David A Puts

Pennsylvania State University | Penn State · Department of Anthropology

As the earliest measure of social communication in rodents, ultrasonic vocalizations (USVs) in response to maternal separation are critical in preclinical research on neurodevelopmental disorders (NDDs). While sex differences in both USV production and behavioral outcomes are reported, many studies overlook sex as a biological variable in preclinic...

Fundamental frequency ( f o ) is the most perceptually salient vocal acoustic parameter, yet little is known about how its perceptual influence varies across societies. We examined how f o affects key social perceptions and how socioecological variables modulate these effects in 2,647 adult listeners sampled from 44 locations across 22 nations. Low...

People quickly and involuntarily form impressions of others based on their facial physical attributes, which can modulate critical social interactions. Skin pigmentation is one of the most variable and conspicuous facial traits among human populations. Empirical evidence suggests that these variations reflect ancestral ecological selective pressure...

Sexual attraction to males or females is perhaps the largest behavioral sex difference across animal species. Experiments in laboratory mammals show that prenatal androgens mediate this sex difference, but ethical considerations preclude such experimentation in humans. Multiple lines of converging correlational evidence are therefore needed to demo...

Vocalizations differ substantially between the sexes in many primates, and low-frequency male vocalizations may be favored by sexual selection because they intimidate rivals and/or attract mates. Sexual dimorphism in fundamental frequency may be more pronounced in species with more intense male mating competition and in those with large group size,...

Research on heterosexual mating has demonstrated that acoustic parameters (e.g., pitch) of men’s voices influence their attractiveness to women and appearance of status and formidability to other men. However, little is known about how men’s tendency to clearly articulate their speech influences these important social perceptions. In the current st...

Accumulating evidence suggests that the phenotypes of human males were shaped by contest competition, the mode of sexual selection in which mating opportunities are obtained by using force or threat of force to exclude same-sex competitors. Phylogenetic, paleontological, and archaeological data indicate a great antiquity for male–male violence in o...

Acoustic features of the voice, including fundamental and formant frequencies, communicate qualities of the vocalizer. The preferences of potential mates may have shaped these vocal parameters in both sexes over human evolution, contributing not only to large sexual dimorphisms but also to patterns of vocal modulation that leverage mate preferences...

Benenson et al. postulate that human females evolved unique survival adaptations to facilitate maternal and grandmaternal care. This hypothesis is consistent with the broader hypothesis that female phenotypes are more ecologically optimal, but further evidence is needed to make a compelling case that sex differences in self-protection are not prima...

Objective The dual-hormone hypothesis (DHH) posits that some effects of testosterone on human behavior and psychology related to status-seeking are moderated by cortisol, such that they are stronger when cortisol levels are low. In support of the DHH, studies have found that cortisol negatively moderated the relationship between testosterone and su...

Low perinatal androgens predict recalled childhood gender nonconformity in men Abstract The contributions of gonadal hormones to the development of human behavioral sex differences are subjects of intense scientific and social interest. Isolated GnRH deficiency (IGD) is a rare endocrine disorder that can reveal a possible role of early gonadal horm...

In laboratory animals, exposure to gonadal steroid hormones before and immediately after birth can exert permanent effects on many behaviors, particularly reproductive behaviors. The extent to which such effects occur in humans remains an open question, but several lines of evidence indicate that perinatal levels of both androgens and estrogens may...

The human voice is dynamic, and people modulate their voices across different social interactions. This article presents a review of the literature examining natural vocal modulation in social contexts relevant to human mating and intrasexual competition. Altering acoustic parameters during speech, particularly pitch, in response to mating and comp...

Individual variation in the age of pubertal onset is linked to physical and mental health, yet the factors underlying this variation are poorly understood. Life history theory predicts that individuals at higher risk of mortality due to extrinsic causes such as infectious disease should sexually mature and reproduce earlier, whereas those at lower...

Ovarian estrogens may influence the development of the human brain and behavior, but there are few opportunities to test this possibility. Isolated GnRH deficiency (IGD) is a rare endocrine disorder that could provide evidence for the role of estrogens in organizing sexually differentiated phenotypes: Unlike typical development, development in indi...

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Research on links between peoples’ personality traits and their voices has primarily focused on other peoples’ personality judgments about a target person based on a target person’s vocal characteristics, particularly voice pitch. However, it remains unclear whether individual differences in voices are linked to actual individual differences in per...

