Women's preferences for men's scents associated with testosterone and cortisol levels: Patterns across the ovulatory cycle

Women in the fertile phase of their menstrual cycle show an enhanced sexual preference for masculine expressions in behavioral, morphological and scent traits. These masculinity preferences may be associated with testosterone (T) levels in males and hence connote male quality as a sire. Thus, a scent preference of fertile-phase women for T is predicted. A recent study, however, found no evidence for this, but reported that women prefer the scent of men with high cortisol (C). That study had low power to detect the predicted effect, as well as other methodological limitations. We tested women's preferences across their ovulatory cycle for the body scent of men who varied in T and C, using a larger sample of men and methods used in research on cycle preferences for symmetry-related male body scent. Conception risk in the cycle positively predicted women's scent ratings of men's T; scent ratings of C or T × C interaction were not robustly related to conception risk. Conception risk is related positively to a preference for scent of men's symmetry. This preference is distinct from that arising from a preference for the scent of T. The male-emitted chemical(s) responsible for these preferences shifts across women's cycle remain unknown.     

Hormonal and morphological predictors of women’s body attractiveness

Does women’s body attractiveness predict indices of reproductive capacity? Prior research has provided evidence that large breast size and low waist-to-hip ratio (WHR) are positively associated with women’s estrogen and progesterone concentrations, but no previous studies appear to have directly tested whether ratings of women's body attractiveness are predicted by higher concentrations of ovarian hormones measured across broad regions of the menstrual cycle. Here, we collected daily saliva samples across 1–2 menstrual cycles from a sample of young women; assayed the samples for estradiol, progesterone, and testosterone; obtained anthropometric measurements of the women’s bodies; and also obtained attractiveness ratings of the women’s bodies from photographs of them taken in standardized clothing with faces obscured. Contrary to previous research, mean hormone concentrations were uncorrelated with breast size and WHR. Body mass index (BMI) was a very strong negative predictor of body attractiveness ratings, similar to previous findings. Zero-order associations between women’s mean hormone concentrations and mean attractiveness ratings were not significant; however, after controlling for BMI, attractiveness ratings were independently and positively associated with both estradiol and testosterone concentrations. Discussion focuses on the implications of these findings for whether attractiveness assessment mechanisms are specialized for the detection of cues of differential fecundity in young women’s bodies.     

Menstrual cycle variation in women's preferences for the scent of symmetrical men.

Evidence suggests that female sexual preferences change across the menstrual cycle. Women''s extra-pair copulations tend to occur in their most fertile period, whereas their intra-pair copulations tend to be more evenly spread out across the cycle. This pattern is consistent with women preferentially seeking men who evidence phenotypic markers of genetic benefits just before and during ovulation. This study examined whether women''s olfactory preferences for men''s scent would tend to favour the scent of more symmetrical men, most notably during the women''s fertile period. College women sniffed and rated the attractiveness of the scent of 41 T-shirts worn over a period of two nights by different men. Results indicated that normally cycling (non-pill using) women near the peak fertility of their cycle tended to prefer the scent of shirts worn by symmetrical men. Normally ovulating women at low fertility within their cycle, and women using a contraceptive pill, showed no significant preference for either symmetrical or asymmetrical men''s scent. A separate analysis revealed that, within the set of normally cycling women, individual women''s preference for symmetry correlated with their probability of conception, given the actuarial value associated with the day of the cycle they reported at the time they smelled the shirts. Potential sexual selection processes and proximate mechanisms accounting for these findings are discussed.     

Changes in salivary estradiol predict changes in women's preferences for vocal masculinity

Although many studies have reported that women's preferences for masculine physical characteristics in men change systematically during the menstrual cycle, the hormonal mechanisms underpinning these changes are currently poorly understood. Previous studies investigating the relationships between measured hormone levels and women's masculinity preferences tested only judgments of men's facial attractiveness. Results of these studies suggested that preferences for masculine characteristics in men's faces were related to either women's estradiol or testosterone levels. To investigate the hormonal correlates of within-woman variation in masculinity preferences further, here we measured 62 women's salivary estradiol, progesterone, and testosterone levels and their preferences for masculine characteristics in men's voices in five weekly test sessions. Multilevel modeling of these data showed that changes in salivary estradiol were the best predictor of changes in women's preferences for vocal masculinity. These results complement other recent research implicating estradiol in women's mate preferences, attention to courtship signals, sexual motivation, and sexual strategies, and are the first to link women's voice preferences directly to measured hormone levels.     

Do more attractive women show stronger preferences for male facial masculinity?

