Reproductive strategy, sexual development and attraction to facial characteristics

Sexual reproduction strategies vary both between and within species in the level of investment in offspring. Life-history theories suggest that the rate of sexual maturation is critically linked to reproductive strategy, with high investment being associated with few offspring and delayed maturation. For humans, age of puberty and age of first sex are two developmental milestones that have been associated with reproductive strategies. Stress during early development can retard or accelerate sexual maturation and reproduction. Early age of menarche is associated with absence of younger siblings, absence of a father figure during early life and increased weight. Father absence during early life is also associated with early marriage, pregnancy and divorce. Choice of partner characteristics is critical to successful implementation of sexual strategies. It has been suggested that sexually dimorphic traits (including those evident in the face) signal high-quality immune function and reproductive status. Masculinity in males has also been associated with low investment in mate and offspring. Thus, women's reproductive strategy should be matched to the probability of male investment, hence to male masculinity. Our review leads us to predict associations between the rate of sexual maturation and adult preferences for facial characteristics (enhanced sexual dimorphism and attractiveness). We find for men, engaging in sex at an early age is related to an increased preference for feminized female faces. Similarly, for women, the earlier the age of first sex the greater the preference for masculinity in opposite-sex faces. When we controlled sexual dimorphism in male faces, the speed of sexual development in women was not associated with differences in preference for male facial attractiveness. These developmental influences on partner choice were not mediated by self-rated attractiveness or parental relationships. We conclude that individuals assort in preferences based on the rapidity of their sexual development. Fast developing individuals prefer opposite-sex partners with an increased level of sexually dimorphic facial characteristics.     

Intergenerational context discontinuity affects the onset of puberty

The assumption that the onset of puberty is a context-sensitive marker of a reproductive strategy is tested by comparing parental and filial childhood context and somatic development in West and East Germany. Sixty-eight mother-daughter dyads and 35 father-son dyads were taken from two samples of families from Osnabrück in West Germany and Halle in East Germany. According to the observed context discontinuity between the generations in the male dyads, linear regression models show that no indicator of male sexual maturation was influenced significantly by the somatic development of the father. Instead of an inherited timing of maturation, antecedent distal factors like socioeconomic childhood context variables and critical life events lead to an acceleration of male sexual maturation. Finally we test the effect of two different conditions of childhood context continuity on daughter’s age at menarche with maternal age at menarche controlled. Linear regression models show that mother’s age at menarche predicts daughter’s age at menarche only under the condition of contextual continuity between generations, which was the case in the West German sample only. In East Germany, where mother’s age at menarche had no significant effect, the amount of variance explained by childhood context variables was almost the same. These results indicate the context sensitivity of somatic development which seems to follow an evolutionary rationale.     

Facial attractiveness ratings from video-clips and static images tell the same story

Most of what we know about what makes a face attractive and why we have the preferences we do is based on attractiveness ratings of static images of faces, usually photographs. However, several reports that such ratings fail to correlate significantly with ratings made to dynamic video clips, which provide richer samples of appearance, challenge the validity of this literature. Here, we tested the validity of attractiveness ratings made to static images, using a substantial sample of male faces. We found that these ratings agreed very strongly with ratings made to videos of these men, despite the presence of much more information in the videos (multiple views, neutral and smiling expressions and speech-related movements). Not surprisingly, given this high agreement, the components of video-attractiveness were also very similar to those reported previously for static-attractiveness. Specifically, averageness, symmetry and masculinity were all significant components of attractiveness rated from videos. Finally, regression analyses yielded very similar effects of attractiveness on success in obtaining sexual partners, whether attractiveness was rated from videos or static images. These results validate the widespread use of attractiveness ratings made to static images in evolutionary and social psychological research. We speculate that this validity may stem from our tendency to make rapid and robust judgements of attractiveness.     

