors. Using an experimental paradigm, we investigated how these two states interact. Women (final N=76) were assigned to one of four conditions: rate disgust stimuli then watch a pornographic clip; watch a pornographic clip then rate disgust stimuli; rate fear stimuli then watch a pornographic clip; or watch a pornographic clip then rate fear stimuli. Women’s genital sexual arousal was measured with vaginal photoplethysmography and their disgust and fear reactions were measured via self-report. We did not find that baseline disgust propensity predicted sexual arousal in women who were exposed to neutral stimuli before erotic content. In the Erotic-before-Disgust condition we did not find that sexual arousal straightforwardly predicted decreased image disgust ratings. However, we did find some evidence that sexual arousal increased self-reported disgust in women with high trait disgust and sexual arousal decreased self-reported disgust in women with low trait disgust. Women who were exposed to disgusting images before erotic content showed significantly less sexual arousal than women in the control condition or women exposed to fear-inducing images before erotic content. In the Disgust-before-Erotic condition the degree of self-reported disgust was negatively correlated with genital sexual arousal. Hence, in the conflict between the ultimate goals of reproduction and disease avoidance, cues of the presence of pathogens significantly reduce the motivation to engage in mating behaviors that, by their nature, entail a risk of pathogen transmission.     

On the intricate relationship between sexual motivation and arousal

Sexual motivation and sexual arousal are widely used concepts. While there seem to be considerable agreement as to the meaning of sexual motivation, there is certain confusion about the exact meaning of sexual arousal. Some use it as a synonym to sexual motivation and others make it equivalent to erection or vaginal lubrication. An unresolved question is the relationship between sexual arousal and general arousal as well as that between arousal and motivation. I present arguments for the view that arousal refers to the general state of alertness of the organism. Consequently, there is no such thing as a specific sexual arousal. I suggest that this term should be abandoned, or if that is not feasible, to make it a synonym to enhanced genital blood flow. The notion of a subjective sexual arousal, some kind of vaguely described mental state, seems to lack all explanatory value. I then show that general arousal is an important determinant of sexual motivation, and that the execution of copulatory acts leads to increased general arousal. This increase leads to enhanced sexual motivation, making the activation of sexual reflexes requiring high levels of motivation possible. Examples of such reflexes may be ejaculation in males of many species, and perhaps the psychic state of orgasm in women.     

Females prefer to associate with males with longer intromittent organs in mosquitofish

Sexual selection is a major force behind the rapid evolution of male genital morphology among species. Most within-species studies have focused on sexual selection on male genital traits owing to events during or after copulation that increase a male''s share of paternity. Very little attention has been given to whether genitalia are visual signals that cause males to vary in their attractiveness to females and are therefore under pre-copulatory sexual selection. Here we show that, on average, female eastern mosquitofish Gambusia holbrooki spent more time in association with males who received only a slight reduction in the length of the intromittent organ (‘gonopodium’) than males that received a greater reduction. This preference was, however, only expressed when females chose between two large males; for small males, there was no effect of genital size on female association time.     

Complex patterns of grooming and sexual activity in Barbary macaques (Macaca sylvanus)

Grooming in primates is often considered a “currency” that can be exchanged for other “services” or “commodities” such as reciprocal grooming, coalitionary support, infant handling, tolerance around food sources, active food sharing, or mating opportunities. Previous studies on primate grooming‐for‐sex exchange viewed the males as the demanding class, with the females as suppliers of mating opportunities. In this study, we examine the broader context of grooming‐for‐mating exchange in Barbary macaques in Gibraltar. Our data show that Barbary macaque males groom females with whom they are mating more frequently and for longer periods than other females, and the relationship between grooming and mating remains significant in both sexual and nonsexual contexts. In addition, females groomed males with whom they were mating more frequently and for longer periods than other males. In both sexes, grooming was observed to be far more frequent and to occur for longer durations in sexual compared to nonsexual contexts. We did not find any difference in grooming behavior between presexual and postsexual contexts. Our data suggest that there is no simple model to describe Barbary macaque grooming patterns in sexual contexts. Although our results are partly consistent with male use of grooming as payment for mating, broadly assessed grooming‐mating patterns cannot be solely explained by a male‐driven grooming‐for‐mating exchange.     

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.     

