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.     

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.     

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.     

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.     

Women's sexual arousal: effects of high alcohol dosages and self-control instructions

The basic relationship between alcohol and women's sexual arousal – especially genital arousal – received little research attention for nearly 30 years (e.g. Wilson and Lawson, 1978) until very recently (e.g. George et al., 2009). To investigate hypotheses based on earlier findings and Alcohol Myopia Theory (AMT), two experiments evaluated the effects of high blood alcohol concentrations (BACs) and arousal instructional demands on indices of vaginal responding and self-reported sexual arousal. In Experiment 1, self-control instructions to maximize (versus suppress) arousal increased peak and average Vaginal Pulse Amplitude (VPA) change. Self-control also interacted with a target BAC of .08% (versus .00%) to influence latency to peak arousal onset: Intoxicated women instructed to maximize showed a shorter latency to peak arousal than did intoxicated women instructed to suppress; however, sober women showed an undifferentiated pattern. Also, in Experiment 1, the target BAC of .08% had no effect on VPA or subjective arousal measures. In Experiment 2, a target BAC of .10% (versus .00%) attenuated peak change and average change in VPA, but this dosage had no effects on latency to peak achieved arousal, or on subjective arousal. Instructions to maximize arousal (versus no instruction) had no effect on any arousal measures. Overall, among young moderate drinking women, alcohol had attenuating effects but only at the higher dosage. Maximize versus suppress instructions about arousal had predicted effects on arousal and interactive effects on latency, but only at the lower dosage. The findings highlight the importance of dosage and contextual factors in alcohol's impact on the variability of women's sexual responding.     

Sex differences in viewing sexual stimuli: an eye-tracking study in men and women

Men and women exhibit different neural, genital, and subjective arousal responses to visual sexual stimuli. The source of these sex differences is unknown. We hypothesized that men and women look differently at sexual stimuli, resulting in different responses. We used eye tracking to measure looking by 15 male and 30 female (15 normal cycling (NC) and 15 oral contracepting (OC)) heterosexual adults viewing sexually explicit photos. NC Women were tested during their menstrual, periovulatory, and luteal phases while Men and OC Women were tested at equivalent intervals, producing three test sessions per individual. Men, NC, and OC Women differed in the relative amounts of first looks towards, percent time looking at, and probability of looking at, defined regions of the pictures. Men spent more time, and had a higher probability of, looking at female faces. NC Women had more first looks towards, spent more time, and had a higher probability of, looking at genitals. OC Women spent more time, and had a higher probability of, looking at contextual regions of pictures, those featuring clothing or background. Groups did not differ in looking at the female body. Menstrual cycle phase did not affect women's looking patterns. However, differences between OC and NC groups suggest hormonal influences on attention to sexual stimuli that were unexplained by subject characteristic differences. Our finding that men and women attend to different aspects of the same visual sexual stimuli could reflect pre-existing cognitive biases that possibly contribute to sex differences in neural, subjective, and physiological arousal.     

Feelings of Disgust and Disgust-Induced Avoidance Weaken following Induced Sexual Arousal in Women

Background

Sex and disgust are basic, evolutionary relevant functions that are often construed as paradoxical. In general the stimuli involved in sexual encounters are, at least out of context strongly perceived to hold high disgust qualities. Saliva, sweat, semen and body odours are among the strongest disgust elicitors. This results in the intriguing question of how people succeed in having pleasurable sex at all. One possible explanation could be that sexual engagement temporarily reduces the disgust eliciting properties of particular stimuli or that sexual engagement might weaken the hesitation to actually approach these stimuli.

Methodology

Participants were healthy women (n = 90) randomly allocated to one of three groups: the sexual arousal, the non-sexual positive arousal, or the neutral control group. Film clips were used to elicit the relevant mood state. Participants engaged in 16 behavioural tasks, involving sex related (e.g., lubricate the vibrator) and non-sex related (e.g., take a sip of juice with a large insect in the cup) stimuli, to measure the impact of sexual arousal on feelings of disgust and actual avoidance behaviour.