Researchers have suggested that more attractive women will show stronger preferences for masculine men because such women are better placed to offset the potential costs of choosing a masculine mate. However, evidence for correlations between measures of women's own attractiveness and preferences for masculine men is mixed. Moreover, the samples used to test this hypothesis are typically relatively small. Consequently, we conducted two large-scale studies that investigated possible associations between women's preferences for facial masculinity and their own attractiveness as assessed from third-party ratings of their facial attractiveness (Study 1, N = 454, laboratory study) and self-rated attractiveness (Study 2, N = 8972, online study). Own attractiveness was positively correlated with preferences for masculine men in Study 2 (self-rated attractiveness), but not Study 1 (third-party ratings of facial attractiveness). This pattern of results is consistent with the proposal that women's beliefs about their own attractiveness, rather than their physical condition per se, underpins attractiveness-contingent masculinity preferences.     

The Scent of Symmetry: A Human Sex Pheromone that Signals Fitness?

A previous study by the authors showed that the body scent of men who have greater body bilateral symmetry is rated as more attractive by normally ovulating (non-pill-using) women during the period of highest fertility based on day within the menstrual cycle. Women in low-fertility phases of the cycle and women using hormone-based contraceptives do not show this pattern. The current study replicated these findings with a larger sample and statistically controlled for men's hygiene and other factors that were not controlled in the first study. The current study also examined women's scent attractiveness to men and found no evidence that men prefer the scent of symmetric women. We propose that the scent of symmetry is an honest signal of phenotypic and genetic quality in the human male, and chemical candidates are discussed. In both sexes, facial attractiveness (as judged from photos) appears to predict body scent attractiveness to the opposite sex. Women's preference for the scent associated with men's facial attractiveness is greatest when their fertility is highest across the menstrual cycle. The results overall suggest that women have an evolved preference for sires with good genes.     

Facial attractiveness is related to women's cortisol and body fat,but not with immune responsiveness

Recent studies suggest that facial attractiveness indicates immune responsiveness in men and that this relationship is moderated by stress hormones which interact with testosterone levels. However, studies testing whether facial attractiveness in women signals their immune responsiveness are lacking. Here, we photographed young Latvian women, vaccinated them against hepatitis B and measured the amount of specific antibodies produced, cortisol levels and percentage body fat. Latvian men rated the attractiveness of the women''s faces. Interestingly, in women, immune responsiveness (amount of antibodies produced) did not predict facial attractiveness. Instead, plasma cortisol level was negatively associated with attractiveness, indicating that stressed women look less attractive. Fat percentage was curvilinearly associated with facial attractiveness, indicating that being too thin or too fat reduces attractiveness. Our study suggests that in contrast to men, facial attractiveness in women does not indicate immune responsiveness against hepatitis B, but is associated with two other aspects of long-term health and fertility: circulating levels of the stress hormone cortisol and percentage body fat.     

The role of facial hair in women's perceptions of men's attractiveness,health, masculinity and parenting abilities

Facial hair strongly influences people's judgments of men's socio-sexual attributes. However, the nature of these judgments is often contradictory. The levels of intermediate facial hair growth presented to raters and the stage of female raters' menstrual cycles might have influenced past findings. We quantified men's and women's judgments of attractiveness, health, masculinity and parenting abilities for photographs of men who were clean-shaven, lightly or heavily stubbled and fully bearded. We also tested the effect of the menstrual cycle and hormonal contraceptive use on women's ratings. Women judged faces with heavy stubble as most attractive and heavy beards, light stubble and clean-shaven faces as similarly less attractive. In contrast, men rated full beards and heavy stubble as most attractive, followed closely by clean-shaven and light stubble as least attractive. Men and women rated full beards highest for parenting ability and healthiness. Masculinity ratings increased linearly as facial hair increased, and this effect was more pronounced in women in the fertile phase of the menstrual cycle, although attractiveness ratings did not differ according to fertility. Our findings confirm that beardedness affects judgments of male socio-sexual attributes and suggest that an intermediate level of beardedness is most attractive while full-bearded men may be perceived as better fathers who could protect and invest in offspring.     

