Female sexual arousal in amphibians

Rather than being a static, species specific trait, reproductive behavior in female amphibians is variable within an individual during the breeding season when females are capable of reproductive activity. Changes in receptivity coincide with changes in circulating estrogen. Estrogen is highest at the point when females are ready to choose a male and lay eggs. At this time female receptivity (her probability of responding to a male vocal signal) is highest and her selectivity among conspecific calls (measured by her probability of responding to a degraded or otherwise usually unattractive male signal) is lowest. These changes occur even though females retain the ability to discriminate different acoustic characteristics of various conspecific calls. After releasing her eggs, female amphibians quickly become less receptive and more choosy in terms of their responses to male sexual advertisement signals. Male vocal signals stimulate both behavior and estrogen changes in amphibian females making mating more probable. The changes in female reproductive behavior are the same as those generally accepted as indicative of a change in female sexual arousal leading to copulation. They are situationally triggered, gated by interactions with males, and decline with the consummation of sexual reproduction with a chosen male. The changes can be triggered by either internal physiological state or by the presence of stimuli presented by males, and the same stimuli change both behavior and physiological (endocrine) state in such a way as to make acceptance of a male more likely. Thus amphibian females demonstrate many of the same general characteristics of changing female sexual state that in mammals indicate sexual arousal.     

Fatherhood, pairbonding and testosterone in the Philippines

In species with a high level of paternal care, including humans, testosterone (T) is believed to help mediate the trade-off between parenting and mating effort. This hypothesis is supported by the observation of lower T in pairbonded men or fathers compared to single, non-fathers; however, prior work has highlighted population variation in the association between T and pairbonding or fatherhood status. Here we evaluate this hypothesis in a large (n = 890), representative birth cohort of young men (age range 20.5–22.5 years) living in Cebu City, the Philippines. Bioavailable T was measured in saliva collected prior to bed and immediately upon waking the following morning. Plasma T and luteinizing hormone (LH) were measured in morning plasma samples. In this sample, 20% of men were pairbonded, defined as living with a partner or married, 13% were fathers, and roughly half of fathers reported involvement in childcare. Pairbonded men had significantly lower T at both times of day. Unlike in other populations, this relationship was accounted for entirely by fatherhood status: among the large sub-sample of non-fathers, mean T was nearly identical among pairbonded and single men. There was a strong association between self-reported involvement in childcare and lower evening T, supporting the idea that the evening nadir in T is related to social interactions across the day. Similar relationships were found for total plasma T and LH, suggesting that these relationships are coordinated by centrally-mediated changes in LH secretion. The relatively modest T difference in relation to fatherhood at Cebu, in comparison to other studies, may reflect a lower level of paternal involvement in childcare activities in this population. Our findings using a large, well-characterized birth cohort support the hypothesized role of T as a mediator of mating and parenting effort in humans, while contributing evidence for cultural variation in the relative importance of pairbonding and fathering to these relationships.     

Socio-bioenergetics and sexual dimorphism in primates

Socio-bioenergetics is presented as a practical method of estimating energy budgets of primates in a social context. Energy budgets are estimated on the basis of behavioral observations and a series of empirical formulae, which consider body weight, activity, and reproductive status. Data on a captive colony of Sykes' monkeys and baboons are incorporated as illustrations of the possible effects of group composition, body size, reproductive status, and activity patterns on energy requirements.Supported by the Wenner-Gren Foundation for Anthropological Research Incorporated and the National Science Foundation Grant GU-1598.     