Orexin mediates initiation of sexual behavior in sexually naive male rats, but is not critical for sexual performance

The hypothalamic neuropeptide orexin mediates arousal, sleep, and naturally rewarding behaviors, including food intake. Male sexual behavior is altered by orexin receptor-1 agonists or antagonists, suggesting a role for orexin-A in this naturally rewarding behavior. However, the specific role of endogenous orexin-A or B in different elements of male sexual behavior is currently unclear. Therefore, the current studies utilized markers for neural activation and orexin cell-specific lesions to test the hypothesis that orexin is critical for sexual motivation and performance in male rats. First, cFos expression in orexin neurons was demonstrated following presentation of a receptive or non-receptive female without further activation by different elements of mating. Next, the functional role of orexin was tested utilizing orexin-B conjugated saporin, resulting in orexin cell body lesions in the hypothalamus. Lesions were conducted in sexually naive males and subsequent sexual behavior was recorded during four mating trials. Lesion males showed shortened latencies to mount and intromit during the first, but not subsequent mating trials, suggesting lesions facilitated initiation of sexual behavior in sexually naive, but not experienced males. Likewise, lesions did not affect sexual motivation in experienced males, determined by runway tests. Finally, elevated plus maze tests demonstrated reduced anxiety-like behaviors in lesioned males, supporting a role for orexin in anxiety associated with initial exposure to the female in naive animals. Overall, these findings show that orexin is not critical for male sexual performance or motivation, but may play a role in arousal and anxiety related to sexual behavior in naive animals.     

A contextual definition of male sexual arousal

Sexual arousal is a construct without a widely shared definition. Historically, sexual arousal has usually referred to a central physiological state, but there has been much less agreement on its relation to motivation, emotion, and - for males - penile erection and ejaculation. Many behavioral and physiological measures have been used as operational definitions of sexual arousal, but the relation of the measure to arousal is often assumed rather than tested. For men, penile erection in the presence of erotic stimuli has been considered the most reliable and valid indicator of sexual arousal. The adoption of analogous criteria is recommended for research on other male mammals in order to establish a minimal basis for inferring that they are sexually aroused. That is, sexual arousal should be inferred only when penile erection is observed in a sexual context. A sexual context is provisionally defined as an environment that tends in most reproductively active males of the species to provoke further sexual stimulation, e.g., copulation or self-stimulation to ejaculation. Erection occurring outside of a sexual context, as during REM sleep or from injection of drugs, is not grounds for inferring arousal. Conversely, males engaging in behavior directed toward estrous females may be sexually motivated, but in the absence of erection, the males should not be assumed to be sexually aroused. Implications of other erection-context interactions are also considered. Adoption of these more conservative criteria for inferring sexual arousal may promote greater precision in identifying the physiological systems mediating this hypothetical construct.     

Homosexual Fellatio: Erect Penis Licking between Male Bonin Flying Foxes Pteropus pselaphon

A recent focus of interest has been on the functional significance of genital licking (fellatio and cunnilingus) in relation to sexual selection in Pteropodid bats. In the present paper, a form of fellatio in wild Bonin flying foxes, Pteropus pselaphon, performed between adult males has been reported. During the mating season, adult flying foxes roost in same-sex groups, forming ball-shaped clusters which provide warmth. The female clusters may also contain a few males. Unassociated with allogrooming, same-sex genital licking occurred among males in the all male clusters. As such, male-male fellatio can be considered as homosexual behavior, two functional explanations could account for this behavior; the social bonding and the social tension regulation hypotheses suggested in a previous review. Given that neither the simpler alternative that in all male groups such fellatio may represent misdirected sexual behavior, nor the two previously proposed functional hypotheses were supported by the data, I propose another functional hypothesis. Homosexual fellatio in this species could help males solve inconsistent situations in the roost when there are conflicts between cooperative behavior for social thermoregulation and competition for mating.     