Early stress predicts age at menarche and first birth,adult attachment,and expected lifespan

Life history theory suggests that in risky and uncertain environments the optimal reproductive strategy is to reproduce early in order to maximize the probability of leaving any descendants at all. The fact that early menarche facilitates early reproduction provides an adaptationist rationale for our first two hypotheses: that women who experience more risky and uncertain environments early in life would have (1) earlier menarche and (2) earlier first births than women who experience less stress at an early age. Attachment theory and research provide the rationale for our second two hypotheses: that the subjective early experience of risky and uncertain environments (insecurity) is (3) part of an evolved mechanism for entraining alternative reproductive strategies contingent on environmental risk and uncertainty and (4) reflected in expected lifespan. Evidence from our pilot study of 100 women attending antenatal clinics at a large metropolitan hospital is consistent with all four hypotheses: Women reporting more troubled family relations early in life had earlier menarche, earlier first birth, were more likely to identify with insecure adult attachment styles, and expected shorter lifespans. Multivariate analyses show that early stress directly affected age at menarche and first birth, affected adult attachment in interaction with expected lifespan, but had no effect on expected lifespan, where its original effect was taken over by interactions between age at menarche and adult attachment as well as age at first birth and adult attachment. We discuss our results in terms of the need to combine evolutionary and developmental perspectives and the relation between early stress in general and father absence in particular. This work was supported by The University of Melbourne Department of Obstetrics and Gynaecology. James S. Chisholm is Professor in the School of Anatomy and Human Biology at the University of Western Australia. He is an anthropologist whose interests lie in the fields of human behavioral biology, evolutionary ecology, life history theory, and parental investment theory, where he focuses on infant social-emotional development, the development of reproductive strategies, and the integration of evolutionary, developmental, and cultural psychology and public health. Julie A. Quinlivan is Associate Professor in Obstetrics and Gynaecology at the University of Melbourne and Head of the Maternity Care Program at the Royal Women’s Hospital, Melbourne. Her interests are teenage pregnancy, domestic violence, child abuse prevention, and high-risk pregnancy. Rodney W. Petersen is Senior Lecturer in Obstetrics and Gynaecology at the University of Melbourne and Senior Consultant in Obstetrics and Gynaecology at the Royal Women’s Hospital and Sunshine Hospital in Melbourne. His interests are in psychosocial aspects of women’s health and cancer. David A. Coall is a Ph.D. student in the School of Anatomy and Human Biology at the University of Western Australia. His main interest lies in the application of evolutionary theory within an epidemiological framework. He is currently working on the synthesis of life history theory, parental investment theory, and parent-offspring conflict theory in exploring factors that influence variation in human birth weight and placental weight.     

Sexual Conflict and Gender Gap Effects: Associations between Social Context and Sex on Rated Attractiveness and Economic Status

Human mate choice research often concerns sex differences in the importance of traits such as physical attractiveness and social status. A growing number of studies indicate that cues to social context, including other people who appear in stimulus photographs, can alter that individual’s attractiveness. Fewer studies, however, consider judgements of traits other than physical attractiveness, such as wealth. Here we manipulate the presence/absence of other people in photographs of target models, and test the effects on judgments of both attractiveness and earnings (a proxy for status). Participants (N = 2044) rated either male or female models for either physical attractiveness or social/economic status when presented alone, with same sex others or with opposite sex others. We collectively refer to this manipulation as ‘social context’. Male and female models received similar responses for physical attractiveness, but social context affected ratings of status differently for women and men. Males presented alongside other men received the highest status ratings while females presented alone were given the highest status ratings. Further, the status of females presented alongside a male was constrained by the rated status of that male. Our results suggests that high status may not directly lead to high attractiveness in men, but that status is more readily attributed to men than to women. This divide in status between the sexes is very clear when men and women are presented together, possibly reflecting one underlying mechanism of the modern day gender gap and sexist attitudes to women’s economic participation. This adds complexity to our understanding of the relationship between attractiveness, status, and sex in the light of parental investment theory, sexual conflict and economic theory.     

Father absence, parental care, and female reproductive development

This study examines female reproductive development from an evolutionary life history perspective. Retrospective data are for 10,847 U.S. women. Results indicate that timing of parental separation is associated with reproductive development and is not confounded with socioeconomic variables or phenotypic correlations with mothers' reproductive behavior. Divorce/separation between birth and 5 years predicted early menarche, first sexual intercourse, first pregnancy, and shorter duration of first marriage. Separation in adolescence was the strongest predictor of number of sex partners. Multiple changes in childhood caretaking environment were associated with early menarche, first sex, first pregnancy, greater number of sex partners, and shorter duration of marriage. Living with either the father or mother after separation had similar effect on reproductive development. Living with a stepfather showed a weak, but significant, association with reproductive development, however, duration of stepfather exposure was not a significant predictor of development. Difference in amount and quality of direct parental care (vs. indirect parental investment) in two- and single-parent households may be the primary factor linking family environment to reproductive development.     