Autonomic nervous system responses to odours: the role of pleasantness and arousal

Perception of odours can provoke explicit reactions such as judgements of intensity or pleasantness, and implicit output such as skin conductance or heart rate variations. The main purpose of the present experiment was to ascertain: (i) the correlation between odour ratings (intensity, arousal, pleasantness and familiarity) and activation of the autonomic nervous system, and (ii) the inter-correlation between self-report ratings on intensity, arousal, pleasantness and familiarity dimensions in odour perception. Twelve healthy volunteers were tested in two separate sessions. Firstly, subjects were instructed to smell six odorants (isovaleric acid, thiophenol, pyridine, L-menthol, isoamyl acetate, and 1-8 cineole), while skin conductance and heart rate variations were being measured. During this phase, participants were not asked to give any judgement about the odorants. Secondly, subjects were instructed to rate the odorants on dimensions of intensity, pleasantness, arousal and familiarity (self-report ratings), by giving a mark between 1 (not at all intense, arousing, pleasant or familiar) and 9 (extremely intense, arousing, pleasant or familiar). Results indicated: (i) a pleasantness factor correlated with heart rate variations, (ii) an arousal factor correlated with skin conductance variations, and (iii) a strong correlation between the arousal and intensity dimensions. In conclusion, given that these correlations are also found in other studies using visual and auditory stimuli, these findings provide preliminary information suggesting that autonomic variations in response to olfactory stimuli are probably not modality specific, and may be organized along two main dimensions of pleasantness and arousal, at least for the parameters considered (i.e. heart rate and skin conductance).     

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.     

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.     

Sexual size dimorphism and sexual selection in primates

1. In most animals, females are larger than males. Paradoxically, sexual size dimorphism is biased towards males in most mammalian species. An accepted explanation is that sexual dimorphism in mammals evolved by intramale sexual selection. I tested this hypothesis in primates, by relating sexual size dimorphism to seven proxies of sexual selection intensity: operational sex ratio, mating system, intermale competition, group sex ratio, group size, maximum mating percentage (percentage of observed copulations involving the most successful male), and total paternity (a genetic estimate of the percentage of young sired by the most successful male).
2. I fitted phylogenetic generalised least squares models using sexual size dimorphism as the dependent variable and each of the seven measures of intensity of sexual selection as independent variables. I conducted this comparative analysis with data from 50 extant species of primates, including Homo sapiens, Pan troglodytes, and Gorilla spp.
3. Sexual dimorphism was positively related to the four measures of female monopolisation (operational sex ratio, mating system, intermale competition, and group sex ratio) and in some cases to group size, but was not associated with maximum mating percentage or total paternity. Additional regression analyses indicated that maximum mating percentage and total paternity were negatively associated with group size.
4. These results are predicted by reproductive skew theory: in large groups, males can lose control of the sexual behaviour of the other members of the group or can concede reproductive opportunities to others. The results are also consistent with the evolution of sexual size dimorphism before polygyny, due to the effects of natural, rather than sexual, selection. In birds, the study of molecular paternity showed that variance in male reproductive success is much higher than expected by behaviour. In mammals, recent studies have begun to show the opposite trend, i.e. that intensity of sexual selection is lower than expected by polygyny.
5. Results of this comparative analysis of sexual size dimorphism and sexual selection intensity in primates suggest that the use of intramale sexual selection theory to explain the evolution of polygyny and sexual dimorphism in mammals should be reviewed, and that natural selection should be considered alongside sexual selection as an evolutionary driver of sexual size dimorphism and polygyny in mammals.

     

'Concealed ovulation' and sexual signals in primates.

The absence of conspicuous sexual signals in some primates, particularly humans and vervets, has been interpreted as evidence that females of these species are 'concealing' ovulation from males. This conclusion is unjustified: the null hypothesis of no adaptation, that the absence of conspicuous sexual signals has resulted from the absence of selective pressures maintaining such adaptations, is both more parsimonious and better fits the facts. The related suggestion that there has been adaptation among females to conceal ovulation from their own consciousness is also unjustified. What, then, maintains sexual signals in those species that do have them? Many proposed hypotheses for the function of sexual signals do not account for their most puzzling feature: their conspicuousness. According to current theory on the evolution of communication, two explanations seem most plausible: conspicuous sexual signals function to communicate to distant receivers and/or to convince reluctant receivers. There is some empirical support for both hypotheses, but not overwhelming support for either.     

