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American J Phys Anthropol - 2013 - Gray - Evolution and Human Sexuality
American J Phys Anthropol - 2013 - Gray - Evolution and Human Sexuality
KEY WORDS evolution; sexual behavior; sexual function; sexual selection; hominin;
Homo; life course; life history; genetics; neuroendocrine; polygyny
ABSTRACT The aim of this review is to put core fea- uality. As examples, the availability of rapidly increasing
tures of human sexuality in an evolutionary light. genomic information aids comparative approaches to dis-
Toward that end, I address five topics concerning the cern signals of selection in sexuality-related phenotypes,
evolution of human sexuality. First, I address theoretical and neuroendocrine studies of human responses to sex-
foundations, including recent critiques and develop- ual stimuli provide insight into homologous and derived
ments. While much traces back to Darwin and his view mechanisms. Fourth, I consider some of the most recent,
of sexual selection, more recent work helps refine the large, and rigorous studies of human sexuality. These
theoretical bases to sex differences and life history allo- provide insights into sexual behavior across other
cations to mating effort. Second, I consider central mod- national samples and on the Internet. Fifth, I discuss
els attempting to specify the phylogenetic details the relevance of a life course perspective to understand-
regarding how hominin sexuality might have changed, ing the evolution of human sexuality. Most research on
with most of those models honing in on transitions from the evolution of human sexuality focuses on young
a possible chimpanzee-like ancestor to the slightly polyg- adults. Yet humans are sexual beings from gestation to
ynous and long-term bonded sociosexual partnerships death, albeit in different ways across the life course, and
observed among most recently studied hunter-gatherers. in ways that can be theoretically couched within life his-
Third, I address recent genetic and physiological data tory theory. Am J Phys Anthropol 57:94–118,
contributing to a refined understanding of human sex- 2013. VC 2013 Wiley Periodicals, Inc.
The aim of this review is to put core features of David Buss’ (2003) The Evolution of Desire. The field
human sexuality in an integrative, evolutionary light. can benefit from a current overview of the evolution of
Given the central importance of sexuality to the evolu- human sexuality that integrates many of the disparate
tionary imperative—reproductive success—there is strands of work and that helps orient ongoing scholar-
strong theoretical impetus for understanding how evo- ship. If the field does not do this, others will fill that
lution has shaped human sexuality in the past, and for gap, leaving open the potential for best-selling but theo-
how the influence of past selective forces continue to retically and empirically distorted works such as Ryan
manifest in the present. For biological anthropologists, and Jetha’s (2010) Sex at Dawn to shape how people
interdisciplinary social and biomedical scientists, and think human sexuality evolved.
even wider audiences, questions about human sexual- A second reason for a review is new theoretical and
ity can be among the most compelling and meaningful, empirical work on the evolution of human sexuality. The
another reason to try informing answers to fundamen- theoretical foundations to sex differences in mating
tal questions of human sexuality with the most effort, for example, can also reflect demographic factors
scientifically supported and current views. An under- that had been less appreciated in previous theoretical
standing of human sexuality is central to topics as diverse formulations. New data from neuroimaging, hormonal
as predicting the dynamics of a sexually transmitted infec- studies, and genetics and genomics contribute to an
tion (STI) outbreak to the reasons why people pour so enriched understanding of the mechanisms of human
much of their time and resources into mating effort. sexual behavior. The availability of more recent sex sur-
Human sexuality, as much as any other topical focus vey data drawing upon large, international, and even
within biological anthropology, warrants ongoing evolu- Internet-based content helps provide new insights into
tionary scrutiny. the patterning of human mating and interest in sexual
For several reasons, this is arguably an opportune stimuli. How these new lines of evidence fit with other
time for providing a current overview of the evolution of lines of evidence and evolutionary theory is worthy of
human sexuality. For one, many of the foundational the- investigation. New theory and data could potentially
oretical and empirical touchstones in biological anthro- transform, or slightly modify, or even affirm earlier per-
pology that focus on the evolution of human sexuality spectives regarding the evolutionary foundations of
are dated. Darwin’s (1871) seminal contributions con-
tinue to shape our thinking regarding sexual selection,
sex differences, and evolutionary models of human mat- *Correspondence to: Peter B. Gray; Department of Anthropology,
ing. So too do important contributions from the 1970s University of Nevada, Las Vegas, 4505 Maryland Parkway, Box 455003,
and 1980s such as Symons (1979) Evolution of Human Las Vegas, NV 89154-5003. E-mail: peter.gray@unlv.edu
Sexuality and Hrdy’s (1981) The Woman that Never
Evolved. Of more recent and prominent works focused DOI: 10.1002/ajpa.22394
on the evolution of human sexuality, however, many Published online in Wiley Online Library
draw heavily from evolutionary psychology, such as (wileyonlinelibrary.com).
