Male sexual attractiveness predicts differential ovulatory shifts in female extra-pair attraction and male mate retention

Because ancestral women faced trade-offs in choosing mates, they may have evolved to pursue a dual-mating strategy in which they secured investment through one partner and obtained good genes through others. The dual-mating theory predicts that women will display greater interest in extra-pair sex near ovulation, especially if they are mated to a primary male partner who is low in sexual attractiveness. Forty-three normally ovulating women rated their partner's sexual attractiveness and separately reported their own desires and their partner's mate retention behaviors at high and low fertility (confirmed using luteinizing hormone tests). In the high-fertility session relative to the low, women who assessed their partners as being lower in sexual attractiveness reported greater extra-pair desires and more expressed love and attention from their male partners. Women's desire for their own partners did not differ significantly between high and low-fertility sessions.     

Women's sexual interests across the ovulatory cycle depend on primary partner developmental instability

Normally ovulating women have been found to report greater sexual attraction to men other than their own partners when near ovulation relative to the luteal phase. One interpretation is that women possess adaptations to be attracted to men possessing (ancestral) markers of genetic fitness when near ovulation, which implies that women's interests should depend on qualities of her partner. In a sample of 54 couples, we found that women whose partners had high developmental instability (high fluctuating asymmetry) had greater attraction to men other than their partners, and less attraction to their own partners, when fertile.     

Fertility in the cycle predicts women's interest in sexual opportunism

Research over the past decade has documented clear, robust changes in women's sexual preferences and interests across the ovarian cycle. When fertile, women are particularly attracted to a number of masculine male features (e.g., masculine faces, voices, scents and bodies) and other traits, and especially when they evaluate men's “sexiness” rather than their attractiveness as long-term partners. The current research extended this line of research by examining changes in women's self-reported sexual interests across the cycle. We asked 68 normally ovulating women in committed romantic relationships to fill out questionnaires about their sexual preferences and interests (at that time, not in general) twice across their cycles: once when fertile and once during the luteal phase. Relative to during the luteal phase, fertile women expressed (a) greater emphasis on the physical attractiveness of a partner; (b) greater arousal at the sight or thought of attractive male bodily features; (c) greater willingness to engage in and interest in sex with attractive men, even ones who they do not know well (interest in sexual opportunism). These findings importantly extend our understanding of women's fertile-phase sexuality.     

Hormonal influences on women's extra-pair sexual interests: The moderating impact of partner attractiveness

Based on the idea that women are especially attracted to ancestral markers of male genetic quality when conceptive in their cycle, scholars have conjectured that increases in women's extra-pair sexual interests during the conceptive phase of the cycle are moderated by their primary partners' sexual attractiveness. Multiple studies have examined this prediction, with largely supportive but mixed results. The current study is the first to examine whether hormonal influences—thought to mediate cycle shifts—on women's extra-pair sexual interests are moderated by partner attractiveness. 213 naturally cycling, romantically involved women (181 attending multiple sessions) participated in up to four sessions over about a month. Estrogen and progesterone levels were measured multiple times across women's cycles. Male partner attractiveness moderated associations between progesterone levels (though not estrogen levels) and women's extra-pair sexual interests. A negative association between progesterone levels and extra-pair sexual interests, a composite measure, was observed for women with relatively unattractive partners. For women with relatively attractive partners, no clear association emerged. The interaction between progesterone and partner attractiveness was robust for women's interest in extra-pair sex, a component preregistered as exploratory. The interaction effect was also significant for the absolute intensity of women's extra-pair attraction (a component item referenced in the preregistration) but was non-significant for the frequency of women's extra-pair attraction relative to typical days (a composite component specified in the preregistration). These findings inform theoretical understandings of women's sexuality.     

