Face perception is modulated by sexual preference

Face perception is mediated by a distributed neural system in the human brain . The response to faces is modulated by cognitive factors such as attention, visual imagery, and emotion ; however, the effects of gender and sexual orientation are currently unknown. We used fMRI to test whether subjects would respond more to their sexually preferred faces and predicted such modulation in the reward circuitry. Forty heterosexual and homosexual men and women viewed photographs of male and female faces and assessed facial attractiveness. Regardless of their gender and sexual orientation, all subjects similarly rated the attractiveness of both male and female faces. Within multiple, bilateral face-selective regions in the visual cortex, limbic system, and prefrontal cortex, similar patterns of activation were found in all subjects in response to both male and female faces. Consistent with our hypothesis, we found a significant interaction between stimulus gender and the sexual preference of the subject in the thalamus and medial orbitofrontal cortex, where heterosexual men and homosexual women responded more to female faces and heterosexual women and homosexual men responded more to male faces. Our findings suggest that sexual preference modulates face-evoked activation in the reward circuitry.     

Neural activation in women in response to masculinized male faces: mediation by hormones and psychosexual factors

Women's preference for masculine faces varies with hormonal state, sociosexuality, and relationship status, but the underlying mechanisms are poorly understood. We hypothesized that hormones and psychosexual factors (sociosexuality, sexual inhibition/excitation) mediate the perception and evaluation of male faces thereby influencing women's preferences. We used functional magnetic resonance imaging to measure brain activity in 12 women as they evaluated pictures of male faces (half 30% masculinized, half 30% feminized). Participants were heterosexual women, age 23–28 years, who were not in a committed relationship and not using hormonal contraception. Women were tested during both the follicular and luteal phase of their menstrual cycle. We found five brain regions related to face and risk processing that responded more to the masculinized than to the feminized faces, including the superior temporal gyrus, precentral gyrus, posterior cingulate cortex, inferior parietal lobule, and anterior cingulate cortex. Increased activation in the anterior cingulate cortex, specifically, may indicate that women perceive masculinized faces to be both more risky and more attractive. We did not see any areas that were more strongly activated by feminized faces. Levels of activation were influenced by hormonal and psychosexual factors. The patterns of hormonally and psychosexually mediated neural activation observed may offer insight into the cognitive processes underlying women's partner preferences.     

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 comparison of heterosexual and homosexual mating preferences in personal advertisements

Human heterosexual mating preferences have been shown to conform to predictions drawn from evolutionary theory, with men and women adopting broadly distinct strategies. Attempts to reconcile sexual selection theory with homosexual behavior have been less consistent, however, and have largely focussed on addressing two alternative perspectives: (i) that gay men and lesbians display phenotypic traits in common with opposite sex heterosexual individuals or (ii) that homosexual individuals display sex-typical, or exaggerated sex-typical phenotypes. Testing these hypotheses is complicated by sampling issues involved in the study of human sexual orientation, since obtaining standardized and comparable samples of heterosexual and non-heterosexual mating preferences is a prerequisite to analysis. Here we present a comparison of homosexual and heterosexual mating strategies in men and women using a sample of 1733 personal (‘lonely hearts’) adverts gathered from a single source. We used principal components analysis in order to expose underlying structure of the advertisements, and identify three components involving relative emphasis placed on resources, physical attractiveness and personality when offering or seeking mate characteristics. While homosexual individuals are shown to resemble their own-sex heterosexual counterparts in terms of emphasis placed on partner physical attractiveness relative to partner personality, no clear pattern emerges in other aspects of advertisement strategy. Nevertheless, there we find no evidence in support of the hypothesis that homosexual men and women are intrinsically opposite-sex typical in terms of mate preferences.     