Intersexual cooperation during male clasping of external female genitalia in the spider Physocyclus dugesi (Araneae, Pholcidae)

Sexual conflict may influence the shape and evolution of body structures that males use to grasp females during mating. Not only sexual coercion but also intersexual cooperation may be involved during clasping behavior. Among pholcid spiders, secondary sexual modifications of the male chelicerae, such as apophyses with spines or tooth-like processes, function to grasp the female by specific parts of her external genitalia such as grooves or apophyses of the epigynum. We analyzed how the female and the male respond when their structures for clasping are experimentally modified in the pholcid Physocyclus dugesi. We used three treatment groups for virgin females that differed in the manipulation of the epigynum apophyses (uncovered, partially covered, and fully covered by a plaster) and two groups of males (uncovered and fully covered cheliceral apophyses). We found that females are mainly cooperative to courting males not only when the female genital apophyses were experimentally covered but also when the male cheliceral apophyses were covered. The current data also indicate behavioral flexibility in males during courtship, especially when they had difficulty in genital intromission. Our experimental results, together with previous observational studies, support a modulated-cooperative scenario between the sexes for cheliceral clasping and genital intromission in pholcid spiders.     

Female genital mutilation and monandry in an orb-web spider

Monandry, in which a female has only one mating partner during the reproductive period, is established when a female spontaneously refrains from re-mating, or when a partner male interferes with the attempts of a female to mate again. In the latter case, however, females often have countermeasures against males, which may explain why polyandry is ubiquitous. Here, I demonstrate that the genital appendage, or scape, of the female orb-web spider (Cyclosa argenteoalba) is injured after her first mating, possibly by her first male partner. This female genital mutilation (FGM) permanently precludes copulation, and females appear to have no countermeasures. FGM is considered to confer a strong advantage to males in sexual conflicts over the number of female matings, and it may widely occur in spiders.     

Mate Choice in Experimentally Parasitized Rock Doves: Lousy Males Lose

A recent model by Hamilton and Zuk (1982) suggests that exaggeratedsecondary sexual traits facilitate mate choice for genetic resistanceto parasites. The model predicts that individuals discriminateagainst parasitized mates by scrutinizing traits indicativeof parasite load. In the case of birds and their feather-feedinglice, for example, individuals might avoid parasitized matesbydetecting reduced plumage brightness, reduced courtship display,or increased grooming. I conducted a series of mate choice trialsin which female Rock Doves (Columba livia) were allowed to choosebetween "clean" males without lice and "lousy" males with experimentallyincreased loads. Clean males displayed significantly more oftenthan lousy males and females demonstrated a significant preferencefor clean males. Lousy males were subject to plumage damage;however, none of the damage was externally visible, and thetime spent grooming by clean and lousy males did not differsignificantly. Female louse loads, which were also manipulated,were not significantly related to female mating preferences.These results are consistent with the Hamilton-Zuk model. Theyare also consistent with a model of sexual selection for theavoidance of parasite transmission, which is discussed. Thegeneral relevance of lice and other ectoparasites to modelsof parasite-mediated sexual selection is reviewed.     

Discovery that a melanocortin regulates sexual functions in male and female humans

Melanocortins (MCs) are multifunctional peptide hormones that regulate a diversity of physiological functions. MCs have been implicated in sexual function in animals. We document here that a MC analog, Melanotan II (MTII), can enhance sexual function in human males (erectile activity) and females (increased levels of sexual desire and genital arousal). Unlike other sexual-enhancement drugs, MTII works at the level of the brain, thus eliciting a rather natural sexual response with minimal or no undesirable side effects. The actions of the peptide were discovered accidentally while studying the effects of the peptide and related analogs on human skin pigmentation (tanning).     

The Adaptive Function of Masturbation in a Promiscuous African Ground Squirrel

Background

Studies of animal mating systems increasingly emphasize female multiple mating and cryptic sexual selection, particularly sperm competition. Males under intense sperm competition may manipulate sperm quantity and quality through masturbation, which could waste sperm and decrease fertility. I examined the factors influencing masturbation by male Cape ground squirrels (Xerus inauris) in light of a number of functional hypotheses.

Methodology

Observational data on a marked population of squirrels were collected in east-central Namibia using scan and all-occurrences sampling.

Findings

Masturbation was far more frequent on days of female oestrus and mostly occurred after copulation. Masturbation rates were higher in dominant males, which copulate more, than in subordinates and increased with number of mates a female accepts.

Conclusions

These results suggest that masturbation in this species was not a response to sperm competition nor a sexual outlet by subordinates that did not copulate. Instead masturbation could function as a form of genital grooming. Female Cape ground squirrels mate with up to 10 males in a 3-hr oestrus, and by masturbating after copulation males could reduce the chance of infection. Sexually transmitted infections (STIs) can profoundly affect fertility, and their consequences for mating strategies need to be examined more fully.     