Principal Findings

The sexual arousal group rated the sex related stimuli as less disgusting compared to the other groups. A similar tendency was evident for the non-sex disgusting stimuli. For both the sex and non-sex related behavioural tasks the sexual arousal group showed less avoidance behaviour (i.e., they conducted the highest percentage of tasks compared to the other groups).

Significance

This study has investigated how sexual arousal interplays with disgust and disgust eliciting properties in women, and has demonstrated that this relationship goes beyond subjective report by affecting the actual approach to disgusting stimuli. Hence, this could explain how we still manage to engage in pleasurable sexual activity. Moreover, these findings suggest that low sexual arousal might be a key feature in the maintenance of particular sexual dysfunctions.     

Effects of affiliation and power motivation arousal on salivary progesterone and testosterone

Following up on earlier research suggesting a link between implicit affiliation motivation and progesterone (P) and implicit power motivation and testosterone [T; Schultheiss, O.C., Dargel, A., Rohde, W., 2003. Implicit motives and gonadal steroid hormones: Effects of menstrual cycle phase, oral contraceptive use, and relationship status. Horm. Behav. 43, 293-301.], we tested whether arousal of affiliation motivation increases P levels and whether arousal of power motivation increases T levels. Sixty subjects were randomly assigned to watch 30 min of either Bridges of Madison County (affiliation arousal) or The Godfather II (power arousal), or a documentary about the Amazon (control condition). Levels of P and T were assessed in saliva samples taken before (T1), immediately after (T2), and 45 min after the movie (T3). The efficacy of experimental conditions to differentially arouse motives was verified by assessment of changes in affiliation and power motive imagery expressed in imaginative stories written before and after the movie. After the movie, salivary P levels (T2 and T3) in the affiliation-arousal group were significantly higher than in the control group and marginally higher than in the power-arousal group. Subjects' postmovie T responses (T3) depended on premovie T levels: in men, higher premovie T levels predicted a greater likelihood of postmovie T increases in the Power Arousal condition but not in the other conditions, whereas in women, higher premovie T levels tended to be associated with postmovie T decreases in the Power Arousal condition but not in the other conditions. These findings suggest that aroused affiliation motivation has a specific stimulatory effect on P, whereas aroused power motivation has a specific stimulatory effect on T in men, but not in women, with high baseline T levels.     

Neural activation in the orbitofrontal cortex in response to male faces increases during the follicular phase

Women's sexual interest changes with hormonal fluctuations across the menstrual cycle. It is unclear how hormones modify women's sexual behavior and desire, but one possibility is that they alter women's positive appraisals of stimuli and thus their sexual interest. Using 3 T fMRI, we measured neural activation in women at two time points in their menstrual cycle (late follicular, luteal) while they evaluated photos of men presented as potential sexual partners. Participants were ten heterosexual women aged 23-28 none of who was using hormonal contraceptives or in a committed relationship. In an event-related design, the women were presented with as series of photos of male faces and asked questions to assess their degree of sexual interest in the men depicted. Results demonstrate an overall effect of menstrual cycle phase on neural activation. During their follicular versus luteal phase, women demonstrated increased activation in the right medial orbitofrontal cortex (OFC), suggesting increased positive appraisal. Activation in the OFC was positively correlated with women's estradiol to progesterone ratios. There were no areas that demonstrated increased activation during the luteal versus follicular phase. The observed increase in activation in the OFC during the follicular phase may reflect a hormonally mediated increase in appetitive motivation and may prime women towards increased sexual interest and behavior around ovulation.     