societies sampled, Betzig (2012) also found that males monogamous (lekking species such as peacocks and tur-
had higher variance in reproductive success than keys among those exceptions), it became clear that social
females. Betzig also showed that male reproductive skew and sexual relationships were not identical. These obser-
was most pronounced in socially stratified intensive agri- vations may have helped raise more questions about the
cultural societies such as the Inka. traits females sought in a prospective mate, including
Hanna Kokko and colleagues have also deepened our how this could give rise to mating with multiple part-
theoretical understanding of sexuality. She notes that ners (Clutton-Brock and McAuliffe, 2009; Geary, 2010).
actual, not potential, reproductive rates are what mat- Across primates, current theorizing suggests female
ters. That simple observation reminds us to consider the mate choice is oriented toward protection of herself and
availability of potential mates (just because males may her offspring. Protection may be useful against would-be
gain higher reproductive success than females if having predators, but also against would-be harmful males. In
more mates does not guarantee that mates are avail- species in which females mate with multiple males, such
able). Males have lower parental certainty than females, as chimpanzees and rhesus monkeys, Hrdy (1981) has
and fewer males tend to reproduce than females, factors suggested this represents paternity confusion, designed
that result in males gaining fewer benefits and higher to make all males be kinder toward the female and her
costs to parental care than females (Kokko and Jenn- offspring. For some species with long-term sociosexual
ions, 2003, 2008a). These processes may account for the bonds and paternal care, including primarily small-
origins of as well as positive reinforcement of sex differ- bodied South American owl monkeys, tamarins, and
ences in mating and parenting effort. Additionally, how- marmosets, females may benefit from male support but
ever, demographic factors play an important role in evidence is lacking that females use cues of male invest-
specifying the gradient of sexual selection (Kokko and ment (see Dixson, 2012). Females of various species may
Jennions, 2008b). As an example, if males are subject to also seek to mate with males providing complementary
greater mortality as a result of mating competition, then (e.g., in MHC system) or “good” (e.g., beneficial alleles
that lightens to some degree the mating competition the against local infectious disease ecology) genes. Since few
fewer, surviving males face. primate species have males that provide food resources
Such critiques have not overturned fundamental prin- (some callitrichid monkeys of South America exceptions)
ciples of sexual selection. Kokko’s work refines our theo- or have a sexual division of labor, there are no strong
rizing (e.g., to push from sexual selection to parenting nonhuman primate comparisons of females choosing
rather than from parenting to sexual selection), but ulti- males based on economic (e.g., food acquisition and pro-
mately anticipates sex differences in mating and paren- visioning) criteria.
tal care observed in the natural world. Additionally, Although Darwin emphasized female choice and male–
these kinds of critiques and extensions point to the male competition, it is recognized that female–female
importance of socioecological context and demography in competition and male choice occur (Low, 2000; Geary,
accounting for species and population differences in sex- 2010; Stockley and Bro-Jorgensen, 2011). Given that
ual selection pressures. In populations with more heav- female reproductive success is often closely tied to
ily biased sex ratios toward females, we might expect resources such as food, models of female–female competi-
more female–female competition and enhanced male tion, and cooperation feature reproductively relevant
choice (e.g., Guttentag and Secord, 1983). Research resources such as ripe fruits among chimpanzees or
shows that in many primate species, including humans, resource-bearing males in human agricultural societies.