Men's Preferences for Women's Femininity in Dynamic Cross-Modal Stimuli

Men generally prefer feminine women''s faces and voices over masculine women''s faces and voices, and these cross-modal preferences are positively correlated. Men''s preferences for female facial and vocal femininity have typically been investigated independently by presenting soundless still images separately from audio-only vocal recordings. For the first time ever, we presented men with short video clips in which dynamic faces and voices were simultaneously manipulated in femininity/masculinity. Men preferred feminine men''s faces over masculine men''s faces, and preferred masculine men''s voices over feminine men''s voices. We found that men preferred feminine women''s faces and voices over masculine women''s faces and voices. Men''s attractiveness ratings of both feminine and masculine faces were increased by the addition of vocal femininity. Also, men''s attractiveness ratings of feminine and masculine voices were increased by the addition of facial femininity present in the video. Men''s preferences for vocal and facial femininity were significantly and positively correlated when stimuli were female, but not when they were male. Our findings complement other evidence for cross-modal femininity preferences among male raters, and show that preferences observed in studies using still images and/or independently presented vocal stimuli are also observed when dynamic faces and voices are displayed simultaneously in video format.     

Attunement to the fertility status of same-sex rivals: women's testosterone responses to olfactory ovulation cues

Evolutionary theories of mating suggest that changes in fertility across the menstrual cycle play an important role in sexual selection. In line with this framework, the current research examined whether olfactory cues to the fertility of a same-sex rival would prompt hormonal signs of intrasexual competition in women. Women exposed to the scent of another woman close to ovulation subsequently displayed higher levels of testosterone than women exposed to the scent of a woman far from ovulation. Whereas women exposed to the scent of a woman in the mid-luteal phase displayed sizable decreases in testosterone over time, no such decline was observed among women exposed to the scent of a woman near ovulation. Thus, olfactory cues signaling a rival's heightened level of fertility were associated with endocrinological responses in women that could be linked to intrasexual competition.     

Scent attractiveness and endocrine status in male students before and during a stress situation

Scent attractiveness of sweat samples collected from male students before and during theoretical exams was assessed by female students. Five-rank scale was used for subjective assessment of the scent samples in term unpleasant/pleasant. Scent attractiveness depended on physiological conditions of both recipients and donors. Male students with low basal concentration of salivary cortisol smelt better than students with high level of cortisol. High level of salivary testosterone also was associated with low scent attractiveness of the male students, but only for the recipients in non-receptive phase of menstrual cycle. In all, the females who were in receptive phase of menstrual cycle assessed scent attractiveness of male students higher then the females in non-receptive phase. Exam stress coincided with increase of the salivary cortisol resulting in decline of scent attractiveness in male students. The negative effect of exam was most prominent in students that failed exam and in student who aspired to be the best but failed. So, the exam stress and basal variation of stress-related physiological indexes, such as salivary cortisol, are mirrored in male chemical signals, which are recognized by females.     

Duration of oral contraceptive use predicts women's initial and subsequent subjective responses to sexual stimuli

Recent work suggests that a woman's hormonal state when first exposed to visual sexual stimuli (VSS) modulates her initial and subsequent responses to VSS. The present study investigated whether women's initial hormonal state was related to their subjective ratings of VSS, and whether this relationship differed with VSS content. We reanalyzed previously collected data from 14 naturally cycling (NC) women and 14 women taking oral contraceptives (OCs), who subjectively rated VSS at three hormonal time-points. NC women's ratings of 216 unique sexual images were collected during the menstrual, periovulatory, and luteal phases of their menstrual cycles, and OC women's ratings were collected at comparable time-points across their pill-cycles. NC women's initial hormonal state was not related to their ratings of VSS. OC women's initial hormonal state predicted their ratings of VSS with minimal contextual information and of images depicting female-to-male oral sex. Specifically, women who entered the study in the third week of their pill-cycle (OC-3 women) rated such images as less attractive at all testing sessions than did all other women. OC-3 women were also the only women to rate decontextualized VSS as unattractive at all testing sessions. These results corroborate previous studies in which women's initial hormonal state was found to predict subsequent interest in sexual stimuli. Future work, with larger samples, should more directly investigate whether OC-3 women's negative assessment of specific types of VSS reflects a reaction to the laboratory environment or a broader mechanism, wherein OC women's sexual interests decrease late in their pill-cycle.     

Lack of support for relation between woman's masculinity preference,estradiol level and mating context

It has been proposed that women's preferences for male facial sexual dimorphism are positively correlated with conception probability and differ between short- and long-term mating contexts. In this study, we tested this assumption by analyzing relationships between estradiol levels to the women's preferences of male faces that were manipulated to vary in masculinity. Estradiol was measured in daily saliva samples throughout the entire menstrual cycle collected by Polish women with regular menstrual cycles. In our analyses, we included the three most commonly used definitions of the fertile window in the literature. After computing the overall masculinity preference of each participant and measuring hormone levels, we found that i) the timing of ovulation varied greatly among women (between − 11 and − 17 days from the onset of the next menses, counting backwards), ii) there was no relationship between daily, measured during the day of the test (N = 83) or average for the cycle (N = 115) estradiol levels and masculinity preferences, iii) there were no differences in masculinity preferences between women in low- and high-conception probability phases of the cycle, and iv) there were no differences in masculinity preferences between short- and long-term mating contexts. Our results do not support the idea that women's preferences for a potential sexual partner's facial masculinity fluctuate throughout the cycle.     