Androgenic hormones and aging--the link with female sexual function

In women, sexual function, hormones and aging are inextricably related. Sexual activity in women involves interest and motivation, the ability to become aroused and achieve orgasm, the pleasure of the experience and subsequent personal satisfaction. Androgens, as endogenous hormones or given as a therapy, potentially influence female sexual function, with research into the effects of exogenous androgens in women mostly devoted to effects on sexual desire. Some studies have been conducted to delineate the effects of testosterone on arousal, however arousal determined by laboratory measures does not always correlate with subjective reporting of a sensation of arousal. Overall large randomised controlled trials of exogenous testosterone show benefits over placebo on sexual desire, arousal, orgasm, pleasure and satisfaction. The aspects of consideration of androgen therapy for women that continue to stimulate debate in this therapeutic area include whether female sexual dysfunction is a condition that merits pharmacotherapy, how effective is such treatment and whether testosterone therapy is safe.     

Associations between testosterone secretion and sexual activity in women

Some studies show an increase in testosterone (T) after sexual activity; this literature has inconsistent findings, focuses mostly on men, and does not employ control activities. The present study examined within-subject effects of intercourse versus control activities (cuddling; exercise) on salivary T. The initial sample included 49 women (mostly heterosexual), though not all participants returned all samples or engaged in all activities, leaving a smaller sample for endocrine analyses (n=16). Participants attended an initial session in the laboratory where they completed questionnaires, and then engaged in the activities on their own. On three separate nights, they provided pre-activity, post-activity, and next-morning saliva samples and completed brief questionnaires at the last two timepoints. Women's T was higher pre-intercourse than pre-control activity. Women's T was also higher post-intercourse than post-control activity, though the percent change in T from pre- to post-activity was highest for cuddling, then intercourse, then exercise. Next-morning T did not differ by activity. Data pointed to an association between T and orgasming, sexual desire, and relationship commitment. Analyses on post-activity appraisals suggest that the close intimate physicality of a sexual and non-sexual nature can affect T and be beneficial in short-term and perhaps longer-lasting ways for women's sexuality and relationships.     

Neural correlates of sexual arousal in heterosexual and homosexual women and men

Most men have a category-specific pattern of genital and subjective sexual arousal, responding much more strongly to erotic stimuli depicting their preferred sex than to erotic stimuli depicting their nonpreferred sex. In contrast, women tend to have a less specific arousal pattern. To better understand this sex difference, we used neuroimaging to explore its neural correlates. Heterosexual and homosexual women viewed erotic photographs of either men or women. Evoked neural activity was monitored via fMRI and compared with responses to the same stimuli in heterosexual and homosexual men. Overall, a network of limbic (as well as the anterior cingulate) and visual processing regions showed significantly less category-specific activity in women than men. This was primarily driven by weaker overall activations to preferred-sex stimuli in women, though there was also some evidence of stronger limbic activations to nonpreferred-sex stimuli in women. Primary results were similar for heterosexual and homosexual participants. Women did show some evidence of category-specific responses in the visual processing regions, although even in these regions they exhibited less differential activity than men. In the anterior cingulate, a region with high concentrations of sex-hormone receptors, subjective and neural category specificity measures correlated positively for women but negatively for men, suggesting a possible sex difference in the role of the anterior cingulate. Overall, results suggest that men tend to show more differentiated neural responses than do women to erotic photographs of one sex compared to the other sex, though women may not be entirely indifferent to which sex is depicted.     

A high-powered replication study finds no effect of starting or stopping hormonal contraceptive use on relationship quality

A number of recent studies have implicated that incongruent use of hormonal contraceptives (HCs) negatively affects various aspects of women's romantic relationships. It has been suggested that women with incongruent HC use (a discrepancy in HC use status between when they first met their current partner and the time of study participation) report less sexual satisfaction and higher jealousy scores compared to women with congruent HC use. A similar effect has also been hypothesized for general relationship satisfaction, and recent findings suggest that the association between HC incongruency and women's general relationship satisfaction is moderated by third-party ratings of facial attractiveness of the women's male partners. Using a large convenience sample (N = 948) of Finnish women, we attempted to replicate previously reported findings but found no support for the HC congruency hypothesis, despite excellent statistical power (≥98.7%) to detect previously reported effect sizes. Instead, after dividing our sample into four groups based on HC congruency/incongruency, we found that the largest differences in jealousy, sexual satisfaction, and relationship satisfaction scores tended to be found between women who were consistent HC users and consistent non-users (i.e., between women with different kinds of congruent HC use). We also detected a significant main effect of current HC use on jealousy. We conclude that HC congruency effects reported in previous studies may have spuriously arisen from unequal distributions of current HC users within congruent and incongruent HC user groups.     