female reproductive success is highly attuned to varia- Studies of nonhuman primates reveal that higher-
tion in energetic factors such as food availability. Indeed, ranking female social primates may benefit by having
related theoretical models help situate socioecological more surviving offspring and offspring whose reproduc-
and demographic considerations: Emlen and Oring tive careers are accelerated; this could be due to prefer-
(1977) pointed to the importance of resource distribu- ential food access often available to higher-ranking
tions in accounting for variable mating dynamics, and females (reviewed in Pusey, 2012). At the same time,
Mitani et al. (1996) suggested that the operational sex potential female rank-related variance in female repro-
ratio (ratio of breeding males:females) better represented ductive success should not be oversold; lower-ranking
the gradient of sexual selection than the actual adult females may engage in alternative foraging strategies,
sex ratio. All said, current sexual selection theory aims resulting in no net differences in reproductive success,
to account for the evolutionary past that shaped over- as is also frequently found in primate field studies
arching patterns of sexual selection among species (Pusey, 2012). In humans, much of the female–female
(including humans), as well as inform an understanding competition literature has highlighted competition over
of population variation in sexual selection pressures. acquiring and maintaining desirable mates (Campbell,
1999; Archer and Coyne, 2005; Geary, 2010). Further,
Female promiscuity, mate choice, and reproductive competition can lead to attempts to channel
female–female competition limited resources toward one’s own rather than another
women’s children, as observed with cowife competition
Part of the challenge to principles of sexual selection (Jankowiak et al., 2005). As for male choice, even in
theory stems from data on female mating patterns. multimale, multifemale species such as chimpanzees or
While Darwin and other theoreticians highlighted the rhesus monkeys, males may still attempt to channel
relevance of female choice, how that played out in the their mating effort toward more desirable females, such
natural world expanded theoretical imaginations. The as maximally fertile females; less fertile and experienced
advent of genetically based paternity testing among adolescent females may be relatively shunned as mates,
many avian field studies revealed that a sizable and for example (Manson, 2011; Muller et al., 2006). For
variable fraction of offspring were sired by individuals human resource intensive strategies, males may seek to
other than a social partner (Birkhead, 2000). Even allocate their limited mating effort toward more desira-
though over 90% of avian species tend to be socially ble mates; that can include preferences for younger
and in obtaining attractive females.” Darwin’s observa- compared with other times across the day when they
tions are consistent with many aspects of contemporary may be more fissioned (Martin Muller, personal commu-
theorizing concerning the evolution of male mating nication). Most sexual behavior was likely in a type of
strategies; they leave out female strategies, which can ventral-dorsal position (e.g., males from behind),
be better elaborated, as we shall see below. although bonobos and orangutans are recognized to use
In this section, I discuss evolutionary scenarios con- a wider array of sexual positions, especially compared
cerning hominin sexual behavior. Phylogenetic insights with other primates and mammals (Dixson, 2012). Based
are drawn from great ape comparisons, specifics of the on great ape comparisons, most matings likely took place
hominin fossil and archaeological record, and insights on the ground. If early hominin sexuality resembled that
into recently studied human hunter-gatherer societies, of chimpanzees and bonobos, then it would have entailed
with overarching theoretical guidance provided by sex- mating in multimale, multifemale groups. Further, early
ual selection theory and socioecological principles. These hominin females might have mated with many males,
scenarios will also later be connected with genetic and while also showing preferential mating access to a domi-
physiological data, contemporary international studies of nant male around the time she was most fertile.
human sexual behavior, and insights drawn from a life Females might have mated during parts of their preg-
course perspective on human sexuality. While there is nancy, but exhibited profound reductions in sexual
ongoing debate about specific evolutionary models of behavior when in a state of postpartum lactational
hominin sexuality, not surprisingly in light of the limita- amenorrhea (i.e., not cycling, but heavily lactating and
tions of available data, it is also worth underscoring that caring for an infant). Males may have competed heavily
the models are consistent with these latter bodies of evi- through alliance formation and contest competition with
dence, an indication that there is a coherent and syn- other males for social rank, in turn translating higher
thetic science here. rank into greater mating success. At this time, it is diffi-
cult to discern whether chimpanzees, some other ape
Early hominin and australopithecine sexuality such as gorilla, or no extant ape can serve as a strong
referential model for early hominin sexuality. Reasons
Current genetic and fossil evidence suggests that the for the lack of clarity include questions whether the mul-
closest living relatives of humans are chimpanzees and timale, multifemale mating system of chimpanzees and
bonobos (Goodman et al., 1998). Genetic data suggest bonobos (and related traits such as exaggerated sexual
that chimpanzees and bonobos themselves had a last swellings and heightened evidence of sperm competition)
common ancestor in Africa about 1 million years ago is derived or shared with a common ancestor; over
(Prufer et al., 2012). Humans had a last common ances- reconstructions of the degree of body size sexual dimor-
tor with chimpanzees and bonobos in Africa around 6 phism among early hominins and australopithecines;
million years ago. What was that ancestor like, and and a dearth of relevant genetic analyses that might be
what can we infer about its sexual behavior? As we shall able to address the evolutionary polarity of potential
see, there are clues to help answer these questions, but shifts in hominin sexuality.