Evidence for the stress-linked immunocompetence handicap hypothesis in human male faces

The stress-linked immunocompetence handicap hypothesis (SL-ICHH) of sexual selection incorporates a role of the stress hormone corticosterone (C; cortisol in humans) in relationships between testosterone (T), immunity and secondary sexual trait expression. In support of this, C has been shown to mediate and moderate relationships between T and immune response and to be inversely related to attractiveness in some avian species. We predicted that female preferences for cues to T in human male faces would be contingent upon co-occurring cortisol levels. In study 1, we tested relationships between T and cortisol and attractiveness, masculinity and health ratings of raw male faces. We found cortisol to be inversely related to attractiveness. In study 2, we tested female preferences for male faces that were parametrically manipulated on the basis of cues to naturally co-occurring levels of T and cortisol across the menstrual cycle. Women preferred cues to low cortisol in general and in the fertile phase of the cycle, and there was an interaction between T and cortisol in general and in the non-fertile phase. Results were consistent with the SL-ICHH but not the original immunocompetence handicap model: females expressed preferences for cues to cortisol but not for cues to T, except in interaction with the stress hormone. Results inform the SL-ICHH by demonstrating female preferences for low cortisol and the nature of its interaction with T in humans, as well as indicating the traits that may be signalled by different combinations of the hormones including immune response, current health and resource acquisition characteristics.     

Increasing women's sexual desire: The comparative effectiveness of estrogens and androgens

Both estradiol and testosterone have been implicated as the steroid critical for modulating women's sexual desire. By contrast, in all other female mammals only estradiol has been shown to be critical for female sexual motivation and behavior. Pharmaceutical companies have invested heavily in the development of androgen therapies for female sexual desire disorders, but today there are still no FDA approved androgen therapies for women. Nonetheless, testosterone is currently, and frequently, prescribed off-label for the treatment of low sexual desire in women, and the idea of testosterone as a possible cure-all for female sexual dysfunction remains popular. This paper places the ongoing debate concerning the hormonal modulation of women's sexual desire within a historical context, and reviews controlled trials of estrogen and/or androgen therapies for low sexual desire in postmenopausal women. These studies demonstrate that estrogen-only therapies that produce periovulatory levels of circulating estradiol increase sexual desire in postmenopausal women. Testosterone at supraphysiological, but not at physiological, levels enhances the effectiveness of low-dose estrogen therapies at increasing women's sexual desire; however, the mechanism by which supraphysiological testosterone increases women's sexual desire in combination with an estrogen remains unknown. Because effective therapies require supraphysiological amounts of testosterone, it remains unclear whether endogenous testosterone contributes to the modulation of women's sexual desire. The likelihood that an androgen-only clinical treatment will meaningfully increase women's sexual desire is minimal, and the focus of pharmaceutical companies on the development of androgen therapies for the treatment of female sexual desire disorders is likely misplaced.     

Menstrual cycle shifts in attentional bias for courtship language

The current study investigated whether women show an attentional bias toward courtship language during the fertile phase of their menstrual cycle. Thirty heterosexual women (17 naturally cycling, 13 using hormonal contraceptives) completed a dichotic listening task on both a high and low fertility day of their menstrual cycle. Participants were asked to verbally repeat (shadow) an emotionally neutral target passage played in one ear while either a neutral or courtship distracter was played in the other ear. Courtship distracters were flirtatious in content but not overtly sexual. Shadowing errors were coded as a measure of attentional bias toward the distracter. Saliva samples were taken to determine whether levels of estradiol, progesterone and/or testosterone correlated with task performance. As predicted, naturally cycling women made more shadowing errors when listening to a courtship distracter during the fertile phase of their cycle than during the nonfertile phase. This effect was moderated by relationship status, such that fertile, mated women showed an attentional bias for courtship language but fertile single women did not. However, because of small sample sizes in the analysis, this relationship should be viewed as preliminary. Hormonal analysis revealed that higher levels of salivary estradiol predicted greater attentional bias toward courtship language in naturally cycling women. These results suggest that women's attention is drawn to verbal courtship signals when they are fertile, and that this shift is linked to increased estradiol release during the periovulatory phase.     