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.     

The Roles of Noradrenergic and Glucocorticoid Activation in the Development of Intrusive Memories

Intrusive memories are a common feature of many psychological disorders. Recent evidence has potentially extended cognitive models of intrusions by identifying the role of biological markers of arousal at the time of consolidation in subsequent memory for emotional events. This study investigated the role of arousal during consolidation in the development of intrusive memories. Seventy-eight university students (37 men and 41 women) viewed 20 negative and 20 neutral images. Half the participants then underwent a cold pressor test (High Stress), immersing their hand in ice water, while the remaining participants immersed their hand in warm water (Low Stress). Samples of salivary alpha-amylase (sAA) and cortisol were collected from participants at baseline and following the stressor challenge. Participants completed a delayed free recall test and intrusion questionnaires two days later. Participants in the High Stress condition reported more intrusions of negative images than participants in the Low Stress condition. An interaction variable in a linear regression of increased noradrenergic and cortisol values predicted intrusive memories of emotional stimuli for men but not women. These findings are consistent with recent evidence of the combined effects of noradrenaline and corticoid responses to stress on emotional memories, and also with increasing evidence of gender differences in how stress hormones influence formation of emotional memories. These findings point to possible mechanisms by which development of intrusions may be prevented after consolidation of traumatic experiences.     

Effects of affiliation arousal (hope of closeness) and affiliation stress (fear of rejection) on progesterone and cortisol

Our prior research has suggested a connection between progesterone (PROG) and implicit affiliation motivation, the non-conscious drive for positive social contact. In particular, experimental arousal of affiliation motivation led to relative PROG increase in women and men [Schultheiss, O.C., Wirth, M.M., Stanton, S.J., 2004. Effects of affiliation and power motivation arousal on salivary progesterone and testosterone. Horm. Behav. 46(5), 592-599]. The present study aimed to (1) replicate this effect, (2) simultaneously assess cortisol (CORT) levels in this paradigm in order to rule out non-specific adrenal effects induced by affiliation arousal, and (3) examine effects on PROG and CORT of approach (hope for closeness, HOC) versus avoidance (fear of rejection, FOR) affiliation arousal. These motivational states were experimentally aroused in participants using film segments containing approach- or avoidance-oriented affiliation-related themes; a neutral film segment was used as a control condition. The film segments affected participants' implicit affiliation motivation and self-reported mood, demonstrating effectiveness of the manipulation. In the FOR condition, participants' CORT and PROG were increased post-film, consistent with the idea that fear of rejection is stressful. We did not replicate our prior finding of PROG increase following the HOC manipulation. However, relationships between PROG and implicit affiliation motivation were apparent across conditions. In particular, PROG co-varied positively with affiliation motivation, and baseline affiliation motivation positively predicted PROG increase in the FOR condition. As prior research implicates PROG in down-regulation of stress, we speculate that PROG release during stress may encourage affiliation for stress reduction purposes.     

The role of salivary cortisol and DHEA-S in response to sexual, humorous, and anxiety-inducing stimuli