The contributions of gender socialization and direct hormonal action on the brain in the development of human behavioral sex differences are subjects of intense scientific and social interest. Prior research indicates masculinized behavioral patterns in individuals with high prenatal androgen exposure raised as girls, but complementary evidence reg...

Human voice pitch is highly sexually dimorphic and eminently quantifiable, making it an ideal phenotype for studying the influence of sexual selection. In both traditional and industrial populations, lower pitch in men predicts mating success, reproductive success, and social status and shapes social perceptions, especially those related to physica...

Circulating gonadal hormones have been linked to variation in the structure and function of the adult human brain, raising the question of how cognition is affected by sex hormones in adulthood. The impacts of progestogens and estrogens are of special interest due to the widespread use of hormone supplementation. Multiple studies have analyzed rela...

Sexual selection appears to have shaped the acoustic signals of diverse species, including humans. Deep, resonant vocalizations in particular may function in attracting mates and/or intimidating same-sex competitors. Evidence for these adaptive functions in human males derives predominantly from perception studies in which vocal acoustic parameters...

Men's voices may provide cues to overall condition; however, little research has assessed whether health status is reliably associated with perceivable voice parameters. In Study 1, we investigated whether listeners could classify voices belonging to men with either relatively lower or higher self-reported health. Participants rated voices for spea...

Acoustic signals function in intrasexual mating competition in a wide variety of species, including humans. The low voice pitch of human males has been proposed to represent an honest signal of formidability. Although voice pitch in men affects perceptions of size and dominance, it is relatively weakly associated with objective measures of formidab...

Research on links between peoples’ personality traits and their voices has primarily focused on other peoples’ personality judgments about a target person based on a target person’s vocal characteristics, particularly voice pitch. However, it remains unclear whether individual differences in voices are linked to actual individual differences in per...

Evidence suggests that psychosexuality in humans is modulated by both organizational effects of prenatal and peripubertal sex steroid hormones, and by activational effects of circulating hormones in adulthood. Experimental work in male rodents indicates that sensitivity to androgen-driven organization of sexual motivation decreases across the puber...

Experiments in male rodents demonstrate that sensitivity to the organizational effects of steroid hormones decreases across the pubertal window, with earlier androgen exposure leading to greater masculinization of the brain and behavior. Similarly, some research suggests the timing of peripubertal exposure to sex steroids influences aspects of huma...

Relatively little is known about the effects of endogenous and exogenous steroid hormones on ecologically relevant behavioral and cognitive phenotypes in women, such as emotion recognition, despite the widespread use of steroid hormone-altering hormonal contraceptives (HCs). Though some previous studies have examined the effect of HC use, estradiol...

Previous research suggests that binge drinking among young men serves as a “costly signal” to potential mates, such that the binge drinker is capable of bearing the harmful consequences of alcohol consumption. Here, we propose that binge drinking among young adults is conditionally dependent upon the signaler’s willingness to take risks, which is i...

Past work demonstrates that humans behave differently towards women across their menstrual cycles, even after exclusively visual exposure to women's faces. People may look at women's faces differently as a function of women's menstrual cycles. Analyses of participants' scanpaths (eye movement patterns) while they looked at women at different phases...

Pitch is the most perceptually salient acoustic property of the voice and influences perceptions of characteristics related to social power, such as dominance and leadership abilities. Voice pitch is also highly sexually differentiated; men vocalize approximately one octave below women. We consider the evolution of this sex difference, and how this...

Pitch is the most perceptually salient feature of the voice, yet it is approximately five standard deviations lower in men than in women, a degree of sexual dimorphism exceeding that of all extant nonhuman apes. Evidence from Western samples suggests that low-frequency vocalizations may have augmented male mating success ancestrally by intimidating...

Acoustic analysis and playback studies have greatly advanced our understanding of between-individual differences in nonverbal communication. Yet, researchers have only recently begun to investigate within-individual variation in the voice, particularly how people modulate key vocal parameters across various social contexts, with most of this resear...

This letter (to appear in Behavioral Ecology) reports alternative analyses of Shimoda et al. (2018 Behavioral Ecology https://doi.org/10.1093/beheco/arx124). These new analyses suggest that conception risk has similar effects on in- and extra-pair sexual desire, rather than the effect on extra-pair desire reported by Shimoda et al. (2018).

Research into the characteristics of attractive women’s voices has focused almost exclusively on associations with fundamental or formant frequencies. A recent study of a small sample of voices used a bottom-up approach to identify acoustic characteristics associated with women’s vocal attractiveness, finding that many acoustic characteristics othe...