there are also uncertainties. Most researchers suggest that australopithecines were
The earliest putative hominin fossils indicate body more polygynous than members of the genus Homo
sizes comparable to today’s chimpanzees and bonobos (McHenry, 1994). Debate continues on the number of
(Senut et al. 2001; Brunet et al., 2005; Lovejoy, 2009). gracile australopithecines and their phylogenetic rela-
The finds are all in Africa. The locomotor anatomy sug- tionship to earlier hominins and later members of Homo
gests the earliest hominins, as well as later gracile and (Wood and Lonergan, 2008). However, most reconstruc-
robust australopithecines, were semiterrestrial bipeds. tions of the postcranial remains of gracile australopithe-
They retained both adaptations for spending time mov- cines suggest they exhibited greater degrees of body size
ing in trees (e.g., long finger bones, relatively long arms, sexual dimorphism compared with modern humans and
and mobile shoulder joint) and for moving bipedally. most other members of Homo, and possibly even more
Most models suggest they moved bipedally on the dimorphism compared with chimpanzees and bonobos
ground (Lieberman, 2011), although it has also been sug- (see Plavcan, 2012). The available postcranial remains
gested that bipedalism emerged on tree branches during are scanty, are scattered across time and locations, and
foraging (Crompton et al., 2008). The environmental entail inferences regarding sex in the first place. Such
reconstructions of early and later pre-Homo hominins considerations have led other researchers to question
suggest they inhabited wooded environments. A few whether australopithecine body size dimorphism esti-
early hominins, such as a find at Chad dated to approxi- mates were exaggerated. It has been suggested the Aust.
mately 7 million years ago, appeared to have slightly africanus might have been less dimorphic than Aust.
larger canine teeth (compared with modern humans, but afarensis (Harmon, 2009). It has also been suggested by
less than great apes), though generally among hominins Reno et al. (2003) that gracile forms were only mildly
canine teeth are reduced in size compared with extant dimorphic—comparable with modern humans. If gracile
and presumably ancestral great apes. The shift toward australopithecines were highly dimorphic, the theoreti-
greater bipedalism may have relaxed selection on reten- cal inference would suggest they mated in one-male pol-
tion of larger canine teeth, particularly among males, for ygynous groups, perhaps somewhat similarly to gorillas
use in male–male contest competition; fighting may have (Geary and Flinn, 2001). If they were only mildly dimor-
relied, instead, on punching, grappling, or use of phic, that suggests, instead, they were mildly polygy-
weapons. nous, and perhaps mating in somewhat human-like
If early hominins were similar to extant great apes, ways from this earlier time. The robust australopithe-
these early hominins likely mated primarily diurnally cines, which most scholars suggest became evolutionary
(see Mitani et al., 2012). Among contemporary chimpan- side branches rather than serving as human ancestors,
zees, most matings in the wild occur early morning or appeared to be highly sexually dimorphic in body size.
late afternoon when chimpanzee groups are more fused Aust. robustus, from southern Africa, may have had
foundation, with subsequent layers of hominin economic Berndt, 1951; Shoskak, 1981; Hewlett and Hewlett,
and social behavior building on these. For example, a 2010; Marlowe 2010; Walker et al., 2011). Because the
pronounced hunter-gatherer sexual division of labor is socioecology of hunter-gatherers is thought to resemble
viewed as a more recent derivative of the preceding in some respects (e.g., small group sizes, family relation-
shifts in sexuality. Table 1 illustrates some leading mod- ships) that in which earlier forms of Homo sexuality
els concerning the evolutionary foundations of Homo evolved, hunter-gatherers can serve as analogies for
sexuality, featuring female perspectives, male perspec- more distant patterns. Of course, recently studied
tives, and commentary concerning the possible timing hunter-gatherers are not living fossils; they have differ-
and relevance of each (see also Quinlan, 2008). ent foraging tools than more distant ancestors (e.g.,
Marlowe, 2005), among other differences. Yet observa-
Hunter-gatherer sexuality tions and patterns among foragers offer insights that
otherwise are not accessible.
