The sex/gender conference succeeded in bringing together people “with different ideas and skills, different ways of thinking, that are actually transforming the field,” observed Carol Worthman, chair of Part 3 (“What’s at Stake?”). The earlier sessions (see Parts 1 and 2) provided us with a better sense of the complexities of sex/gender; we also learned some ways to usefully deconstruct – and form new ideas about – old questions. But there’s a lot at stake, Worthman continued. In the following session, speakers addressed the theme (“What counts as adequate function?”) from a variety of perspectives and from individual to macro levels of analysis. The question regarding adequate function is critical, Worthman reminded the audience, “because a lot of what is lurking in the background is frequently this question of ‘not good enough’ or ‘not a real person,’ both exogenously, in terms of how people are viewed, and endogenously, in terms of how they view themselves” by internalizing cultural norms. This suggests the importance of recognizing culture-mind-brain “intra-actions” (Barad, 1998, p. 96, noting “the inseparability of ‘objects’ and ‘agencies of observation’”) that can perpetuate shame, fear, and other forms of suffering.

This post reviews two films shown at the conference (Bitter Honey and Tales of the Waria) and three talks by neuroscientist Melissa Hines and anthropologists Hillard Kaplan and Marcia Inhorn.

Bitter Honey

On Day 1 and 2 of the conference, FPR founder and president, Robert Lemelson, a documentary filmmaker and psychological anthropologist on the UCLA faculty, screened Bitter Honey. Shot over a seven-year period, the film explores polygamous marriages through the lens of three Balinese families headed by Sadra (2 wives), Darma (5 wives) – both working class, and the elderly Tuaji “of royal blood” (10 wives). The film addresses seven themes: love and marriage, power, violence, children, lust and infidelity, divorce, and endurance and freedom.

In Bali, 10 percent of registered marriages are polygamous, although the percent of unofficial unions is likely higher. As the film illustrates, many of these unions are formed and maintained through deep-seated power dynamics that justify men’s infidelity and restrict women’s abilities to leave. For instance, when Sadra’s second wife, Murni, finally discovered his marital status, she was already pregnant and had decided to “[accept] it . . . [because she] took the risk and . . . had to take the responsibility.” This simple statement masks a grimmer reality. Tricking women into polygamous marriages appears to be all too common. To varying degrees Sadra and Darma’s wives have learned to adapt to their surroundings, maintain jobs to support the men’s lifestyles, and raise their children (Tuaji’s wives appear to lead more comfortable lives). But the film wordlessly and beautifully unveils the pain, isolation, and sense of confinement in the wives’ situation in ways that written ethnography cannot (Lemelson & Tucker, 2015).

Hints of infidelity and domestic violence permeate the three men’s marriages. Both Sadra and Darma frequent the red light district. Brothels catering to tourists have sprouted across rural areas of Bali and are also frequented by local men. Although an estimated 25 percent of Balinese sex workers are infected with HIV/AIDS, use of condoms appears infrequent, and the risk of transmission to the men’s wives is high. Regarding domestic violence, at one point in the film the documentary team – along with Dedung, Sadra’s boss, and Anggreni, a women’s rights attorney in Bali – are compelled to stage an intervention to attempt to stop Sadra from beating his wife.

Karma is an excuse many men use to escape from their responsibilities, as Degung Santikarma ­– Balinese anthropologist – explains in the course of the film. In Bali, it is commonly believed that men gain more power from having more wives; this draws from the Hindu concept of Bhairawa in which lust should not be restrained, but rather allowed full expression. However, men’s power is not the only force that constrains women’s abilities to leave these marriages. While in many regions of Indonesia the divorce rate is approximately 50 percent, it is less than 10 percent in Bali. In Balinese culture, the soul of the wife is thought to belong to that of the husband’s family, into whose lineage she will reincarnate; upon divorce, women lose not only their inheritance and custody of their children, but also their soul. (An exception is a nyentana marriage in which the man marries into his wife’s family and takes on the role of the eldest son, in which case the woman retains her inheritance and custody of the children after divorce.)

Bitter Honey poignantly documents the ways in which polygamous marriages in Bali are deeply embedded in cultural dynamics that are disadvantageous to women and trap them in an iterative cycle of vulnerability. In doing so, the film challenges universalistic paradigms that depict women as “coy choosy females” looking for well-resourced mates (Brown, Laland, & Borgerhoff Mulder, 2009) and resonates with Herdt’s documentation of change among the Sambia and Borgerhoff Mulder’s work among the Pimbwe in demanding cultural and historical specificity. While a scholarly accomplishment, Lemelson’s film work is also notable for the additional sensory experiences and, in particular, the sense of emotional intimacy with others that only a filmic medium can convey. Lemelson, who also trained as a clinical psychologist, feels that the use of more cinematic elements in the construction of his films – in combination with the deep, mutually respectful relationships he enters into with his subjects, who frequently participate and provide feedback during the editing process – have allowed him “to tell richer stories about fully fleshed out individuals” in “the multiple cultural and environmental contexts that suffuse any experience” (Lemelson & Tucker, 2015, p. 17, 29). His films push us to confront human suffering and re-consider the more engaged form of anthropology of many long-term practitioners (another example of which is Afflictions, Lemelson’s series on mental illness in Indonesia). Refusing to look away, Lemelson and his team have also begun working with local organizations to set up the first gender-based violence program in Indonesia.

Tales of the Waria

The conference program also included Kathy Huang’s documentary Tales of the Waria, which focuses on four “biological men” in Makassar, Indonesia  – Tiara, Mami Ria, Suharni, and Firman – who self-identify as women. The men are known locally as waria –“ a combination of the terms wanita (woman) and pria (man), which can be roughly translated as ‘male transvestite’” (Boellstorff, 2004, p. 160). Waria live openly as women, mainly engaged as performers (Tiara), in some form of salon work (Mami Ria, Suharni, and Firman), especially bridal makeup and hair styling, or as sex workers; thus, they are far more visible (and, as a “recognizable continuity” dating back to the early 1800s, far older in origin) than Indonesians who identify as gay or lesbi (Boellstorff, 2004).