Stress and anxiety are commonly thought to be detrimental to sexual function. Several studies in both the human and animal literature, however, have found that inducing anxiety can enhance sexual function in women. The mechanisms that explain a negative relationship between physical and psychological stress and sexual functioning are well documented, but little is known about how stress or anxiety might have a facilitatory effect on sexual arousal. As an initial step in exploring the relationship between anxiety and sexual arousal, the present study examined the role of the autonomic nervous system, and the adrenal hormones cortisol and dehydroepiandrosterone-sulfate (DHEA-S) in response to a sexual film, an anxiety-inducing film, and a humorous film. Nineteen premenopausal women (mean age 24.4 years) who were free from sexual difficulties came into the lab on three separate days. At each session they were shown an anxiety-inducing, sexually arousing, or humorous (control) film while their physiological arousal was measured. They also provided saliva samples before and after each film. Cortisol significantly decreased, while DHEA-S increased in the sexual and humorous conditions. Neither hormone changed significantly in the anxiety-inducing condition. Autonomic nervous system activity measured by heart rate and heart rate variability did not change in response to the sexual or anxiety-inducing films, but heart rate variability increased significantly in response to the humorous film. The cortisol/DHEA-S ratio at the post-sexual film time point was significantly negatively correlated with genital arousal (measured by vaginal pulse amplitude). Anxiety-inducing films did not result in a physiological stress response, which can explain why they do not impair sexual function.     

Hormonal responses to different sexually related conditions in male rats

Plasma levels of corticosterone (C) and testosterone (T) increase after sexual activity in males of several species. However, the physiological significance of these increases has not been elucidated. In the present study, hormonal response to different conditions linked to sexual activity was assessed. In the first experiment, plasma levels of C and T were assessed both in sexually experienced and naive male rats after the following conditions: (A) control group, without sexual stimulation; (B) males exposed to ovariectomized females; (C) males exposed to intact, non-receptive females; (D) males exposed to receptive females with the vagina obstructed, to avoid intromission; (E) males exposed to receptive females: but separated by a grid that prevents physical contact; (F) males exposed to receptive females during 30 min. In a second experiment, experienced male rats were allowed to repeatedly copulate until reaching the criteria for sexual exhaustion, and 24 h later, they were allowed to copulate. Once sexually related conditions ended, males were killed and their blood was obtained. C and T plasma levels were assessed by HPLC with ultraviolet (UV) detection. Results indicate that T did not increase significantly in naive male in any sexual condition, while in the experienced males, significant increases were observed with the mere presence of a receptive female and also after ejaculation. These increases were significantly larger in experienced males. On the other hand, C also increased in all sexual conditions, both in experienced and naive rats; however, the increase observed was larger in experienced males. Regarding sexual satiety, both C and T increased after copulating ad libitum to satiety. T increased almost three-fold compared to control, while C increased two-fold. No significant changes were observed in either one of the steroids 24 h after sexual exhaustion, even though males remained with a receptive female during an hour. These results show that sexual experience has an important influence on the hormonal response to sexual activity. C rises could be directly related to sexual arousal involved in the different sexual conditions, while T rises seem to have a direct relationship with both the motivation and execution aspects of masculine sexual behavior.     

Sexual arousal, is it for mammals only?

Sexual arousal has many dimensions and has consequently been defined in various ways. In humans, sexual arousal can be assessed based in part on verbal communication. In male non-human mammalian species, it has been argued that arousal can only be definitively inferred if the subject exhibits a penile erection in a sexual context. In non-mammalian species that lack an intromittent organ, as is the case for most avian species, the question of how to assess sexual arousal has not been thoroughly addressed. Based on studies performed in male Japanese quail, we argue that several behavioral or physiological characteristics provide suitable measures of sexual arousal in birds and probably also in other tetrapods. These indices include, the performance of appetitive sexual behavior in anticipation of copulation (although anticipation and arousal are not synonymous), the activation of specific brain area as identified by the detection of the expression of immediate early genes (fos, egr-1) or by 2-deoxygucose quantitative autoradiography, and above all, by the release of dopamine in the medial preoptic area as measured by in vivo dialysis. Based on these criteria, it is possible to assess in birds sexual arousal in its broadest sense but meeting the more restrictive definition of arousal proposed for male mammals (erection in an explicit sexual context) is and will probably remain impossible in birds until refinement of in vivo imaging techniques such fMRI allow us to match in different species, with and without an intromittent organ, the brain areas that are activated in the presence of specific stimuli.     

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.