As another source of insight into the evolution of From recently studied hunter-gatherers, the most
human sexuality, we can draw upon patterns observed common marital system is slight polygyny (Marlowe,
among recently studied hunter-gatherers (Berndt and 2005; Walker et al., 2011). That is, among foragers, most
for the patterning of genetic data and how proximate Genetically based insights into the evolution of
mechanisms underpinning romantic and sexual behavior human sexuality
are expressed. Many of the recent physiological data on
human sexuality have been generated without explicit One of the most rapidly evolving regions of the human
evolutionary theorizing and interpretation, even though and these other hominids’ genomes involves male repro-
an understanding of proximate mechanisms and adapt- ductive genes (Clark and Swanson, 2005). Given that
ive function are complementary ways to investigate the the bottom line of evolution is reproductive success, it is
system (Tinbergen, 1963). not surprising that genes involved in spermatogenesis or
semen parameters might be rapidly evolving. As an
Recent genetic and genomic data example, the SEMG1 and SEMG2 genes, thought to aid
related to the evolution of the coagulation of semen, appear to have undergone a
human sexuality loss of function in gorillas and gibbons (which have low
sperm competition pressures), while being subject to
From the standpoint of genetics and genomics, several adaptive evolution in chimpanzees (which have high
approaches guide the application of new data to evolu- sperm competition pressures); “Interestingly, the obser-
tionary questions. One is a candidate gene approach, in vation that humans are in a sense intermediate in their
which tests for signatures of selection are undertaken to rates of seminal protein evolution between chimpanzees
determine whether a specific gene thought to be impli- and gorillas parallels that of anatomical and physiologi-
cated in some aspect of human sexuality shows evidence cal correlates of mating systems.” (Carnahan and
of this. As an example, the vasopressin 1a receptor Jensen-Seamen 2008: 946). However, a wider compari-
(AVPR1a) has been implicated in vertebrate male spe- son of the ejaculate components across humans, chim-
cies- and individual differences in mating and paternal panzees, bonobos, and gorillas found that effective
behavior; one report noted that humans and bonobos population size rather than sperm competition pressure
had a distinct AVPR1a compared with chimpanzees, appeared to best explain overall species differences in
raising the prospect of this genetic difference serving as genes expressed in ejaculate (Good et al., 2013). When
a possible contributor to species differences in male related analyses compared subsets of genes involved in
social behavior (Hammock and Young, 2005). However, different ejaculate functions, there was some evidence of
subsequent comparative genomic work has not impli- enhanced selection on genes involved in immune
cated selection upon this gene during hominin or homi- responses and protease, suggesting different components
nid evolution. Another approach is to study genetic of the semen may evolve at different rates. Good et al.
information from other forms, such as herpes or HIV, to (2013) also note that the genetic bases underlying spe-
infer possible evolutionary-related scenarios regarding cies differences in human and African ape male repro-
sexually transmitted disease. Still another is to draw ductive function may rely more on gene expression than
comparisons of genomic sequences (or even entire protein differences and that gamete differences may be
genomes rather than a single or several genes) between more important than semen components.
humans and other species, between human populations, Differences between human androgen receptor and
or within individuals of a human population in order to that of primates have also been noted (Mubiru et al.,
identify genetic differences that might be related to sex- 2012). Humans have more CAG repeats in the androgen
ually relevant phenotypic differences. In a related vein, receptor promoter region than other primates although
quantification of sex-specific genetic contributions to a it is not clear whether the pattern represents phyloge-
population can inform an understanding of mating netic inertia or lineage-specific selective effects. A 2011
dynamics (e.g., specifying the relative variance in male paper noted a deletion upstream of the human androgen
and female reproductive success: Heyer et al., 2012). As receptor after comparison with other primates including
an example, relatively less male genetic variation was chimpanzee (McLean et al., 2011). In vivo work suggests
found among the Yoruba in Nigeria compared with a that the genetic deletion results in loss of penile spines
Chinese sample, consistent with recognized differences situated on the glans penis. Further, Neandertal has the
in marital behavior (Labuda et al., 2010). same version as humans (Hawks, 2011), suggesting this
The availability of a draft human genome in 2003, and deletion may have occurred among a common ancestor,
of the chimpanzee in 2005, has helped fuel interest in possibly in association with some derived aspect of homi-
the growing field of comparative genomics (Chimpanzee nin mating. As to other genetic insights into hominin
Sequencing and Analysis Consortium, 2005). The avail- sexuality, Gentry et al. (1988) note that oral and genital
ability of orangutan, gorilla, and bonobo genomes now herpes strains have separated long ago and remained
means that the genomes of all great ape species can be that way, suggesting that oral sex may not have
compared to discern similarities and differences, and in occurred regularly among our ancestors (since those
turn raise questions about the genetic bases to species strains of herpes would have merged rather than
differences in sexuality. The publication of draft genomes remained distinct).