Increase in Male Reproductive Success and Female Reproductive Investment in Invasive Populations of the Harlequin Ladybird Harmonia axyridis

Reproductive strategy affects population dynamics and genetic parameters that can, in turn, affect evolutionary processes during the course of biological invasion. Life-history traits associated with reproductive strategy are therefore potentially good candidates for rapid evolutionary shifts during invasions. In a series of mating trials, we examined mixed groups of four males from invasive and native populations of the harlequin ladybird Harmonia axyridis mating freely during 48 hours with one female of either type. We recorded the identity of the first male to copulate and after the 48 h-period, we examined female fecundity and share of paternity, using molecular markers. We found that invasive populations have a different profile of male and female reproductive output. Males from invasive populations are more likely to mate first and gain a higher proportion of offspring with both invasive and native females. Females from invasive populations reproduce sooner, lay more eggs, and have offspring sired by a larger number of fathers than females from native populations. We found no evidence of direct inbreeding avoidance behaviour in both invasive and native females. This study highlights the importance of investigating evolutionary changes in reproductive strategy and associated traits during biological invasions.     

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.     

A relationship between attractiveness and performance in professional cyclists

Females often prefer to mate with high quality males, and one aspect of quality is physical performance. Although a preference for physically fitter males is therefore predicted, the relationship between attractiveness and performance has rarely been quantified. Here, I test for such a relationship in humans and ask whether variation in (endurance) performance is associated with variation in facial attractiveness within elite professional cyclists that finished the 2012 Tour de France. I show that riders that performed better were more attractive, and that this preference was strongest in women not using a hormonal contraceptive. Thereby, I show that, within this preselected but relatively homogeneous sample of the male population, facial attractiveness signals endurance performance. Provided that there is a relationship between performance-mediated attractiveness and reproductive success, this suggests that human endurance capacity has been subject to sexual selection in our evolutionary past.     

Effects of putative male pheromones on female ratings of male attractiveness: influence of oral contraceptives and the menstrual cycle

Previous research has revealed that natural and synthetic pheromones can enhance ratings of opposite sex attractiveness. The present study investigated the effects of exposure to male axillary secretions on female ratings of the sexual attractiveness of male stimuli. Thirty-two female undergraduates, half of whom were contraceptive pill users, rated male vignette characters and photographs of male faces on aspects of attractiveness. On two separate study days, corresponding to different phases of their menstrual cycle, stimuli were presented while exposed to male axillary pheromones and under a control condition (no pheromone). The order of testing was balanced with respect to pheromone/control condition and menstrual cycle phase. Pheromone exposure resulted in significantly higher attractiveness ratings of vignette characters and faces. Use of the contraceptive pill or menstrual cycle phase had equivocal effects on some vignette items and neither had any influence on female ratings of male facial attractiveness. The results of this study suggest that exposure to natural male axillary pheromones can significantly enhance female perceptions of various aspects of male attractiveness.     

Hairstyle as an adaptive means of displaying phenotypic quality

Although facial features that are considered beautiful have been investigated across cultures using the framework of sexual selection theory, the effects of head hair on esthetic evaluations have rarely been examined from an evolutionary perspective. In the present study the effects of six hair-styles (short, medium-length, long, disheveled, knot [hair bun], unkempt) on female facial attractiveness were examined in four dimensions (femininity, youth, health, sexiness) relative to faces without visible head hair (“basic face”). Three evolutionary hypotheses were tested (covering hypothesis, healthy mate theory, and good genes model); only the good genes model was supported by our data. According to this theory, individuals who can afford the high costs of long hair are those who have good phenotypic and genetic quality. In accordance with this hypothesis, we found that only long and medium-length hair had a significant positive effect on ratings of women’s attractiveness; the other hairstyles did not influence the evaluation of their physical beauty. Furthermore, these two hairstyles caused a much larger change in the dimension of health than in the rest of the dimensions. Finally, male raters considered the longer-haired female subjects’ health status better, especially if the subjects were less attractive women. The possible relationships between facial attractiveness and hair are discussed, and alternative explanations are presented.     

Human body odour, symmetry and attractiveness.