Being a waria is not a matter of sexual orientation; according to Tiara, “waria exist to make this life more beautiful.” Further, “waria almost never describe themselves as a “third gender” but see themselves as men with women’s souls who therefore dress like women and are attracted to men” (Boellstorff, 2005, p. 57). Although they may take estrogen in the form of birth control pills and use injectable fillers, Tiara explains that “most warias don’t want a sex-change operation because of the teachings of Islam. We believe that we were born as men and must return to God as men.”

The film offers a humanizing portrait of the warias’ pursuit of love with a man (“every waria’s dream”), and in doing so, movingly depicts the dual sense “of belonging to (and exclusion from) national society and popular culture” (Boellstorff, 2004, p. 161) given their visible positions in society, including in the political sphere. At first glance, the love lives of the four warias “flow simply into the mold of [male] power,” to paraphrase Catherine Malabou (2015). Tiara is a performer who likens herself to Madonna or Beyoncé. Despite the general tolerance for warias in Indonesia, Tiara’s family does not openly welcome their son’s adoption of a waria identity; a former boyfriend refused to marry because Tiara wasn’t a “real” woman and couldn’t give him children (“I was just a place to stop until he found a woman”).

Younger warias, like Suharni, juggle achieving financial security and maintaining a relationship with their boyfriends. In Suharni’s case, her boyfriend Madi remains committed even after learning of her HIV-positive status (according to Suharni, “HIV/AIDS is the most feared disease in Indonesia”), but neither is able to make enough money living in Makassar, and Suharni decides during the course of the film to relocate to Bali. The oldest of the four, Mami Ria faces a different challenge in maintaining the interest of her partner Pak Ansar, who is married with children. Pak Ansar thinks warias are “creative” and “really have a passion for life,” and his wife, Ety, appears complacent with the arrangement. But as Mami Ria has gotten older and feels less inclined to wear make-up and maintain long, feminine hair, she believes Pak Ansar’s love has waned. By the end of Huang’s film, Mami and Pak Ansar had not spoken in more than three months.

“I walk two paths”

Some warias, like Firman, opt to marry heterosexually. In an opening scene, Firman prays to God to “change [him] into a real man” because it was “impossible to go back into being a waria,” which was “a terrible [past] mistake.” Firman said his family was ashamed of him as a child (“They always got angry at me and beat me”). His father hit him hard with a rattan switch and told him “to be a man, but I wasn’t able to.” Firman is now “respected” by his family and in-laws for being a good father and husband. Though his wife Mimi has heard stories of his past, she said she accepts him because she believes “he’s left his old ways behind” and is now committed to “making a future together . . . [and] wants to change.” Through tears, she adds, that Firman continues to go on late outings with his waria friends; though he brushes it aside as work, she cries and worries, not sleeping until he returns. Although Firman reaffirms his commitment to his family, he tells Huang that he nonetheless misses the warmth of a male body and contrasts the “ecstatic” feeling of “soaring into the sky” with the reality of waking up next to his wife and children.

Together, the four narratives illustrate the multiple ways in which warias pursue love in Indonesian society – not as a “third gender,” but occupying “a gendered subject position haunting maleness” (Boellstorff, 2004, p. 183) – and, like the wives in Bitter Honey, have learned to deftly negotiate the marginalizing social and cultural dynamics that continue to bound their experiences.

Our “volatile” anatomy

Most warias recount “atypical play” for as children (“I hung out with the girls and played jump rope, played with dolls”; Boellstorff, 2004, p 166), either because their parents didn’t prohibit it or, as many attest, they were simply born “with the soul of a woman.” Their various reflections – in some cases, biologically oriented; in others, social – for the basis of their childhood toy preferences resonated with neuroscientist Melissa Hines’s talk (“Early Androgen Exposure and Human Gender Development: Outcomes and Mechanisms”). Hines focuses primarily on the role of testosterone in influencing human gender development; part of her research program explores gender typical play.

According to Hines, the brain is not hardwired to be male or female. But neither is gender identity wholly a function of “self-socialization or socialization by others” postnatally. Both viewpoints are partial and thus “flawed” because neither takes the other into account. Our anatomy, as Elizabeth Wilson (2015) argues in Gut Feminism, is “volatile enough” to produce many “multifaceted . . . destinies.”

Four factors influence human gender development: genetic factors, gonadal steroids (particularly testosterone during early development), social reinforcement, and self-socialization, Hines continued. Her remarks focused on the influence of testosterone on brain and behavior, whose “enduring masculinizing and defeminizing effects during early sensitive periods of rapid brain development” have been well-studied in non-human animals (Hines et al., 2016a).

Prior to puberty, testosterone is higher in males than in females during two periods of rapid brain development: prenatally (from about 8–24 weeks) and postnatally (also referred to as “mini-puberty,” from about 1–6 months; Hines et al., 2016b). In humans, evidence suggests the first (prenatal) elevation influences three main behaviors that differ by sex (1) gender identity, (2) sexual orientation, and (3) gender-typical play. The second elevation contributes to later gender-typical play in particular.

How can we study the influence of hormones in human development?

The strongest evidence comes from studies of individuals with genetic conditions. Hines’s research focuses mainly on congenital adrenal hyperplasia (CAH), a genetic disorder. CAH disrupts the adrenal glands’ ability to make cortisol. The resulting dysregulation in cortisol-mediated adrenocorticotropic (ACTH) secretion by the pituitary gland causes the overproduction of adrenal androgens, beginning prenatally. Despite some gender ambiguity in external genitalia at birth due to prenatal exposure to high levels of testosterone, girls with CAH, who are treated postnatally with glucocorticoids, are usually raised as girls. But girls and women with CAH show differences in all three gendered behaviors.