Conditional expression of women's desires and men's mate guarding across the ovulatory cycle

Thirty-eight normally cycling women provided daily reports of sexual interests and feelings for 35 days. Near ovulation, both pair-bonded and single women reported feeling more physically attractive and having greater interest in attending social gatherings where they might meet men. Pair-bonded women who were near ovulation reported greater extra-pair flirtation and greater mate guarding by their primary partner. As predicted, however, these effects were exhibited primarily by women who perceived their partners to be low on hypothesized good genes indicators (low in sexual attractiveness relative to investment attractiveness). Ovulation-contingent increases in partner mate guarding were also moderated by female physical attractiveness; midcycle increases in mate guarding were experienced primarily by less attractive women, whereas more attractive women experienced relatively high levels of mate guarding throughout their cycle. These findings demonstrate ovulation-contingent shifts in desires and behaviors that are sensitive to varying fitness payoffs, and they provide support for the good genes hypothesis of human female extra-pair mating. The daily assessment method provides an important supplement to existing studies using scheduled laboratory visits as the purpose of the study (examining cycle-related variation) is not known by participants.     

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.     

Women’s sexual strategies in pregnancy

Humans exhibit an unusual pattern of sexual behavior compared to other mammalian females. Women's extended sexuality has been hypothesized to be related to a variety of possible benefits, especially non-genetic reproductive benefits, such as securing male investment via reinforced pairbonds or paternity confusion. But sexual behavior also comes at a cost, particularly for pregnant women, in terms of energetic costs, potential disease, and possible harm to the fetus. We hypothesize, therefore, that sexual behavior in pregnant women should reflect adaptive strategies and that pregnant women will be particularly strategic about their sexual behavior in order to maximize potential benefits while minimizing potential costs. One hundred twelve pregnant women completed a survey of their partners' qualities and their sexual desires toward their primary partners and men other than their primary partners. Results showed that women's perceptions of relationship threat positively predicted sexual desire for primary partners, while their perceptions of their partner's investing qualities negatively predicted sexual desire for extra-pair mates. These qualities, as well as cues to partner's genetic quality and gestation age, also interacted in ways that suggest that pregnant women's sexual desires are sensitive to cues of future investment and relationship stability.     

Women Ornament Themselves for Intrasexual Competition near Ovulation,but for Intersexual Attraction in Luteal Phase

The present study examined women''s attentional bias toward ornamental objects in relation to their menstrual phase as well as to motivations of intersexual courtship or intrasexual competition. In Experiment 1, 33 healthy heterosexual women were tested in a bias-assessment visual cuing task twice: once on a high-fertility day (during the ovulatory phase) and once on a low-fertility day (during the luteal phase). They paid greater attention to pictures of ornamental objects than to pictures of non-ornamental objects near ovulation, but not during the luteal phase, suggesting an ornamental bias during the high-fertility phase. In Experiment 2, before the visual cuing task, 40 participants viewed 10 same-sex or opposite-sex facial photographs with either high or low attractiveness as priming tasks to activate the intrasexual competition or intersexual courtship motives. Results showed that women''s ornamental bias was dependent on the interaction of menstrual phase and mating motive. Specifically, the ornamental bias was observed on the high-fertility day when the subjects were primed with high-attractive same-sex images (intrasexual competition) and was observed on the low-fertility day when they were primed with high-attractive opposite-sex photographs (intersexual courtship). In conclusion, the present findings confirm the hypothesis that, during the high-fertility phase, women have an attentional bias toward ornamental objects and further support the hypothesis that the ornamental bias is driven by intrasexual competition motivation near ovulation, but driven by intersexual courtship motivation during the luteal phase.     

Hormonal predictors of women's extra-pair vs. in-pair sexual attraction in natural cycles: Implications for extended sexuality

In naturally cycling women, Roney and Simmons (2013) examined hormonal correlates of their desire for sexual contact. Estradiol was positively associated, and progesterone negatively associated, with self-reported desire. The current study extended these findings by examining, within a sample of 33 naturally cycling women involved in romantic relationships, hormonal correlates of sexual attraction to or interests in specific targets: women's own primary partner or men other than women's primary partner. Women's sexual interests and hormone (estradiol, progesterone, and testosterone) levels were assessed at two different time points. Whereas estradiol levels were associated with relatively greater extra-pair sexual interests than in-pair sexual interests, progesterone levels were associated with relatively greater in-pair sexual interests. Both hormones specifically predicted in-pair sexual desire, estradiol negatively and progesterone positively. These findings have implications for understanding the function of women's extended sexuality — their sexual proceptivity and receptivity outside the fertile phase, especially during the luteal phase.     