Temporal and sexual variation of leaf-produced pollinator-attracting odours in the dwarf palm

Information on intra-specific variation in pollinator-attracting floral traits provides clues to selective pressures imposed by pollinators. However, these traits also reflect constraints related to floral phenology or morphology. The specific weevil pollinator Derelomus chamaeropsis of the dioecious Mediterranean dwarf palm Chamaerops humilis is attracted by volatile compounds that leaves, and not flowers, release during anthesis. Production of these olfactory cues is thus probably not constrained by any other floral function. This provides the opportunity to study variation of a floral trait that is not produced by a floral organ. We studied volatile compounds emitted by leaves of 12 individual C. humilis over the whole flowering season. The quantity of volatile compounds emitted by leaves reached a maximum when plants required pollinator visits. The relationship between odour emission and floral phenology was slightly different between male and female plants, probably reflecting differences in the exact time at which females and males benefit from pollinator visitation. Male plants produced higher quantities of volatile compounds than females. Odour composition was highly variable among individuals but did not differ between male and female plants. In this system, female C. humilis are pollinated by deceit and pollinators should be selected to avoid visiting them. The absence of sexual difference in blend composition may thus prevent pollinators from discriminating between male and female plants.An erratum to this article can be found at     

Spatial autocorrelation,phylogenetic constraints,and the causes of sexual dimorphism in primates

Cheverud et al. (1985) apply the important and relatively new methodology of spatial autocorrelation to the quantification of phylogenetic constraints on adaptation and illustrate the use of these methods in an allometric study of sexual dimorphism in body size among extant nonhuman primates. Though of potentially broad applicability, the technique was completely overlooked in a recent review of methods to control for the effects of common descent in comparative studies (Bell, 1989). Their approach therefore deserves a wider recognition. However, their specific conclusion, that phytogeny is the primary determinant of patterns of sexual dimorphism among primates, has been uncritically accepted. We present four main methodological problems with their approach that should temper the interpretation of their analysis: biased phylogenetic relatedness scores, biased sample selection, size dependence in sex dimorphism measurement, and deficiencies in selection of a structural path model. We conclude that, even in terms of the analysis by Cheverud and co-workers (1985), phylogenetic inertia is not the primary reason for body size dimorphism.     

Fetal and infant head circumference sexual dimorphism in primates

Studies have shown that after controlling for the effects of body size on brain size, the brains of adult humans, rhesus monkeys, and chimpanzees differ in relative size, where males have a greater volume of cerebral tissue than females. We assess whether head circumference sexual dimorphism is present during early development by evaluating sex differences in relative head circumference in living fetuses and infants within the first year of life. Head circumference is used as a proxy for brain size in the fetus and infant. Femur length is used as a proxy for body length in the fetus. Ultrasonography was used to obtain fetal measures, and anthropometry was used to obtain postnatal measures in humans, rhesus monkeys, baboons, and common marmosets. We show that statistically significant but low levels of head circumference sexual dimorphism are present in humans, rhesus monkeys, and baboons in early life. On average, males have head circumferences about 2% larger than females of comparable femur/body length in humans, rhesus monkeys, and baboons. No evidence for head circumference sexual dimorphism in the common marmoset was found. Dimorphism was present across all body size ranges. We suggest that head circumference sexual dimorphism emerges largely postnatally and increases throughout maturation, particularly in humans who reach adult dimorphism values greater than the monkeys. We suggest that brain dimorphism is not likely to impose an additional energetic burden to the gestating or lactating mother. Finally, some of the problems with ascribing functional significance to brain size sexual dimorphism are discussed, and the energetic implications for brain size sexual dimorphism in infancy are assessed.     