Hines discussed toy preferences, which her research has related to both prenatal androgen exposure and gender-related models (persons of the same or opposite sex choosing gender-neutral objects) and labels (being told certain objects were “for girls’ and others were “for boys”; see Hines et al., 2016a). Girls with CAH spent more time with the boys’ toys and less time playing with girls’ toys than their unaffected relatives. To determine whether the effects were due to socialization (e.g., parent support of children’s toy preferences), Hines and colleagues also studied toy preferences among vervet monkeys and found similar sex differences (Alexander & Hines, 2002; see also Hassett, Siebert, & Wallen, 2008). Further, girls with CAH appeared to be less susceptible than other children to same-sex modeling and to gender labels. To Hines and colleagues, this suggests an interaction between prenatal androgen exposure and cognitive processes related to gender identity. Her group also measured urinary testosterone in infants during “mini-puberty,” which predicted male-typical play behavior at ages 3–4 (Hines et al., 2016b). Hines concluded that testosterone contributes to the development of human gender-related behavior, but “there are many dimensions of gendered behavior and different dimensions are influenced by different combinations of factors,” including self-socialization in response to same-gender models or gender labels as wells the influence of parents and peers.

Next, the focus widened as anthropologist Hillard Kaplan (“Embodied Capital and the Sexual Division of Labor: Evolution of Multiple Time Scales”) presented data on societies that practice hunting and gathering or a mix of foraging and horticulture. These societies offer “one particular lens on the evolved biology of our species,” according to Kaplan. “Comparisons with modern societies can shed light on the interactions of genes, environments, and lifestyles on behavior, health, and longevity and, in particular, . . . modern health conditions.”

Kaplan briefly reviewed fertility, mortality, and net caloric production (“how much food you produce less how much you consume”). Marriages in hunter-gather societies are mostly monogamous, reflective of “some strong complementarities in the life history of the two sexes,” which is relatively rare among other mammals. Women are primarily committed to maternal caregiving and men to learning- / skill-intensive hunting (the peak in male muscular strength well precedes the peak in men’s hunting ability). Men provide most of the surplus calories in the form of hunted meat, a new and valuable resource from an evolutionary standpoint, but one that is “very expensive in terms of care.” Kaplan’s talk helped us understand how men’s and women’s pursuits are delicately balanced (“the value of what women do for their families depends upon what the men do, and vice versa, and their life histories become linked”). The sex difference results because childbearing women have less time (between births) to acquire the necessary skills to be successful at hunting.

The remainder of the talk focused on the Tsimané of Bolivia, a foraging-horticulturalist population that still follows a “traditional subsistence pattern.” In particular Kaplan discussed various adaptations as men and women age. Beyond the mid-thirties, men tend to hunt less and farm more. As women age, they also engage more in agricultural activities, becoming major contributors to total caloric production. Overall, resources flow downward from grandparents to their children and grandchildren – or, with fewer dependents, to closely related, frequently younger households (Hooper, Gurven, Winking, & Kaplan, 2015). Kaplan uses a unified model (time-path of production and inclusive fitness theory) to account for these “exquisitely patterned” flows. Elsewhere he has likened them to a form of “indirect” reproduction, “which has allowed selection to favour the evolution of significant post-reproductive lifespan” (Hopper et al., 2015). He particularly emphasized the role of grandfathers (as well as grandmothers) in the downward flows to grandchildren; these men tend not to seek out new reproductive relationships. Like Monique Borgerhoff Mulder, he stressed the importance of marriage (and the relative rarity of polygamy), especially the importance of choosing a good partner, and of cooperation between the sexes and in terms of intergenerational investment. The final segment of his talk focused on the adaptability of the division of labor to changing social and economic contexts and gendered effects on health and mortality. Today, women tend to live longer than men but report worse health, reflecting the interaction of ontogeny and selection for certain “physiological and psychological pre-commitments.”

Cultural and medical anthropologist Marcia Inhorn’s talk (“Male Infertility, Assisted Reproductive Technologies [ART], and Emergent Masculinities in the Arab World”) focused on the globalization of technologies addressing male infertility, which Inhorn argues has been accompanied by the “emergence” of new forms of manhood in the Arab Muslim Middle East.

Inhorn’s work challenges stereotypes of Middle Eastern men as “fanatically religious” and “brutal oppressors of women.” Her long-term ethnographic research illuminates transformed attitudes about love, marriage, and fatherhood. Inhorn’s large amount of data, which is pooled from more than 330 Arab men from 14 Arab countries over 15 years, demonstrate a reality that bears no resemblance to caricatures of Arab Muslim men in the media. For the most part, the men are family-oriented, seek love/partnership in marriage, and highly value fatherhood.

Regarding the reproductive piece, in vitro fertilization (IVF), a “platform technology,” has been joined by an “unbelievable array” of ARTs, including third party reproductive assistance, gestational surrogacy, cryopreservation (freezing or vitrification), preimplantation genetic diagnosis (PGD), human embryonic stem cell (hESC) research, and even the possibility of human reproductive cloning. Inhorn’s talk focused on intracytoplasmic sperm injection (ICSI), a variant of IVF designed specifically to address male infertility, which she described as an “underappreciated global reproductive health problem.” Across the world about 9–10 percent of couples are infertile, and about half of those cases are due to male infertility. In the Middle East, however, male infertility rates are much higher, primarily for genetic reasons due to high rates of marriages among kin (sub-Saharan Africa has even higher rates due to reproductive tract infections and various other complications, according to the World Health Organization). Before ICSI, male infertility was highly stigmatizing, due to the conflation of infertility and sexuality, as well as incurable. ICSI addresses low sperm count by directly injecting a single spermatozoon into the oocyte.