Several studies have found body and facial symmetry as well as attractiveness to be human mate choice criteria. These characteristics are presumed to signal developmental stability. Human body odour has been shown to influence female mate choice depending on the immune system, but the question of whether smell could signal general mate quality, as do other cues, was not addressed in previous studies. We compared ratings of body odour, attractiveness, and measurements of facial and body asymmetry of 16 male and 19 female subjects. Subjects wore a T-shirt for three consecutive nights under controlled conditions. Opposite-sex raters judged the odour of the T-shirts and another group evaluated portraits of the subjects for attractiveness. We measured seven bilateral traits of the subject's body to assess body asymmetry. Facial asymmetry was examined by distance measurements of portrait photographs. The results showed a significant positive correlation between facial attractiveness and sexiness of body odour for female subjects. We found positive relationships between body odour and attractiveness and negative ones between smell and body asymmetry for males only if female odour raters were in the most fertile phase of their menstrual cycle. The outcomes are discussed in the light of different male and female reproductive strategies.     

Why father absence might precipitate early menarche: The role of polygyny

An evolutionary model of conditional reproductive strategies argues that girls whose fathers are absent or make little parental investment experience early puberty. However, such a conditional strategy cannot be adaptive unless the absence of the girl's father at the microlevel is predictive of some recurrent feature of the macrosocial system and early puberty is advantageous in the system. I argue that father absence is indicative of the degree of polygyny (simultaneous and serial) in society. Polygyny of both kinds creates a shortage of women in reproductive age, and thus, early puberty will be advantageous. Available comparative data indicate that the degree of polygyny is associated with a decrease in the mean age of menarche across societies, as is the divorce rate a presumptive index of serial polygyny, in strictly monogamous societies.     

Explanatory Models of Female Pubertal Timing: Discordances Between Cultural Models of Maturation and the Recollection and Interpretation of Personal Developmental Experiences

Given the ambiguity surrounding the source of the continuing trend toward earlier menarche observed in Westernized nations, several competing explanatory models have emerged regarding variation in pubertal timing. While a biomedical model proposes that predominantly constitutional characteristics shape the maturation timetable, an alternative framework derived from Life History Theory (LHT) evolutionary principles emphasizes the influence of psychosocial factors on development. Working with a sample of women 19?C25?years of age (N?=?103) drawn from two Southeastern U.S. colleges, we combined cultural consensus analysis with retrospective self-report regarding childhood stress and menarcheal timing to investigate whether reported developmental experiences align with cultural models regarding factors that should drive pubertal timing. Results suggest a robust cultural model consistent with a biomedical framework concentrating principally on constitutional characteristics. However, participants?? personal developmental recollections support an association between higher childhood stress and earlier menarche. These findings support LHT predictions that early reproductive maturation is an evolutionary adaptive response to chronic childhood stress as well as clarify the extent to which cultural models of factors contributing to puberty concord with developmental experiences.     

Human grief: Is its intensity related to the reproductive value of the deceased?

Thurstone's method of comparative judgement was used to measured the intensity of grief that parents of high-, moderate-, and low-reproductive value were expected to experience at the death of male and female children of different ages. The results were correlated with reproductive values for male and female British Columbians and for !Kung Bushwomen. Grief ratings were more highly correlated with reproductive value than with age and more highly correlated with reproductive values of !King Bushwomen than with those of British Columbians. The correlations were higher for male- than for female-stimulus children. The correlations of female ratings with reproductive value were higher than male ratings with reproductive value, although not as high as expected. However, the correlation between grief ratings and reproductive value did not increase as the reproductive value of the raters declined.     

Evolution of floral dimorphism in a cleistogamous annual, <Emphasis Type="Italic">Impatiens noli-tangere</Emphasis> L. occurring under different environmental conditions

The resource allocation models for the evolution of a mixed production of cleistogamous and chasmogamous flowers were tested in four natural populations of Impatiens noli-tangere L. When we incorporated the estimated parameter values into our basic model, under the assumption that the environmental conditions did not change through the reproductive season, the model predicted the obligate chasmogamy in all four populations. However, including the fluctuation of the fertility of chasmogamous flowers in the model, the seasonal switching of chasmogamous and cleistogamous flower production becomes an evolutionary stable strategy, which explains the reproductive behavior of the individuals in two populations. Only after the relationship between the geitonogamy of chasmogamous flowers and the number of chasmogamous flower production was considered in the model did the simultaneous production of both of the dimorphic flowers become an evolutionary stable strategy. Further, the complete model clarifies the reproductive behavior of the individuals in the other two populations.     