Testing multiple hypotheses for the maintenance of male homosexual copulatory behaviour in flour beetles

Diverse animal groups exhibit homosexual interactions, yet the evolutionary maintenance of such behaviours remains enigmatic as they do not directly increase reproductive success by generating progeny. Here, we use Tribolium castaneum flour beetles, which exhibit frequent male homosexual copulations, to empirically test several hypotheses for the maintenance of such behaviours: (1) establishing social dominance; (2) practice for future heterosexual encounters; and (3) indirect sperm translocation. We found no evidence that Tribolium males use homosexual copulations either to establish dominance or to practice behaviours that increase their subsequent heterosexual reproductive performance. Our results provide limited support for the hypothesis of indirect sperm translocation: when males from two genetic strains mated with females immediately following a homosexual copulation, females produced progeny sired not only by the directly mating male, but also by that male’s homosexual partner. However, this phenomenon was detected in only 7% of homosexual pairs, and in each case such indirectly sired progeny accounted for < 0.5% of females’ total progeny. Direct observations indicated that mounting males often released spermatophores during homosexual copulations. These observations suggest that homosexual copulations may be a behavioural mechanism that allows males to expel older, potentially low‐quality sperm. Additional work is needed to test this new hypothesis, and to determine whether sperm release during homosexual copulations occurs in other groups.     

Homosexual incest avoidance among females in captive Japanese macaques

This paper reports a systematic pattern of homosexual incest avoidance among females in a captive group of Japanese macaques (Macaca fuscata) culled from the Arashiyama-West troop known for its high rates of female homosexuality. The study group included three matrilines and two generations. Between eight and 11 females were sexually active over four consecutive mating seasons, and all engaged in both heterosexual and homosexual activity. While all females performed homosexual acts with almost all possible non-kin partners, they systematically avoided homosexual interactions with their mother, daughters, and sisters. This pattern could not be explained either in terms of kin not being simultaneously in estrus, kin avoiding affiliative interactions in general, or non-kin utilizing the tension-reducing effect of estrus to affiliate exclusively with each other. In contrast to homosexual females, heterosexual pairs of relatives (brother-sister, mother-son) were sometimes incestuous. Assuming that female homosexuality expresses the reproductive strategy of females unconstrained by male influence, the present results point to the strong tendency of females to avoid incest and suggest that males are primarily responsible for the reported exceptions to incest avoidance.     

Courtship behaviour in Japanese macaques during heterosexual and homosexual consortships

Female Japanese macaques (Macaca fuscata) in the Arashiyama population near Kyoto, Japan, are unusual, in that they exhibit what many would consider to be male-typical sexual characteristics. Specifically, they mount other females within the context of temporary, but exclusive, sexual relationships (i.e., homosexual consortships) and they sometimes exhibit a preference for female sexual partners, even when given the choice of a sexually motivated male alternative. In this study, we examined whether female Japanese macaques also exhibited male-typical patterns of courtship behaviour during homosexual consortships. Data were collected on courtship behaviour during heterosexual and homosexual consortships in free-ranging Japanese macaques from the Arashiyama (Japan) population. We analyzed the occurrence of 12 different courtship behaviours during 3374 heterosexual inter-mount intervals and 1412 homosexual inter-mount intervals. Sex differences between heterosexually consorting males and females existed for only two of the 12 courtship behaviours we investigated: inclined-back presentations and sexual vocalizations. Dominant and subordinate homosexually consorting females were sex-typical in their expression of inclined-back presentations and sexual vocalizations. Consequently, facultative same-sex sexual partner preference, mounting and consortships do not co-occur with male-typical patterns of courtship behaviour in female Japanese macaques.     

Birth weight, sexual orientation and the sex of preceding siblings.

This study's first objective was to compare the mean birth weights of homosexual and heterosexual men and women. Its second objective was to investigate whether prior male and female fetuses have different effects on the birth weight of subsequent fetuses. The subjects were 3229 adult men and women (the probands), who weighed at least 2500 g at birth, and whose mothers knew the sex of the child (or fetus) for each pregnancy prior to the proband. Information on birth weight, maternal gravidity and other demographic variables was reported on questionnaires completed by the probands' mothers. The results confirmed earlier reports that boys with older brothers weigh less at birth than boys with older sisters, but they did not confirm reports that girls with older brothers weigh less than girls with older sisters. The results did not show across-the-board differences in the mean birth weights of homosexual versus heterosexual women or homosexual versus heterosexual men. However, the homosexual males with older brothers weighed about 170 g less at birth than the heterosexual males with older brothers. It is suggested that this pattern of results may reflect a maternal immune response to Y-linked minor histocompatibility antigens (H-Y antigens). According to this hypothesis, when the maternal immune response is mild, it produces only a slightly reduced birth weight, but when it is stronger, it produces a markedly reduced birth weight as well as an increased probability of homosexuality.     