Stimulation of reproductive activity in anovulatory Alpine goats exposed to bucks treated only with artificially long days

Two experiments were conducted in a subtropical latitude to determine the response of Alpine male goats to a treatment with artificially long days (experiment 1), and the response of anovulatory lactating Alpine does exposed to males treated only with artificially long days (experiment 2). In experiment 1, one group of males was kept under natural photoperiod (n = 4) while another was exposed to 2.5 months of artificially long days (16 h of light/day) from 1 December (n = 4). Plasma testosterone concentrations were determined weekly. Intensity of odor of males was determined every 2 weeks. Sexual behavior of bucks was observed during 3 days about 90 days after the end of the long day treatment. A treatment-by-time interaction was detected for testosterone secretion (P < 0.001). In control males, low plasma concentrations of testosterone were observed from March to June. In contrast, in long-day treated males, high levels of testosterone were observed from March to June (P < 0.05). A treatment-by-time interaction was detected for the intensity of male odor (P < 0.01). The male odor was stronger in long-day treated bucks than in untreated ones from March to June (P < 0.05). The number of ano-genital sniffing, nudging and flehmen was greater in long-day treated males than in untreated ones when exposed to anestrous does (P < 0.05). In experiment 2, one group of males was left under natural photoperiod variations (n = 5) and the other (n = 5) was submitted to the same photoperiodic treatment described in experiment 1. On 3 May, three untreated and three long-day treated males were put in contact with anestrous Alpine does left under natural photoperiod. Fertility was higher in does exposed to light-treated males (36/45, 80%) than those in contact with untreated ones (3/45, 7%; P < 0.05). Prolificacy was similar (P > 0.05) in does exposed to treated (1.8 ± 0.1) and untreated males (1.7 ± 0.3). These results indicate that the sexual activity of Alpine male goats raised in subtropical latitudes can be induced using only artificially long days and that such males are effective in stimulating reproductive activity in anovulatory females in late spring.     

Identification of putative pheromones in bovine (Bos taurus) faeces in relation to estrus detection

Ten different volatile compounds were detected in bovine faeces using three chromatograms. The chemical profiles of estrus faeces were distinguished significantly from other phases by the presence of three specific substances, viz. acetic acid and propionic acid and 1-iodo undecane. The estrus specific synthetic compounds were rubbed onto the genital region of nonestrus animals (dummy cows), and the bulls were allowed to sniff the genital region and observed sexual behaviours. The statistical significance was higher (P<0.001) in bulls exhibiting repeated flehmen and mounting behaviours towards the mixture of acetic acid, propionic acid and 1-iodo undecane. The bioassay revealed that the fatty acids viz., acetic acid; propionic acid and 1-iodo undecane produced in faeces during estrus appear to be estrus specific. The results suggest that these compounds may be used as estrus indicator in cow, probably involved in bovine biocommunication.     

Female Choice or Male Sex Drive? The Advantages of Male Body Size during Mating in Drosophila Melanogaster

The mating success of larger male Drosophila melanogaster in the laboratory and the wild has been traditionally been explained by female choice, even though the reasons are generally hard to reconcile. Female choice can explain this success by virtue of females taking less time to mate with preferred males, but so can the more aggressive or persistent courtships efforts of large males. Since mating is a negotiation between the two sexes, the behaviors of both are likely to interact and influence mating outcomes. Using a series of assays, we explored these negotiations by testing for the relative influence of male behaviors and its effect on influencing female courtship arousal threshold, which is the time taken for females to accept copulation. Our results show that large males indeed have higher copulation success compared to smaller males. Competition between two males or an increasing number of males had no influence on female sexual arousal threshold;—females therefore may have a relatively fixed ‘arousal threshold’ that must be reached before they are ready to mate, and larger males appear to be able to manipulate this threshold sooner. On the other hand, the females’ physiological and behavioral state drastically influences mating; once females have crossed the courtship arousal threshold they take less time to mate and mate indiscriminately with large and small males. Mating quicker with larger males may be misconstrued to be due to female choice; our results suggest that the mating advantage of larger males may be more a result of heightened male activity and relatively less of female choice. Body size per se may not be a trait under selection by female choice, but size likely amplifies male activity and signal outputs in courtship, allowing them to influence female arousal threshold faster.