Facial attractiveness and fertility in populations with low levels of modern birth control

Evolutionary models of human mate choice generally assume that physical attractiveness has evolved through sexual selection, i.e., it has been associated with higher mating opportunities and subsequent reproductive success across our evolutionary history. Here we investigate whether facial attractiveness is related to fertility in order to understand the extent to which selection can operate on attractive traits in modern populations. We used data from two populations where the prevalence of modern birth control methods is low and thus unlikely to disconnect mating opportunities from reproductive success: men and women from contemporary rural Senegal and men from the West Point Military Academy in the USA who graduated in 1950. We found that facial attractiveness negatively predicts age-specific reproduction in both sexes in Senegal and is independent from lifetime reproductive success in men from the USA. Overall, the results suggest that facial attractiveness is not under positive selection and raise questions about methodological approaches currently used to assess attractiveness.     

Genetic diversity revealed in human faces

From an evolutionary perspective, human facial attractiveness is proposed to signal mate quality. Using a novel approach to the study of the genetic basis of human preferences for facial features, we investigated whether attractiveness signals mate quality in terms of genetic diversity. Genetic diversity in general has been linked to fitness and reproductive success, and genetic diversity within the major histocompatibility complex (MHC) has been linked to immunocompetence and mate preferences. We asked whether any preference for genetic diversity is specific to MHC diversity or reflects a more general preference for overall genetic diversity. We photographed and genotyped 160 participants using microsatellite markers situated within and outside the MHC, and calculated two measures of genetic diversity: mean heterozygosity and standardized mean d(2). Our results suggest a special role for the MHC in female preferences for male faces. MHC heterozygosity positively predicted male attractiveness, and specifically facial averageness, with averageness mediating the MHC-attractiveness relationship. For females, standardized mean d(2) at non-MHC loci predicted facial symmetry. Thus, attractive facial characteristics appear to provide visual cues to genetic quality in both males and females, supporting the view that face preferences have been shaped by selection pressures to identify high-quality mates.     

Viewing time as a measure of sexual interest

Based on the hypothesis that unobtrusively measured viewing time reflects sexual interest, it was predicted that viewing times should correlate with ratings of sexual attractiveness and with phallometrically measured age and gender preferences. Four additional predictions were derived from the Symons (1979) evolutionary theory of human mate preferences: (1) male and female subjects should view slides of young adults of the opposite sex longest and adults of the same sex and prepubescent children of both sexes the shortest, (2) the correlation between viewing time and sexual attractiveness ratings should be higher among males than females (3) males should look at slides of pubescent females longer than females look at slides of pubescent males, and (4) males should look longer at adult females than females look at adult males. Two studies involving 24 heterosexual females and 58 heterosexual males provided statistically significant support for all predictions except the last one.     

Is facial width-to-height ratio reliably associated with social inferences?

Theoretical considerations and early empirical findings suggested facial width-to-height ratio (fWHR) may be relevant to person perception because it is associated with behavioral dispositions. More recent evidence failing to find fWHR-behavior links suggests that mismatch or byproduct hypotheses may be necessary to explain fWHR-based trait inferences; however, these explanations may not be needed because it is not clear that fWHR is reliably associated with trait inferences. To investigate the robustness of fWHR-inference links, we conducted secondary analyses of a cross-national dataset consisting of ratings by 11,481 participants across 11 world regions who judged 60 male and 60 female faces on one of 13 social traits (ns per trait range from 760 to 975). In preregistered analyses—and exploratory analyses of a subset of traits in the larger sample of 597 faces from which the 120 faces were drawn—we found mixed evidence for fWHR-based social judgments. In multilevel models, fWHR was not reliably linked to raters' judgments of male faces for any of the 13 trait-inferences but was negatively associated with ratings of female faces' dominance, trustworthiness, sociability, emotional stability, responsibility, confidence, attractiveness, and intelligence. In exploratory analyses of a subset of traits using the larger sample of faces, fWHR was associated positively with perceptions of meanness and aggressiveness in male but not female faces, negatively with attractiveness and dominance in female but not male faces, and negatively with trustworthiness in male but not female faces. We interpret these mixed findings to suggest that (1) fWHR-inference links are likely to be smaller and less reliable than expected from prior research; (2) fWHR may play a larger role in perceptions of female faces than would be predicted from the theory underpinning fWHR hypotheses; and (3) future research should more closely examine the extent to which robust fWHR-inferences reflect mismatch in the reliability of fWHR-behavior links between ancestral and modern environments versus byproducts of other person perception mechanisms.