Research into the characteristics of attractive women’s voices has focused almost exclusively on associations with fundamental or formant frequencies. A recent study of a small sample of voices used a bottom-up approach to identify acoustic characteristics associated with women’s vocal attractiveness, finding that many acoustic characteristics othe...

Women's mating psychology may have evolved to track reproductive conditions, including conception risk, across and between ovulatory cycles. Alternatively, within-woman correlations between mating psychology and ovarian hormones may be byproducts of between-women relationships. Here, we examined associations between steroid hormones and two facets...

Significance Facial masculinity has been considered a sexual ornament in humans, akin to peacock trains and stag antlers. Recently, studies have questioned the once-popular view that facial masculinity is a condition-dependent male ornament signaling immunocompetence (the immunocompetence handicap hypothesis). We sought to rigorously test these ide...

Are estrous mate preference shifts robust? This question is the subject of controversy within human evolutionary sciences. For nearly two decades, mate preference shifts across the ovulatory cycle were considered an important feature of human sexual selection, directing women’s attention towards mates with indicators of “good genes” in their fertil...

Human sexual orientation is an active topic of psychological research, with much attention focused on possible underlying endocrine factors. Promising results derive from studies of animal models, “natural experiments” involving human endocrine disorders, genetic variation in hormone receptor function, and biomarkers of hormonal signaling. Here, we...

Objectives Across the menstrual cycle, women exhibit fluctuations in psychosocial motivations. Some evidence suggests that near ovulation, women exhibit increased status concerns and behaviors that could be considered intrasexually competitive in nature. Women are sensitive to other women’s fertility, which may be useful for refining expectations a...

Prior work suggests that women’s preferences for sexually dimorphic traits, such as preferences for masculine facial and vocal characteristics, may be modulated by a multitude of factors related to reproductive potential including breastfeeding status. In the present study, we investigated women’s preferences for a highly sexually dimorphic vocal c...

A robust body of research has demonstrated shifts in women's sexual desire and arousal across the menstrual cycle, with heightened desire and arousal coincident with heightened probability of conception (POC), and it is likely that ovarian hormones modulate these shifts. However, studies in which women are exposed to audiovisual sexual stimuli (AVS...

Facial masculinity is thought to be a condition-dependent male ornament, reflecting immunocompetence in humans. To test this hypothesis, we calculated an objective measure of facial masculinity/femininity using three-dimensional images in a large sample (N = 1,233) of people of European ancestry. We show that facial masculinity is positively correl...

Are estrous mate preference shifts robust? This question is the subject of controversy within human evolutionary sciences. For nearly two decades, mate preference shifts across the ovulatory cycle were considered an important feature of human sexual selection, directing women's attention towards mates with indicators of "good genes" in their fertil...

Are estrous mate preference shifts robust? This question is the subject of controversy within human evolutionary sciences. For nearly two decades, mate preference shifts across the ovulatory cycle were considered an important feature of human sexual selection, directing women’s attention towards mates with indicators of “good genes” in their fertil...

Recent evidence suggests that in sexual selection on human males, intrasexual competition plays a larger role than female choice. In a sample of men (N = 164), we sought to provide further evidence on the effects of men's physical dominance and sexual attractiveness on mating success and hence in sexual selection. Objective measures and subjective...

Recent evidence suggests that in sexual selection on human males, intrasexual competition plays a larger role than female choice. In a sample of men (N = 164), we sought to provide further evidence on the effects of men’s physical dominance and sexual attractiveness on mating success and hence in sexual selection. Objective measures and subjective...

Previous research with hunter-gatherers has found that women perceive men with voices manipulated to be lower in pitch to be better hunters, and men perceive women with lower pitch to be better gatherers. Here, we test if actual voice pitch is associated with hunting and gathering reputations in men and women, respectively. We find that voice pitch...

Human genetic males are unlike rodent males in that neither the ability to convert testosterone to estrogen nor a functional estrogen receptor (ER) appears necessary for male-typical gender identity and sexuality, but a functional androgen receptor (AR) is required. Brain masculinization is probably mainly AR-mediated in human genetic males. ER bin...

Attractiveness plays a central role in human non-verbal communication and has been broadly examined in diverse subfields of contemporary psychology. Researchers have garnered compelling evidence in support of the evolutionary functions of physical attractiveness and its role in our daily lives, while at the same time, having largely ignored the sig...