Effects of partner conception risk phase on male perception of dominance in faces

Several studies have suggested that women may prefer to engage in extra-pair copulations with males who appear dominant and to do so near ovulation. While there is some evidence that males are more jealous of dominant rivals and more proprietary when their partners are near ovulation, there is none that suggests the existence of counterstrategic perceptual shifts that mirror those seen in women. We provide such evidence here. Composites of male faces that were either high or low in rated dominance were presented to male participants who provided ratings of dominance. A three-way interaction between stimulus-face dominance, partner conception risk phase, and partner oral contraceptive use was found; men whose partners did not use an oral contraceptive and were in the high conception risk phase of their cycle displayed increased dominance ratings of high-dominance male faces. We conclude that males have evolved counterstrategies to deal with female infidelity that include an overattribution of dominance to those rivals most likely to present a threat at times when that threat is greatest. This overattribution is likely to lead to increases in jealousy and mate-retention behaviors.     

Ovulatory Shifts in Women��s Attractions to Primary Partners and Other Men: Further Evidence of the Importance of Primary Partner Sexual Attractiveness

Previous research has documented shifts in women’s attractions to their romantic partner and to men other than their partner across the ovulation cycle, contingent on the degree to which her partner displays hypothesized indicators of high-fitness genes. The current study set out to replicate and extend this finding. Forty-one couples in which the woman was naturally cycling participated. Female partners reported their feelings of in-pair attraction and extra-pair attraction on two occasions, once on a low-fertility day of the cycle and once on a high-fertility day of the cycle just prior to ovulation. Ovulation was confirmed using luteinizing hormone tests. We collected two measures of male partner sexual attractiveness. First, the women in the study rated their partner’s sexual attractiveness. Second, we photographed the partners and had the photos independently rated for attractiveness. Shifts in women’s in-pair attractions across the cycle were significantly moderated by women’s ratings of partner sexual attractiveness, such that the less sexually attractive women rated their partner, the less in-pair attraction they reported at high fertility compared with low fertility (partial r = .37, p _dir = .01). Shifts in women’s extra-pair attractions across the cycle were significantly moderated by third-party ratings of partner attractiveness, such that the less attractive the partner was, the more extra-pair attraction women reported at high relative to low fertility (partial r = −.33, p _dir = .03). In line with previous findings, we found support for the hypothesis that the degree to which a woman’s romantic partner displays indicators of high-fitness genes affects women’s attractions to their own partner and other men at high fertility.     

Women's pupillary responses to sexually significant others during the hormonal cycle

Women's sexual preferences can change over the hormonal cycle, as several studies, based on responses to questionnaires, diaries, and ratings of photographs, have indicated increased sexual interests around the time of ovulation. However, fewer studies have measured changes in attention or interest to sexually significant stimuli in terms of physiological responses that are not under voluntary control and measure sexual interest indirectly (i.e., without mention of sexual feelings or activities). In the present study, we indexed changes in sexual interest in terms of changes in the eye pupil's size. Pupillary diameter is known to have a proportional relation to the observer's level of interest and attention to a visual stimulus as well as to physical pleasure. Fourteen women (7 being "pill" users) viewed photos on a computer screen while their pupil diameters were recorded using an infrared eye-tracking device. Three measures were taken for each participant during three time windows that estimated the ovulatory, luteal, and menstrual phase of the cycle. We found an increase in mean pupil diameter for sexually significant stimuli during the fertile phase and this pupillary change was also specific to pictures of the participants' actual sexual partners. Moreover, this effect was only seen for women who did not use oral contraceptives. These findings confirm that women's attention for sexually significant stimuli is higher during their fertile phase of the menstrual cycle, and that changes in sexual interest are implicitly measurable using pupillometry.     