In the Sunni-majority Muslim countries, which prohibit the use of donor sperm (or any third party insemination), the advent of ICSI in the early 1990s was “a watershed event.” After years of struggle, men in the Middle East view ICSI as a “hope technology” (although Inhorn pointed out that it also shifts the genetic burden onto their male offspring). The “double” emergence – ICSI and “emerging changes in gender relations and, ultimately, masculinity” – arises by making long-term love-marriages and reproduction viable for more men.

Overall, the collection of talks and films presented human life on multiple time scales. The film Bitter Honey suggested a disintegrating past (the children in the polygamous families seem averse to the practice), while Tales of the Waria and Marcia Inhorn’s talk provided glimpses of a more generous future that for some – like Mami Ria and Sadra and Darma’s wives, with few social anchors –remains out of reach. Hillard Kaplan and Melissa Hines grounded us in evolutionary and biological histories that deftly wove the social and biological together. While all the elements may not fit together perfectly, the composite picture addressing what counts as “adequate function” drawn from many disciplines and interdisciplinary programs, is far richer and more dynamic than we anticipated. Our final blog post addresses what’s at stake.


Alexander, G. M., & Hines, M. (2002). Sex differences in response to children’s toys in non-human primates (cercopithecus aethiops sabaeus). Evolution and Human Behavior, 23, 467479.

Barad, K. (1998). Getting real: Technoscientific practices and the materialization of reality. differences: A Journal of Feminist Cultural Studies, 10(2), 87–128.

Boellstorff, T. (2004). Playing back the nation: Waria, Indonesian transvestites. Current Anthropology, 19(2), 159–195.

Boellstorff, T. (2005). The gay archipelago: Sexuality and nation in Indonesia. Princeton, NJ: Princeton University Press.

Brown, G. R., Laland, K. N. & Borgerhoff Mulder, M. (2009). Bateman’s principles and human sex roles. Trends in Ecology & Evolution, 24(6), 297–304.

Hassett, J. M., Siebert, E R., & Wallen, K. (2008). Sex differences in rhesus monkey toy preferences parallel those of children. Hormones and Behavior, 54(3), 359–64.

Hines, M., Pasterski, V., Spencer, D. Neufeld, S. Patalay, P., Hindmarsh, P. C. . . . Acerini, C. L. (2016a). Prenatal androgen exposure alters girls’ responses to information indicating gender-appropriate behavior. Philosophical Transactions B, 371, 20150125.

Hines, M., Spencer, D., Kung, K. T.-F., Browne, W. V., Constantinescu, M., & Noorderhaven, R. M. (2016b). The early postnatal period, mini-puberty, provides a window on the role of testosterone in human neurobehavioural development. Current Opinion in Neurobiology, 38, 69–73.

Hooper, P. L., Gurven, M., Winking, J., & Kaplan, H. S. (2015). Inclusive fitness and differential productivity across the life course determine intergenerational transfers in a small-scale human society. Proceeding of the Royal Society of London B: Biological Sciences, 282(1803), 20142808.

Lemelson, R., & Tucker, A. (2015a). Lemelson, R., & Tucker, A. (2015). Steps toward an integration of psychological and visual anthropology: Issues raised in the production of the film series Afflictions: Culture and Mental Illness in Indonesia. Ethos, 43(1), 6–39.

Malabou, C. (2015). One life only: Biological resistance, political resistance. Critical Inquiry. This essay was originally published in French as Catherine Malabou, “Une Seule Vie: résistance biologique, résistance politique,” Esprit (January 2015), 30–40.

Wilson, E. A. (2015). Gut feminism. Durham, NC: Duke University Press.

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Part 2 of the FPR-UCLA conference on sex/gender, which was chaired by cultural anthropologist Gilbert Herdt, explored aspects of brain and behavior that are “fixed” by evolution and biology and other aspects that create, reflect, and respond to human social and cultural environments. Speakers in the first session addressed, in Darwin’s phrase, the “entangled bank” of biological, evolutionary, and cultural contexts of sex/gender differences in brain and behavior, while the second session offered a closer examination of “intimacies”– partnerships, marriage, sexual orientations, desires, and practices. A common theme throughout was the instability of the sex/gender binary or, as Carol Worthman observed, the “loss of easy dichotomies” more generally. Perspectives varied widely depending on level of analysis, but there was a general willingness to “work with and speak across difference” (Worthman).

For neurobiologist Donald Pfaff, who presented experimental research focusing on autism, sex is a biological category/variable. Other speakers were more willing to extrapolate from biology, and in so doing, challenge what Sarah Richardson referred as our “essentialist” understandings. For social neuroendocrinologist Sari van Anders, even a “quintessential” male hormone like testosterone can be deconstructed (van Anders, 2013). Another common theme was gender-role and sexual fluidity, addressed from evolutionary (Fessler), hormonal (Rilling, van Anders), and situational/contextual (Diamond) perspectives. Finally, field research by anthropologists (Borgerhoff Mulder and Boellstorff) in non-Western and virtual settings underscored human flexibility and adaptiveness.

In addition, on Friday afternoon, FPR founder and president, Robert Lemelson, a documentary filmmaker and psychological anthropologist on the UCLA faculty, screened Bitter Honey. Shot over a seven-year period, the film explores polygamous marriages through the lens of three Balinese families (Sadra, Darma, and Tuaji). Next, filmmaker Kathy Huang presented her film Tales of the Waria, which focuses focused on a group of biological men who self-identify as women—known locally as waria in Indonesia. Part III of our series reviews the two films in depth.