Homosexual women have less grey matter in perirhinal cortex than heterosexual women

Is sexual orientation associated with structural differences in the brain? To address this question, 80 homosexual and heterosexual men and women (16 homosexual men and 15 homosexual women) underwent structural MRI. We used voxel-based morphometry to test for differences in grey matter concentration associated with gender and sexual orientation. Compared with heterosexual women, homosexual women displayed less grey matter bilaterally in the temporo-basal cortex, ventral cerebellum, and left ventral premotor cortex. The relative decrease in grey matter was most prominent in the left perirhinal cortex. The left perirhinal area also showed less grey matter in heterosexual men than in heterosexual women. Thus, in homosexual women, the perirhinal cortex grey matter displayed a more male-like structural pattern. This is in accordance with previous research that revealed signs of sex-atypical prenatal androgenization in homosexual women, but not in homosexual men. The relevance of the perirhinal area for high order multimodal (olfactory and visual) object, social, and sexual processing is discussed.     

Sexual behaviour of young and middle aged men in England and Wales.

Establishing patterns of sexual behaviour is essential in predicting the future spread of HIV. The sexual behaviour patterns of a randomly selected sample of 480 white men aged 15 to 49 were obtained by interview and analysed in relation to age, social class, and area of residence. Over half of the men had first had intercourse before the age of 18 and over three quarters had done so before the age of 20. Age at first intercourse tended to be lower in more recent birth cohorts and in social classes III, IV, and V. Men in earlier birth cohorts tended to have had fewer heterosexual partners, both regular and casual, than those born more recently, but there were no social class or regional differences in the number of partners. Eight of the 480 men (1.7%, 95% confidence interval 1.1 to 2.3%) stated that they had had homosexual intercourse, four of whom had had one casual homosexual partner. Only one man had had more than 25 homosexual partners. The number of men having had homosexual intercourse was lower than is widely thought. When possible, data were checked with national data or other surveys and thought to be accurate. The difficulties in collecting data of this nature are discussed.     

Testing Miller's theory of alleles preventing androgenization as an evolutionary explanation for the genetic predisposition for male homosexuality

The genetic background of male homosexuality presents an evolutionary paradox, since homosexuality could be considered a reproductive disadvantage. We tested E.M. Miller's (2000) balanced polymorphism explanation, which states that alleles partially preventing androgenization in male fetuses during pregnancy would be associated with a homosexual orientation. Having all the alleles produces homosexuality, while heterosexual carriers with only a few of these alleles instead have a reproductive advantage; that is, they have more traits, which, by controlling for excessive aggressiveness and psychopathy, make them more attractive mates. Pairs of brothers were used to test these assumptions. If homosexuality is due to having all the androgenization-preventing alleles, then heterosexual men with homosexual brothers are more likely to also have some of the these alleles compared to heterosexual men with heterosexual brothers. These two categories were compared on variables related both directly and indirectly to reproductive success, with heterosexual men with a homosexual brother hypothesized to have an advantage on the variables. However, no statistically significant findings in support of the theory were detected. The results were discussed together with alternative explanations.     

Evidence against a relationship between dermatoglyphic asymmetry and male sexual orientation

Hall and Kimura (1994) studied the relation between dermatoglyphic asymmetry and male sexual orientation in a sample of 66 homosexual and 182 heterosexual men. They found that more homosexual men possessed a leftward dermatoglyphic asymmetry than did heterosexual men. In this paper, we report a comparative study about the relationship between sexual orientation and dermatoglyphic characteristics, including 60 homosexual men, 76 heterosexual men, and 60 heterosexual women, recruited from the general population, and also from a gay-rights nongovernmental organization, in Salvador, Brazil. Ulnar loops were the most frequent dermatoglyphic pattern in all groups, followed by whorls, arches, and radial loops. A chi-square analysis comparing the frequencies of the patterns in the three groups only showed an excess of ulnar loops in women (p < 0.05) and arches in men (p < 0.01), but did not reveal significant differences between homosexuals and the other groups studied. There was no significant difference between gay and straight men on total ridge count. We found a preponderance of rightward asymmetry in homosexual and heterosexual men, as well as in heterosexual women. Our results do not agree with Hall and Kimura's data indicating that more gay men possessed the minority leftward asymmetry than did straight men. There was no significant difference in leftward asymmetry in the sample studied. The results reported in this paper do not support any relation between dermatoglyphic asymmetry and male sexual orientation, and, thus, any hypothesis concerning a biological intrauterine contribution to adult sexual orientation somehow associated with dermatoglyphic development.     