Men report stronger attraction to femininity in women's faces when their testosterone levels are high

Many studies have shown that women's judgments of men's attractiveness are affected by changes in levels of sex hormones. However, no studies have tested for associations between changes in levels of sex hormones and men's judgments of women's attractiveness. To investigate this issue, we compared men's attractiveness judgments of feminized and masculinized women's and men's faces in test sessions where salivary testosterone was high and test sessions where salivary testosterone was relatively low. Men reported stronger attraction to femininity in women's faces in test sessions where salivary testosterone was high than in test sessions where salivary testosterone was low. This effect was found to be specific to judgments of opposite-sex faces. The strength of men's reported attraction to femininity in men's faces did not differ between high and low testosterone test sessions, suggesting that the effect of testosterone that we observed for judgments of women's faces was not due to a general response bias. Collectively, these findings suggest that changes in testosterone levels contribute to the strength of men's reported attraction to femininity in women's faces and complement previous findings showing that testosterone modulates men's interest in sexual stimuli.     

Intelligence and mate choice: intelligent men are always appealing

What role does a man's intelligence play in women's mate preferences? Selecting a more intelligent mate often provides women with better access to resources and parental investment for offspring. But this preference may also provide indirect genetic benefits in the form of having offspring who are in better physical condition, regardless of parental provisioning. Intelligence then may serve as both a cue of a mate's provisioning abilities and his overall heritable phenotypic quality. In the current study, we examined the role of a man's intelligence in women's long- and short-term mate preferences. We used a rigorous psychometric measure (men's WAIS scores) to assess intelligence (the first study to our knowledge), in addition to women's subjective ratings to predict mate appeal. We also examined the related trait of creativity, using women's ratings as a first step, to assess whether creativity could predict mate appeal, above and beyond intelligence. Finally, we examined whether preferences for intelligent and creative short-term mates shifted according to a woman's conception risk. Multilevel modeling was used to identify predictors of mate appeal. Study participants (204 women) assessed the long- and short-term mate appeal of videos of 15 men with known measures of intelligence performing verbal and physical tasks. Findings indicate that both intelligence and creativity independently predicted mate appeal across mating contexts, but no conception-risk effects were detected. We discuss implications of these findings for the role of intelligence and creativity in women's mate choices.     

Salivary hormone concentrations in mothers and fathers becoming parents are not correlated

A time- and date-matched set of saliva samples (N = 229) from nine couples first expecting, and then caring for, their first child were used to test whether hormone changes in the father could be predicted by the hormonal status of the mother. Testosterone, cortisol, and estradiol were quantified from saliva. Neither testosterone nor estradiol concentration was correlated within couples before or after the birth, although there was a positive correlation for cortisol concentration in the mother and father before the birth. As the hormone that might be influenced by chemical signals, that already played a similar role in men and women, and that had been empirically linked to paternal behavior, cortisol concentration was also compared with sex steroid concentrations. The mother's cortisol concentration was positively correlated to the father's testosterone concentration, and the mother's testosterone concentration was positively correlated with the father's cortisol concentration. However, both effects were similar in magnitude to the cortisol to cortisol correlation, and all could parsimoniously be explained by similar responses to a shared environment. Thus, this analysis rejects parallels in peripheral hormone concentrations of estradiol, testosterone, and cortisol in mothers and fathers. However, the available data were not able to test or reject hypotheses about local neuroendocrine homology, nor to control for masking effects of other hormonal demands on men and women, nor to determine the relative importance of shared environment versus mother-father signaling.     

Women's behavioural engagement with a masculine male heightens during the fertile window: evidence for the cycle shift hypothesis

Previous research suggests that women may alter their behaviour during the fertile window of the menstrual cycle to attract a mate who has traits that indicate high-quality genes. We tested whether fertile women demonstrate greater behavioural engagement with a masculine compared to a less masculine male. The test was performed using a quiz show paradigm, in which a male host asked female participants general knowledge questions. The masculinity of the host was varied between participants. Women's performance on the quiz, as well as their romantic attraction to the host, was examined in relation to women's estimated cycle phase and host masculinity. Fertile compared to nonfertile women were more romantically attracted to the host and were faster to answer his questions, but only when he was portrayed as masculine. The results of the study are interpreted as being in keeping with Gangestad and Thornhill's cycle shift hypothesis (Menstrual cycle variation in women's preferences for the scent of symmetrical men. Proceedings of the Royal Society of London B: Biological Sciences, 1998;265:727–733. doi:10.1098/rspb.1998.03801998).