The talks and the two films revealed significant advances in our understanding of the underlying mechanisms and dynamic aspects of sex/gender-related behavior and their exquisite attunement to historically and culturally specific environments. As conference keynote speaker Anne Fausto-Sterling reminds us, “neither the body nor experience is prior, but each acts continuously upon the other as individual patterns of behavior and of neuronal connections appear” (Fausto-Sterling, 2014, p. 315).


Prenatal and perinatal factors, including maternal stress, profoundly influence brain and behavioral development. Neurobiologist Donald Pfaff (“Neuroendocrine Mechanisms Underlying Prenatal Stress in Effects and Sex Differences in Autism”) focused on one way in which iterative development can go awry. Autism Spectrum Disorders (ASD) involve motor, language, and social deficits, with the latter considered “the core concept” of the diagnosis. ASD is both highly heritable and sex-related (4:1 male-to-female ratio, increasing to 11:1 “with the highest levels of intellectual capacity; Gillberg et al., 2006, as cited in Schaafsma & Pfaff, 2014). Pfaff and colleagues study the sex-differentiating factors underlying male bias in ASD.

Pfaff briefly outlined several non-mutually exclusive pathways to sex-differentiation in brain function that may have possible implications for sex-specific susceptibilities in or protection from ASD: (1) via Y-linked genes, such as Sry, which, as Art Arnold explained, has both direct and indirect (via gonadal hormones) effects – i.e., some are directly expressed in the brain and play a role in catecholaminergic functions; (2) X-inactivation “in the service of dosage compensation,” which can be balanced or skewed; (3) genetic imprinting – the epigenetic silencing of one of the two working copies of a gene inherited from parents; and (4) other epigenetic-related factors (see Schaafsma & Pfaff, 2014, for more detail).

Male sex and the “3-hit” theory of autism”

Pfaff described the “3-hit” theory of autism as an interaction between: (1) androgenic hormones; (2) early stress; and (3) one of the genetic mutations associated with autism in males, such as the CNTNAP2 mutation. Regarding Steps 1–2, testosterone affects arousal neurons in the forebrain; which results in greater activity in their ascending pathways, especially increased deposition of norepinephrine, dopamine, and serotonin in the amygdala. The amygdala becomes “supersensitive” and responds more acutely to social stressors, pre- and postnatally. Chronic overactivation in response to stress leads to social anxiety and ultimately the social avoidance characteristic of ASD.

Pfaff and colleagues use a mouse model to test exposures to prenatal stress/no stress on a 0–3 event scale in Cntnap2 knockout and wild-type mice. (NB: They used maternal immune activation for prenatal stress because infection in pregnancy predisposes to autism.) The researchers found differences between the 0-hit and 3-hit mice based on three tests: ultrasonic vocalizations, social recognition (a series of home-cage exposures to an intruder mouse, followed by an exposure to a novel mouse), and social approach in a three-chambered cage. Interestingly, there was a slight increase in social approach for the 3-hit mice, and no differences in anxiety between wild-type no-stress females and the male mice in an open field test with various numbers of hits. The researchers sacrificed the animals and found significantly lower expression of mRNA receptors for stress-related corticotropin releasing hormone (CRH-R1) in the left hippocampus in the 3-hit mice. In the final piece of the story, the researchers harvested mRNA and found developmental sex differences in mRNA levels of connexin-36, which is important for mediating electrical synapses and promoting neuronal synchrony. Males express more connexin-36 in the amygdala than females, which has possible implications regarding the sex-related (male) amygdala hypersensitivity of the 3-hit model.

This emerging body of research focuses on the underlying neuroendocrine and epigenetic causes of the sex-related differences in ASD and also links the social behavior consequences (e.g., anxiety) to the number of “hits” over time, underscoring the iterative nature of sex-differentiation, a major theme of this conference.

Biocultural anthropologist James Rilling (“Neural, Hormonal, and Genetic Correlates of Human Paternal Behavior”) shifted the morning’s focus from mother–infant dyads to fatherhood. As Ruth Feldman (2015) observed in a recent review, parenting may be the most delicately poised among all social phenomena between evolutionarily conserved components on the one hand and “the greatest plasticity” on the other. Fathers are less well studied, but Rilling’s research suggests significant plasticity in adult male as well as female brains in response to caring for offspring.

“Mothers and others”

Some biologists refer to humans as “cooperative breeders,” that is, the source of care and feeding of offspring comes from both “mothers and others” (Hrdy, 2009), albeit with significant cultural and interfamilial variation. According to Rilling, in modern Western societies that often consist of isolated nuclear families, fathers are often the greatest source of help to mothers. Emerging evidence indicates that paternal involvement is associated with “multiple positive developmental outcomes in children” in Western settings, he said.

Rilling’s talk addressed two questions: “Why is it that some men are more involved as fathers than others? Can we identify variables that are correlated with or influence paternal involvement?” His biocultural approach is based on an evolutionary and life-history perspective, more specifically for the purposes of his talk, on variation in and tradeoffs between investing energy in mating and in parenting.

Rilling’s research focuses on the hormonal (testosterone, vasopressin, oxytocin, and prolactin) and neural (mesolimbic dopaminergic pathway) mechanisms regulating behavior in fathers of young children (ages 1–3). In rats, the medial preoptic area (MPOA) of the hypothalamus, which is subject to the influence of steroid hormones, regulates parallel systems for pup approach/avoidance. Bathing the area in hormones activates the mesolimbic dopamine system (including the ventral tegmental area in the midbrain, the nucleus accumbens [part of the ventral striatum], and the medial orbitofrontal cortex), “a classic reward system pathway,” enhancing maternal motivation to nurture.