Disclosure of sexual preference to physicians by black lesbian and bisexual women.

Physicians'' ability to diagnose and treat health care problems, particularly those with a psychosocial component, is facilitated by accurate information concerning the life-styles of their patients. White lesbians have been shown to be generally reluctant to disclose sexual orientation to their physicians, but little, if anything, is known about black lesbians. Black women, self-identified as bisexuals (N = 65) and lesbians (N = 529), were asked whether they had disclosed their homosexual behavior to their physicians. In the sample, only a third of the women had. Previous sexual experiences, both heterosexual and homosexual, were also queried to illuminate patterns of gynecologic health risk factors. Nearly all of the women reported previous heterosexual experiences.     

Androsteroneetiocholanolone Ratios in Male Homosexuals

Analyses of 24-hour specimens of urine from healthy adult males for androsterone and etiocholanolone produced values which, when calculated as discriminant scores, discriminated between heterosexual and exclusively homosexual individuals. This confirms a previous study.No significant differences were found between heterosexuals and homosexuals in parental ages, secondary sex characteristics, genitalia, anthropometry, 17-ketosteroids, and 17-ketogenic steroids.A significant difference was found between the heterosexual group and homosexual group in the number of homosexual relatives in the immediate and extended families.     

Neural correlates of hate

In this work, we address an important but unexplored topic, namely the neural correlates of hate. In a block-design fMRI study, we scanned 17 normal human subjects while they viewed the face of a person they hated and also faces of acquaintances for whom they had neutral feelings. A hate score was obtained for the object of hate for each subject and this was used as a covariate in a between-subject random effects analysis. Viewing a hated face resulted in increased activity in the medial frontal gyrus, right putamen, bilaterally in premotor cortex, in the frontal pole and bilaterally in the medial insula. We also found three areas where activation correlated linearly with the declared level of hatred, the right insula, right premotor cortex and the right fronto-medial gyrus. One area of deactivation was found in the right superior frontal gyrus. The study thus shows that there is a unique pattern of activity in the brain in the context of hate. Though distinct from the pattern of activity that correlates with romantic love, this pattern nevertheless shares two areas with the latter, namely the putamen and the insula.     

Is male homosexuality maintained via kin selection?

We tested the kin selection hypothesis of male homosexuality using questionnaire data in a community sample of homosexual and heterosexual men. Homosexual men were no more likely than heterosexual men to channel resources toward family members. Indeed, heterosexual men tended to give more financial resources to siblings than homosexual men. Furthermore, homosexual men were somewhat more estranged from family members, especially from fathers and oldest siblings.     

Patterns of Brain Activation when Mothers View Their Own Child and Dog: An fMRI Study

Neural substrates underlying the human-pet relationship are largely unknown. We examined fMRI brain activation patterns as mothers viewed images of their own child and dog and an unfamiliar child and dog. There was a common network of brain regions involved in emotion, reward, affiliation, visual processing and social cognition when mothers viewed images of both their child and dog. Viewing images of their child resulted in brain activity in the midbrain (ventral tegmental area/substantia nigra involved in reward/affiliation), while a more posterior cortical brain activation pattern involving fusiform gyrus (visual processing of faces and social cognition) characterized a mother''s response to her dog. Mothers also rated images of their child and dog as eliciting similar levels of excitement (arousal) and pleasantness (valence), although the difference in the own vs. unfamiliar child comparison was larger than the own vs. unfamiliar dog comparison for arousal. Valence ratings of their dog were also positively correlated with ratings of the attachment to their dog. Although there are similarities in the perceived emotional experience and brain function associated with the mother-child and mother-dog bond, there are also key differences that may reflect variance in the evolutionary course and function of these relationships.     