Fluctuating asymmetry (FA), deviation from perfect bilateral symmetry, is thought to reflect an organism's relative inability to maintain stable morphological development in the face of environmental and genetic stressors. Previous research has documented negative relationships between FA and attractiveness judgments in humans, but scant research h...

Across three studies, we tested the hypothesis that women exhibit greater jealousy and mate guarding toward women who are in the high (vs. low) fertility phase of their cycle. Women who imagined their partner with a woman pictured at high fertility reported more jealousy than women who imagined their partner with a woman pictured at low fertility (...

The literature on human sexual selection has historically focused on the role of female mate choice, but cumulating experimental, correlational, and cross-cultural evidence suggests that male contest competition may have been more influential in shaping men's phenotypes. Cross-species comparison has shown similarities between humans and our closest...

Hormones orchestrate and coordinate human female sexual development, sexuality, and reproduction in relation to three types of phenotypic changes: life history transitions such as puberty and childbirth, responses to contextual factors such as caloric intake and stress, and cyclical patterns such as the ovulatory cycle. Here, we review the endocrin...

Polymorphisms in the dopamine D4 receptor (DRD4) have previously been shown to associate with a variety of human behavioral phenotypes, including ADHD pathology, alcohol and tobacco craving, financial risk-taking in males, and broader personality traits such as novelty seeking. Recent research has linked the presence of a 7-repeat (7R) allele in a...

In many primates, including humans, the vocalizations of males and females differ dramatically, with male vocalizations and vocal anatomy often seeming to exaggerate apparent body size. These traits may be favoured by sexual selection because low-frequency male vocalizations intimidate rivals and/or attract females, but this hypothesis has not been...

In their widely publicized paper, Joel et al. (1) make two empirical claims about sex differences in features of the human brain: (i) “…internal consistency [in individuals’ sex-differentiated brain features] is rare” (p. 15472) and (ii) the amount of overlap in sex-differentiated features of male and female brains “undermines any attempt to distin...

Over the past two decades, a large literature examining psychological changes across women's ovulatory cycles has accumulated, emphasizing comparisons between fertile and non-fertile phases of the cycle. While some studies have verified ovulation using luteinizing hormone (LH) tests, counting methods – assessments of conception probability based on...

Status competition among female mammals tends to intensify near ovulation. Females compete selectively, targeting females who most threaten their own likelihood of conception. The present study explored the extent to which regularly cycling women differentially compete with other women in a behavioral economic game as a function of both women’s fer...

For women, forming close, cooperative relationships with other women at once poses important opportunities and possible threats-including to mate retention. To maximize the benefits and minimize the costs of same-sex social relationships, we propose that women's mate guarding is functionally flexible and that women are sensitive to both interperson...

In this chapter, we explore how men’s phenotypes, including their psychologies, have been shaped by contest competition, a form of sexual selection involving the use of force or threat of force to exclude same-sex competitors from mates. We first evaluate the intensity of sexual selection in men, finding evidence of a positive and moderately strong...

The difficulty of inducing orgasm in women, its variability between women, and the lack of an obvious relationship with women’s reproductive success have led some researchers to conclude that female orgasm is a nonfunctional by-product of developmental pathways that women share with men. Other researchers have presented evidence that orgasm is an a...

In some laboratory rodents, males’ brains and behavior become less sensitive to the organizing effects of androgens across the time window surrounding puberty. Later puberty in human males has also been associated with less male-typical psychology and behavior, such as lower performance on mental rotation tasks and lower risk of substance abuse and...

The human face displays a wealth of information, including information about dominance and fecundity. Dominance and fecundity are also associated with lower concentrations of the stress hormone cortisol, suggesting that cortisol may negatively predict facial dominance and attractiveness. We digitally photographed 61 women's faces, had these images...

Across human societies and many nonhuman animals, males have greater interest in uncommitted sex (more unrestricted sociosexuality) than do females. Testosterone shows positive associations with male-typical sociosexual behavior in nonhuman animals. Yet, it remains unclear whether the human sex difference in sociosexual psychology (attitudes and de...

Women’s preferences for men’s masculinized faces and voices were assessed after women (n = 331) were primed with images of male-on-male aggression, male-on-female aggression, pathogens, and neutral scenes. Male-on-male aggression and pathogen primes were associated with increased preference for masculine traits, but the same effect emerged in the n...