Rilling and colleagues recruited three groups of men: nonfathers, more involved fathers, and less involved fathers of children (ages 1–3) to study both hormone levels and brain function in response to visual child-related and sexual stimuli in order to explore tradeoffs between mating and parenting. Fathers had lower levels of plasma testosterone than nonfathers. They were also about 20 pounds heavier than nonfathers, putatively reflecting a significant negative correlation between testosterone and body fat. Fathers also had significantly higher levels of plasma oxytocin than nonfathers. In fathers baseline plasma oxytocin appears to enhance certain aspects of caregiving, such as stimulatory parent-infant contact (Feldman, 2012; as cited in Young & Rilling, 2014).

Rilling and colleagues also compared the more and less involved fathers (using the Parental Responsibility Scale). They noted weak negative correlations between testosterone and instrumental caregiving and testes size and caregiving, although the relationship between testosterone and testes size was not statistically significant.

In terms of brain function, fathers viewed images of their own children (with happy, sad, and neutral facial expressions), unknown children, unknown adults, and sexually provocative stimuli. The investigators also primed an empathic response from the fathers by asking them to “try to share the emotions of the person in the picture.” Across the board, fathers had stronger responses than nonfathers to the images of unknown children in such areas as the medial orbitofrontal cortex (part of the “reward” system) and the temporoparietal junction, which plays a putative role in theory of mind, or the ability to make inferences about the mental states of others. Across the board, the nonfathers had stronger responses to the sexually provocative visual stimuli, particularly in brain regions related to goal-related motivation and reward (e.g., the nucleus accumbens).

More involved fathers had higher activation of the ventral tegmental area (VTA), part of the network motivating approach to offspring in the rat studies, when viewing own children. There was also a more robust relationship – a negative correlation – between testes size and VTA response to viewing images of own children. Finally, fathers listened to infant cry stimuli, which among fathers notably activated the anterior insula (AI) bilaterally. Among other functions, the AI tracks sympathetic autonomic arousal. “Less involved” fathers had a lower or higher AI response, respectively, than fathers who were “most involved in instrumental caregiving” suggesting empathic under or over-arousal in the less involved group (Young & Rilling, 2014).

The final speaker of the afternoon, anthropologist Monique Borgerhoff Mulder (“Gender Roles in Mpimbwe: Re-evaluating Bateman’s Gradient”), explored the effects of environment on intra- and extra-marital relations among the Pimbwe of the Rukwa Valley in western Tanzania. Her data challenge universalist gender roles drawn from standard sexual selection theory.

Due to conservation strategies, Pimbwe men can no longer legally hunt or fish in what is now Katavi National Park, and job scarcity makes education an unsatisfactory strategy for making a living in the Rukwa Valley. As a result, Pimbwe women are relatively powerful players within the marriage market. Not only do women control much of the gardening, environmental factors – such as unreliable rainfall, agricultural pests, depredating wildlife, and theft from other villagers – have compelled them to engage in other economic activities that can help offset intermittent food shortages. “Under these tough economic, ecological, and environmental conditions “marriage is problematic,” Borgerhoff Mulder continued, and finding a good provider is critical. Marital decisions are particularly important because Pimbwe are increasingly unable to rely on cooperation from neighbors and kin (Kasper & Borgerhoff Mulder, 2015); cooperation with a marital partner becomes key, placing a lot of pressure on this relationship. Against this background Borgerhoff Mulder analyses the considerable variability in gender roles and a strong prevalence for marital systems that are “very flexible.” Monogamy, serial polygyny, and serial polyandry are common arrangements; extramarital affairs also abound (if a pregnancy results, it typically leads to marriage), and there is increasing polygyny.

“Why do Pimbwe men suffer from multiple spouses?”

Networks of various marriages (which the Pimbwe effectively define as pairs who cohabit), divorces, and extramarital mating patterns produce outcomes that provide an exception to Angus J. Bateman’s (Arnold 1994) foundational claim that number of mates will have stronger effects on the fitness of males than females. This famous “Bateman’s Third Principle” follows from his first two principles – that males will have higher variance in fitness (measured as number of surviving offspring) than females, and that males will have higher variance in the number of their mates than females. All of these differences are predicated on the fact that, by definition, males have smaller gametes than females.

Using longitudinal, cross-sectional, demographic, and economic data for all households in one village in the Rukwa Valley, Borgerhoff Mulder examined variation in reproductive success (Bateman’s first principle) and mate number (his second principle), before modeling the effects on reproductive success of the number of an individual’s mates (the third principle). Regarding reproductive success below age 40, the variance for men and women was “pretty similar.” In accordance with Bateman’s theory, as they aged beyond 40, men had a higher variance than women. Similarly mate number was equally variable between the sexes. But regarding Bateman’s third principle, Borgerhoff Mulder found that having multiple spouses had a positive effect on mating success in women (i.e., production of surviving offspring) and a negative effect in men.

The Pimbwe challenge the “conventional view of promiscuous, undiscriminating males and coy, choosey females” (Brown, Laland, & Borgerhoff Mulder, 2009).  Some Pimbwe women can “discard” – and benefit from discarding – husbands who are not good providers, in poor health, or just considered “lazy.” According to Borgerhoff Mulder, these women tend to be hard workers who can afford “to cycle through multiple marriages.” Despite the fact that some Pimbwe maintain successful polygynous marriages over a number of years, many Pimbwe men face the opposite fate. Reproductive success falters as they cycle through marriages, unable to keep their spouses from leaving them; they seem to make poor fathers. Borgerhoff Mulder theorized that the constant shift between multiple spouses by females, and the higher variance in mate quality that accompanies it, reflects the value system around pair bonding. That is, rather than aligning with conventional accounts, which suggest that in societies where divorce occurs, “pair bonds” must not be “very important,” Borgerhoff Mulder proposed the opposite: “Divorce may actually be an indicator of the importance of pair bonds” rather than their non-importance. That is, in societies where pair bonding is valued highly, stakes are higher, and therefore there is higher pressure to cycle through multiple spouses until the best one is found.