Female sexual arousal: genital anatomy and orgasm in intercourse

In men and women sexual arousal culminates in orgasm, with female orgasm solely from sexual intercourse often regarded as a unique feature of human sexuality. However, orgasm from sexual intercourse occurs more reliably in men than in women, likely reflecting the different types of physical stimulation men and women require for orgasm. In men, orgasms are under strong selective pressure as orgasms are coupled with ejaculation and thus contribute to male reproductive success. By contrast, women's orgasms in intercourse are highly variable and are under little selective pressure as they are not a reproductive necessity. The proximal mechanisms producing variability in women's orgasms are little understood. In 1924 Marie Bonaparte proposed that a shorter distance between a woman's clitoris and her urethral meatus (CUMD) increased her likelihood of experiencing orgasm in intercourse. She based this on her published data that were never statistically analyzed. In 1940 Landis and colleagues published similar data suggesting the same relationship, but these data too were never fully analyzed. We analyzed raw data from these two studies and found that both demonstrate a strong inverse relationship between CUMD and orgasm during intercourse. Unresolved is whether this increased likelihood of orgasm with shorter CUMD reflects increased penile–clitoral contact during sexual intercourse or increased penile stimulation of internal aspects of the clitoris. CUMD likely reflects prenatal androgen exposure, with higher androgen levels producing larger distances. Thus these results suggest that women exposed to lower levels of prenatal androgens are more likely to experience orgasm during sexual intercourse.     

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.     

Sexual desire, sexual arousal and hormonal differences in premenopausal US and Dutch women with and without low sexual desire

The interaction between women's hormonal condition and subjective, physiological, and behavioral indices of desire or arousal remains only partially explored, in spite of frequent reports from women about problems with a lack of sexual desire. The present study recruited premenopausal women at two sites, one in the United States and the other in the Netherlands, and incorporated various measures of acute changes in sexual desire and arousal. A sample of 46 women who met criteria for Hypoactive Sexual Desire Disorder (HSDD) was compared to 47 women who experienced no sexual problems (SF). Half of each group used oral contraceptives (OCs). The specific goal was to investigate whether there is a relationship between women's hormone levels and their genital and subjective sexual responsiveness. Background demographics and health variables, including oral contraceptive (OC) use, were recorded and hormones (total testosterone (T), free testosterone (FT), SHBG, and estradiol) were analyzed along with vaginal pulse amplitude and self-report measures of desire and arousal in response to sexual fantasy, visual sexual stimuli, and photos of men's faces. Self-reported arousal and desire were lower in the HSDD than the SF group, but only for women who were not using oral contraceptives. Relationships between hormones and sexual function differed depending on whether a woman was HSDD or not. In line with prior literature, FT was positively associated with physiological and subjective sexual arousal in the SF group. The HSDD women demonstrated the opposite pattern, in that FT was negatively associated with subjective sexual responsiveness. The findings suggest a possible alternative relationship between hormones and sexual responsiveness in women with HSDD who have characteristics similar to those in the present study.     

Sexy thoughts: effects of sexual cognitions on testosterone, cortisol, and arousal in women

Previous research suggests that sexual stimuli increase testosterone (T) in women and shows inconsistent effects of sexual arousal on cortisol (C), but effects of cognitive aspects of arousal, rather than behaviors or sensory stimuli, are unclear. The present study examined whether sexual thoughts affect T or C and whether hormonal contraceptive (HC) use moderated this effect, given mixed findings of HC use confounding hormone responses. Participants (79 women) provided a baseline saliva sample for radioimmunoassay. We created the Imagined Social Situation Exercise (ISSE) to test effects of imagining social interactions on hormones, and participants were assigned to the experimental (sexual) or one of three control (positive, neutral, stressful) conditions. Participants provided a second saliva sample 15 min post-activity. Results indicated that for women not using HCs, the sexual condition increased T compared to the stressful or positive conditions. In contrast, HC using women in the sexual condition had decreased T relative to the stressful condition and similar T to the positive condition. The effect was specific to T, as sexual thoughts did not change C. For participants in the sexual condition, higher baseline T predicted larger increases in sexual arousal but smaller increases in T, likely due to ceiling effects on T. Our results suggest that sexual thoughts change T but not C, baseline T levels and HC use may contribute to variation in the T response to sexual thoughts, and cognitive aspects of sexual arousal affect physiology.     

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.     

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.