Ovulatory shifts in human female ornamentation: near ovulation, women dress to impress

Humans differ from many other primates in the apparent absence of obvious advertisements of fertility within the ovulatory cycle. However, recent studies demonstrate increases in women's sexual motivation near ovulation, raising the question of whether human ovulation could be marked by observable changes in overt behavior. Using a sample of 30 partnered women photographed at high and low fertility cycle phases, we show that readily-observable behaviors - self-grooming and ornamentation through attractive choice of dress - increase during the fertile phase of the ovulatory cycle. At above-chance levels, 42 judges selected photographs of women in their fertile (59.5%) rather than luteal phase (40.5%) as "trying to look more attractive." Moreover, the closer women were to ovulation when photographed in the fertile window, the more frequently their fertile photograph was chosen. Although an emerging literature indicates a variety of changes in women across the cycle, the ornamentation effect is striking in both its magnitude and its status as an overt behavioral difference that can be easily observed by others. It may help explain the previously documented finding that men's mate retention efforts increase as their partners approach ovulation.     

Human oestrus

For several decades, scholars of human sexuality have almost uniformly assumed that women evolutionarily lost oestrus--a phase of female sexuality occurring near ovulation and distinct from other phases of the ovarian cycle in terms of female sexual motivations and attractivity. In fact, we argue, this long-standing assumption is wrong. We review evidence that women's fertile-phase sexuality differs in a variety of ways from their sexuality during infertile phases of their cycles. In particular, when fertile in their cycles, women are particularly sexually attracted to a variety of features that likely are (or, ancestrally, were) indicators of genetic quality. As women's fertile-phase sexuality shares with other vertebrate females' fertile-phase sexuality a variety of functional and physiological features, we propose that the term oestrus appropriately applies to this phase in women. We discuss the function of women's non-fertile or extended sexuality and, based on empirical findings, suggest ways that fertile-phase sexuality in women has been shaped to partly function in the context of extra-pair mating. Men are particularly attracted to some features of fertile-phase women, but probably based on by-products of physiological changes males have been selected to detect, not because women signal their cycle-based fertility status.     

Intersexual conflict across women's ovulatory cycle

A growing literature shows that the features women find particularly attractive in men vary across the ovulatory cycle. Women furthermore appear to more frequently report attraction to men other than primary partners when they are fertile in their cycle than in the infertile luteal phase. Previous studies have shown that men are more vigilant of or attentive to their primary partners during the fertile phase compared to the luteal phase. This study had several aims: First, to replicate and extend previous findings concerning men's vigilance of partners using male as well as female reports of men's behavior; second, to examine changes in women's behavior toward partners across the cycle; third, to examine ways in which women resist men's attempts to mate-guard across the cycle. Results indicate that (a) men are particularly self-assertive toward partners when their partners are fertile; (b) similarly, women are especially self-assertive toward partners when they are fertile; (c) women report engaging in more behaviors that resist male vigilance and mate guarding when they are fertile, especially in ways that are unobservable to male partners; and (d) these effects are especially strong when women themselves report greater attraction to men other than partners when they are fertile, compared to the luteal phase.     

Within-cycle fluctuations in progesterone negatively predict changes in both in-pair and extra-pair desire among partnered women