Evolutionary anthropologist Dan Fessler (“An Evolutionary Perspective on Sexual Orientation, Same-Sex Attraction, and Affiliation”) opened the second day of the conference with an evolutionary puzzle. Given the role of reproductive success in natural selection, “if there is any heritable contribution to sexual orientation, how have alleles for exclusive same-sex attraction persisted over evolutionary time?”

Human and animal studies find substantial heritability in sexual orientation, including same-sex attraction, which is most likely the effect of a number of genes and/or epigenetics, both of which are subject to natural selection. Existing (partial) explanations, such as pleiotropy or kin selection, focus on male homosexuality and “leave lots of variance unexplained.” Fessler focused on “overdominance,” or the “heterozygote advantage” of having different alleles of the same gene, theorizing that human sexuality is multifunctional. Here, Fessler was careful to distinguish between sexual orientation – “[culturally] patterned sexual desire that may lead to sexual behavior with members of one or the other sex” (Fleischman, Fessler, & Cholakians, 2015), which is closely related to gender, and same-sex attraction, his primary interest, which may or may not align with sexual orientation.

It is difficult to get precise numbers on same-sex sexual behavior; there are “huge” cross-cultural differences. For example, in some cultures same-sex sexuality is unknown, in others it is widespread, and in still others – as in the case of the Sambia – it is “required ritually.” He suggested that, “in the absence of norms prohibiting same-sex behavior, it apparently arises spontaneously,” often during sexual development.

Same-sex sexual behavior occurs across species, although regularity and context vary, and may have two primary nonreproductive functions: one signifying affiliation (e.g., the alliance-enhancing same-sex bonds among the bonobos) and the other dominance (e.g., dogs mounting). Further, Fessler argued that “plasticity and flexibility” in sexual behaviors and corresponding attraction can be subject to natural selection. Returning to the concept of overdominance and particularly “multifunctional sexuality,” he cited work suggesting that same-sex sexual behavior supports the kind of alliance formation Herdt described in his talk. And note, he continued, that “this is compatible with existing evidence suggesting that sexual orientation and sexual psychology are separate things” in that the “target has shifted” to the same sex but the sexual psychology remains the same. Fessler also discussed the fitness-enhancing aspects of affiliative bonds. In nonhuman primates, “social bonds enhance survival and reproductive success.” Alliances were also vital in small-scale human societies for a number of reasons (for enhanced success in violent coalitional conflict, as a buffer against food shortfalls, to facilitate alloparenting, and as insurance against illness and injury), Fessler said.

Where are all the bisexuals?”

Using Darwin’s finches as an example, Fessler showed how three allele pairs yield seven genotypes and a continuously distributed trait, or bell-shaped curve. He also noted the extent to which environment affects the expression of genes: “when the effect of each locus is small, environmental variation will blur genetic differences.” Given the multiple genetic contributions to same-sex arousal, the normal distribution of phenotypes would assume a bell-shaped curve, ranging from a committed same-sex phenotype with no reproduction “but lots of allies” to a committed opposite-sex phenotype with reproduction, but “fewer allies.” According to Fessler, most people would be “flexible” between the two. The “flexible” phenotype “accrues benefits of both same-sex contact and reproduction, depending on context.” Cultural pressures tend to shift normal distribution toward the right. His take-home message was that, aside from “obligate” same-sex and opposite-sex individuals on the far left and right of the bell curve, “for most people, the cultural context and the socialization experiences probably do profoundly influence their self-concept and their experiences of sexuality.”

The next speaker, social neuroendocrinologist Sari van Anders (“Social Neuroendocrinology, Gender/Sex, and Sexual Desire: Testosterone as Socially Constructed and Evolved”) emphasized the iterative or bidirectional relationship between hormones and behavior and, above all, the importance of context. Their dynamic interactions occur within “a space that is both social . . . and responsive to evolution.” Van Anders’s approach to “gender/sex” focuses on the interaction of the two constructs. “We can think of sex as relating to femaleness, maleness, and sexual diversity,” she said. “[S]ex falls into the part of the equation that is evolution. And we can think of gender as falling into the part that is social context and that has to do with gender diversity, femininity, and masculinity. . . . [B]y gender/sex, I mean I’m studying whole men or whole women or whole gender/sex diverse people.”

Van Anders stressed the importance of studying “human specificities,” which often get overlooked, particularly the integration of social constructions (“shared cultural understandings that vary by place and time”) and biological constructions (including contextually sensitive steroid hormones like testosterone), which modify one another. The remainder of her talk focused on testosterone and sexual desire. She described “testosterone” as both a biological material and a social construction. Her point is that “the stories we tell ourselves” about testosterone pretheoretically influence how scientific research is conducted. For example, one of the tacit assumptions of testosterone is that it promotes masculinity and is negatively related to femininity. Van Anders proposed a different model for understanding variability in testosterone based on a broader definition of sexuality that extends beyond reproduction. Her model (the Steroid-Peptide Theory of Social Bonds) suggests “an overarching social role [that is] similar in women and men,” one in which low testosterone is linked to nurturance (“loving, warm contact”) and high testosterone to competition (“acquiring and keeping resources”; see van Anders et al., 2011; van Anders, 2013; Goldey & van Anders, 2015), which brought to mind James Rilling’s earlier talk on life history tradeoffs.

“There are many constructions of desire”

Van Anders discussed the concept of “sexual desire,” taking apart some of its socially constructed assumptions (e.g., sexual desire is prototypically male, dyadic, orgasm-focused, and testosterone-fueled) and drawing out its extraordinary sensitivity to context and particular functions. A straightforward link between testosterone and sexual desire in men or women is tenuous at best. Instead, “there are many constructions of desire,” which have both positive and negative correlations with testosterone. In her nurturance–competition model, nurturance/low testosterone is linked to closeness, pair bonding, and self-comfort, whereas competition/high testosterone is linked to erotic pleasure, power, orgasm, and jealousy. The idea is that “the expectations of erotic pleasure and orgasm actually can differ based on social location and sexual experience and other factors.” For example, in heterosexual women, researchers found a negative correlation between dyadic sexual desire and testosterone, suggesting the women’s “desire” is more closely related to nurturance than expectation of orgasm. The researchers found no correlation between sexual desire and testosterone in men, which also ran counter to our pretheoretical assumptions.