Fecundity of Paternal and Maternal Non-Parental Female Relatives of Homosexual and Heterosexual Men

A variety of social, developmental, biological and genetic factors influence sexual orientation in males. Thus, several hypotheses have attempted to explain the sustenance of genetic factors that influence male homosexuality, despite decreased fecundity within the homosexuals. Kin selection, the existence of maternal effects and two forms of balancing selection, sexually antagonistic selection and overdominance, have been proposed as compensatory mechanisms for reduced homosexual fecundity. Here, we suggest that the empirical support for kin selection and maternal effects cannot account for the low universal frequency and stability of the distribution of homosexuals. To identify the responsible compensatory mechanism, we analyzed fecundity in 2,100 European female relatives, i.e., aunts and grandmothers, of either homosexual or heterosexual probands who were matched in terms of age, culture and sampling strategy. Female relatives were chosen to avoid the sampling bias of the fraternal birth order effect, which occurs when indirectly sampling mothers though their homosexual sons. We observed that the maternal aunts and grandmothers of homosexual probands were significantly more fecund compared with the maternal aunts and maternal grandmothers of the heterosexual probands. No difference in fecundity was observed in the paternal female lines (grandmothers or aunts) from either of the two proband groups. Moreover, due to the selective increase in maternal female fecundity, the total female fecundity was significantly higher in homosexual than heterosexual probands, thus compensating for the reduced fecundity of homosexuals. Altogether, these data support an X-linked multi-locus sexually antagonistic hypothesis rather than an autosomal multi-locus overdominance hypothesis.     

What Does Brain Response to Neutral Faces Tell Us about Major Depression? Evidence from Machine Learning and fMRI

Introduction

A considerable number of previous studies have shown abnormalities in the processing of emotional faces in major depression. Fewer studies, however, have focused specifically on abnormal processing of neutral faces despite evidence that depressed patients are slow and less accurate at recognizing neutral expressions in comparison with healthy controls. The current study aimed to investigate whether this misclassification described behaviourally for neutral faces also occurred when classifying patterns of brain activation to neutral faces for these patients.

Methods

Two independent depressed samples: (1) Nineteen medication-free patients with depression and 19 healthy volunteers and (2) Eighteen depressed individuals and 18 age and gender-ratio-matched healthy volunteers viewed emotional faces (sad/neutral; happy/neutral) during an fMRI experiment. We used a new pattern recognition framework: first, we trained the classifier to discriminate between two brain states (e.g. viewing happy faces vs. viewing neutral faces) using data only from healthy controls (HC). Second, we tested the classifier using patterns of brain activation of a patient and a healthy control for the same stimuli. Finally, we tested if the classifier’s predictions (predictive probabilities) for emotional and neutral face classification were different for healthy controls and depressed patients.

Results

Predictive probabilities to patterns of brain activation to neutral faces in both groups of patients were significantly lower in comparison to the healthy controls. This difference was specific to neutral faces. There were no significant differences in predictive probabilities to patterns of brain activation to sad faces (sample 1) and happy faces (samples 2) between depressed patients and healthy controls.

Conclusions

Our results suggest that the pattern of brain activation to neutral faces in depressed patients is not consistent with the pattern observed in healthy controls subject to the same stimuli. This difference in brain activation might underlie the behavioural misinterpretation of the neutral faces content by the depressed patients.     

Basic Instinct Undressed: Early Spatiotemporal Processing for Primary Sexual Characteristics

This study investigates the spatiotemporal dynamics associated with conscious and non-conscious processing of naked and dressed human bodies. To this effect, stimuli of naked men and women with visible primary sexual characteristics, as well as dressed bodies, were presented to 20 heterosexual male and female participants while acquiring high resolution EEG data. The stimuli were either consciously detectable (supraliminal presentations) or were rendered non-conscious through backward masking (subliminal presentations). The N1 event-related potential component was significantly enhanced in participants when they viewed naked compared to dressed bodies under supraliminal viewing conditions. More importantly, naked bodies of the opposite sex produced a significantly greater N1 component compared to dressed bodies during subliminal presentations, when participants were not aware of the stimulus presented. A source localization algorithm computed on the N1 showed that the response for naked bodies in the supraliminal viewing condition was stronger in body processing areas, primary visual areas and additional structures related to emotion processing. By contrast, in the subliminal viewing condition, only visual and body processing areas were found to be activated. These results suggest that naked bodies and primary sexual characteristics are processed early in time (i.e., <200 ms) and activate key brain structures even when they are not consciously detected. It appears that, similarly to what has been reported for emotional faces, sexual features benefit from automatic and rapid processing, most likely due to their high relevance for the individual and their importance for the species in terms of reproductive success.