Grebe et al. (2016) argued that women's sexual interest in their own partners may be under different hormonal regulation than their sexual desire for other men. They measured partnered women's salivary hormones and reports of attraction to different categories of men at two time points separated by one week. Change in progesterone positively predicted change in women's desire for their own partners, whereas change in estradiol was a negative predictor. These results are opposite to those we previously reported for the hormonal prediction of general sexual desire in a study that employed frequent hormone sampling across multiple menstrual cycles (Roney and Simmons, 2013). Here, to test replication of the Grebe et al. findings, we assessed hormonal predictors of targeted in-pair and extra-pair desire among the subset of the sample from our 2013 paper who reported being in romantic relationships. Contrary to Grebe et al. (2016), we found that within-cycle fluctuations in progesterone were negatively correlated with changes in women's desire for both their own partners and other men. In addition, both in-pair and extra-pair desire were elevated within the fertile window and lowest during the luteal phase. Our findings contradict the idea that partner-specific desire has a unique form of hormonal regulation, and instead support a general elevation of sexual motivation associated with hormonal indices of fecundity. Discussion focuses on possible reasons for the discrepancies in findings between our study and that of Grebe et al. (2016), and on the evolved functions of women's sexual motivation.     

Changes in Sleep Time and Sleep Quality across the Ovulatory Cycle as a Function of Fertility and Partner Attractiveness

Research suggests that near ovulation women tend to consume fewer calories and engage in more physical activity; they are judged to be more attractive, express greater preferences for masculine and symmetrical men, and experience increases in sexual desire for men other than their primary partners. Some of these cycle phase shifts are moderated by partner attractiveness and interpreted as strategic responses to women''s current reproductive context. The present study investigated changes in sleep across the ovulatory cycle, based on the hypothesis that changes in sleep may reflect ancestral strategic shifts of time and energy toward reproductive activities. Participants completed a 32-day daily diary in which they recorded their sleep time and quality for each day, yielding over 1,000 observations of sleep time and quality. Results indicated that, when the probability of conception was high, women partnered with less attractive men slept more, while women with more attractive partners slept less.     

Women's interest in visual sexual stimuli varies with menstrual cycle phase at first exposure and predicts later interest

This study investigated whether women's interest in visual sexual stimuli varied with their hormonal state. Viewing times of 30 women, 15 normal cycling (NC) and 15 oral contracepting (OC), to sexually explicit photos were measured at three different times. NC women were tested during their menstrual, periovulatory, and luteal phases, and OC women were tested at equivalent temporal intervals. Subjects viewed stimuli as long as desired, thus viewing time measured subject interest. Subjective ratings of stimulus sexual attractiveness were obtained on each test. There was no overall relationship between menstrual cycle phase and viewing time. However the participant's menstrual cycle phase during first exposure to sexual stimuli predicted subsequent interest in sexual stimuli during the next two tests. NC women who first viewed stimuli during their periovulatory phase looked longer at the sexual stimuli across all sessions than did women first tested in their luteal phase. OC women first exposed to the sexual stimuli during menstruation looked longer at the stimuli across all sessions than did OC women first exposed at other test phases. Neither current test phase nor initial cycle phase influenced subjective ratings. Women had increased interest in sexual stimuli across all sessions if first exposed to sexual stimuli when endogenous estrogens were most likely highest. These data suggest that women's interest in visual sexual stimuli is modulated by hormones such that the hormonal condition at first exposure possibly determines the stimuli's emotional valence, markedly affecting subsequent interest in sexual stimuli.     

Menstrual cycle phase affects discrimination of infant cuteness

Recent studies have shown that women are more sensitive than men to subtle cuteness differences in infant faces. It has been suggested that raised levels in estradiol and progesterone may be responsible for this advantage. We compared young women's sensitivity to computer-manipulated baby faces varying in cuteness. Thirty-six women were tested once during ovulation and once during the luteal phase of their menstrual cycle. In a two alternative forced-choice experiment, participants chose the baby which they thought was cuter (Task 1), younger (Task 2), or the baby that they would prefer to babysit (Task 3). Saliva samples to assess levels of estradiol, progesterone and testosterone were collected at each test session. During ovulation, women were more likely to choose the cuter baby than during the luteal phase, in all three tasks. These results suggest that cuteness discrimination may be driven by cyclic hormonal shifts. However none of the measured hormones were related to increased cuteness sensitivity. We speculate that other hormones than the ones measured here might be responsible for the increased sensitivity to subtle cuteness differences during ovulation.     

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.