Next, developmental psychologist Lisa Diamond’s talk (“Where Does Sexual Orientation Reside?”) challenged the once-entrenched belief in a binary, innate, and stable distinction between same and other-sex “orientation” (“the general predisposition to experience sexual attraction”), beginning with the observation that individuals who are sexually oriented to the opposite sex are capable of having same-sex desires, and vice versa. Longitudinal studies have uncovered considerable “plasticity” or “fluidity” in same-sex and other-sex attractions and behavior (Diamond, 2012), particularly among women. Diamond defined sexual fluidity as “situational and contextual variability in the experience and expression of same-sex and other-sex sexuality over the life course.” To address this greater complexity, Diamond conceptualized same-sex sexuality in terms of “constitutional same-sex sexuality” and “facultative same-sex sexuality.” The former is rooted in “same-sex desires stemming from one’s orientation.” The latter refers to “same-sex desires facilitated by situational or environmental factors, which can vary significantly in women over time. Elsewhere, Diamond has argued that this variability “may constitute a fundamental feature of female sexual orientation” and may be particularly amenable to a dynamic systems theoretic approach (Diamond, 2012).

Diamond pointed out the difficulty of differentiating between constitutional same-sex sexuality (a “gold star lesbian”) and a situational lesbian or “lesbian until graduation” on the basis of behavior. Even physiologically, “most women possess ‘nonspecific’ patterns of genital arousal” to same-sex and other-sex stimuli, “regardless of their own self-reported sexual orientation” (Diamond, 2012, p. 76; citing Chivers & Bailey, 2005; Chivers et al., 2004; Suschinsky, Lalumiere, & Chivers, 2009). (It’s important to note that “women are more likely than men to show discrepancies between their physiological and subjective arousal [for review, see Chivers et al., 2007],” and these discrepancies are not well understood.)

Diamond’s recent study looked at a different measure of desire: shifts in the responses of women of various self-identified orientations to sexual stimuli during ovulation, when estrogen levels peak. The diversity of responses by women of various sexual orientations suggests the need for a “biopsychosocial understanding” of how genetically influenced constitutions “interact with facultative environments, which range from microlevel processes – individual and dyadic relationships – to broad cultural environments, to produce different phenomenologies of desire, arousal, and behavior and then the pleasure that is taken from behavior.” She characterized “each individual desire” as “its own phenotype because it represents a very specific and dynamic interaction between genetically influence processes, biologically mediated processes, and, obviously, socioculturally facilitated and embedded processes. . . .  [S]exual orientation is neither the genetic part of same-sex sexuality nor the socially categorized part,” she continued, “If [we] want to make a distinction between constitutive and facultative desires, we have to understand the constellation of these different inputs and outputs over time.”

Cultural anthropologist Tom Boellstorff (“Technology and Globalization: Emergent Intersections of Culture, Brain, and Behavior”) discussed two long-term research projects focusing on globalization and technology: one on gaylesbi, and waria Indonesians and the other on culture in virtual worlds (in particular, in the virtual world Second Life). Boellstorff began by remarking on the “inability to fuse or separate sexuality and gender,” which reflects the broader issue of the nature/culture or biology/culture dichotomy that “still dogs us.” For Boellstorff, “long-term engagement with a field site” in Indonesia “allows you to track forms of change . . . how forms of movement and mobility are shaping culture and shaping sexuality and gender.” Both people and ideas are “moving.” The language is incorporating new words (gay, lesbi) that suggest homogenization but also “new forms of difference.” However, like sex/gender being “gay and Indonesian” or in most cases “gay and Muslim” “never fuse but they never become separate; “[i]t’s about the juxtaposition that nonetheless becomes part of everyday experience.” For example, gay and lesbi Indonesians are often married heterosexually – the two worlds needn’t resolve into one, though forms of oppression shape these dynamics. The fragmented globalization of sexual identities is exemplified by the fact that the “coming out” metaphor is quite rare in Indonesia: the more common term, “opening” to the gay, lesbi, or heterosexual “worlds,” has different implications for selfhood and community.

What is changing? Space and place

For Boellstorff, technology, and engagement with digital media in particular, serves as a means to challenge binary thinking, not only sex/gender, gay/straight, real/virtual, but also – as van Anders reminded us – biology/social constructions. His second research project is an ongoing ethnography of Second Life, a 3D virtual world whose members – via avatars – can fashion a less socially or physically encumbered identity, interact with one another in real time, immerse themselves in a range of social activities, and grow socially meaningful communities. In his 2015 preface to Coming of Age in Second Life (Princeton University Press, 2008), Boellstorff writes about an 85-year-old friend with Parkinson’s who is also a Second Life resident. For Fran, both bodies are real – her physical body (with Parkinson’s) and her virtual body, which “made it possible to wear a ball gown,” whirl around a dance floor with Tom, and run a support group in a wooden cabin on Namaste Island for other Second Lifers living with Parkinson’s. Although Boellstorff urged the audience to look beyond the binarism of real/virtual worlds, neither do they resolve into one. Instead, he stressed their coexistence and equal importance to ethnography.

Overall, the transdisciplinary and interdisciplinary sex/gender perspectives in Part 2 challenged many of our theoretical presuppositions about what is fixed by nature and shed light on what is changeable and changing, including how we study sex and gender. As one of the speakers observed, bringing biology and culture together “makes for transformative science.”


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