from Neandertal and Denisova extends the comparisons From a candidate gene approach, some work has
to two extinct and closely related hominins (Disotell, investigated potential human between- and within-
2012). Given that Neandertals and Denisovans are appa- population genetic differences related to sexually rele-
rently sister groups, and that modern humans, apart vant phenotypic differences. Population differences in
from a small degree of population-specific interbreeding dopamine receptor (DRD4) genetic polymorphisms exist,
with them, last shared common ancestry with Neander- with possible explanations advanced to account for those
tals and Denisovans around 600,000 years ago, compari- differences including genetic drift, adaptive migration,
sons with these extinct hominins can help shed light on and mating behavior (Harpending and Cochran, 2002).
the phylogenetic structure of hominin sexuality. What A U.S. study found that polymorphisms in this same
insights have emerged thus far in such comparisons system were associated with differences in sexual prom-
(O’Bleness et al., 2012)? iscuity in a university sample (Garcia et al., 2010).
partnering and other hormones such as cortisol have levels acutely increase in response to sexual stimuli
also been investigated. In one oxytocin study of 38 cou- (reviewed in Archer, 2006; Van Anders and Watson,
ples, Grewen et al. (2005) found that blood oxytocin lev- 2006). In studies of solitary masturbation, men’s testos-
els were higher throughout a 10 min behavioral session terone levels do not consistently change, whereas expo-
among both women and men reporting greater partner sure of men to audiovisual sexual stimuli more reliably
support. In an experimental study in which women and increases testosterone levels. In a naturalistic study of
men were either administered intranasal oxytocin men’s testosterone responses conducted at an adult club,
(which is known to exert effects in the brain) or intra- men’s testosterone levels increased more when engaging
nasal placebo spray, those given oxytocin showed rela- in rather than observing sexual behavior (Escasa et al.,
tively higher ratios of positive to negative partner- 2011). In other studies, among men who masturbated,
directed behavior during a behavioral interaction session prolactin levels did not change (Kruger et al., 2003),
(Ditzen et al., 2009). Studies of both young and older whereas in men and women who engaged in partnered
couples’ blood cortisol levels have been conducted in intercourse their prolactin levels showed acute increases
relation to marital discussion, a paradigm in which cou- (reviewed in Bancroft, 2005). In both men and women,
ples report to a lab and are asked to talk about topics oxytocin levels tend to increase during sexual behavior;
known to elicit conflict (such as sex or finances) (Kiecolt- it may be that oxytocin is involved in smooth muscle
Glaser et al., 2003). As one of the main findings from contraction associated with orgasm as well as the
these studies, women in more positive marital relation- rewarding experience of sexual behavior (Carmichael
ships had rapidly reduced cortisol profiles during the et al., 1987). Studies have typically found no changes in
discussion compared with women in less positive marital women’s estradiol, but in some cases increases in wom-
relationships (Robles et al., 2006). en’s testosterone levels during sexual behavior (see Van
Among studies investigating male testosterone levels Anders et al., 2009). Summarizing much of this litera-
and partnering, nearly all conducted in North America ture on hormonal responses to sexual behavior, major
have found that men involved in committed, romantic features of the system are homologous across a wide
relationships have lower testosterone levels than their phylogenetic scope, and both sex similarities and differ-
unpartnered counterparts (Gray and Campbell, 2009). A ences have been recognized.
longitudinal study of U.S. military veterans showed that
men’s testosterone levels increased after divorce, provid- Human reproductive physiology and anatomy in
ing evidence that changes in relationships can precede comparative context
changes in men’s testosterone levels (Mazur and Micha-
lek, 1998). However, international studies have been less Closing out this section on physiological data, several
consistent in findings linking men’s testosterone and recent syntheses warrant attention. One is Alan Dixson’s
relationship status. In most international studies, (2009) compilation of human reproductive physiology
becoming a father in a social context in which men pro- and anatomy placed within an explicitly comparative
vide paternal care is associated with lower testosterone perspective, and the conclusion from multiple lines of
levels, but a number of studies found no difference in evidence that humans bear signatures of low sperm com-
the testosterone levels between partnered and unpart- petition pressures. While classic comparative research
nered childless men. As an example, in a sample of 126 on relative primate testis size reached a similar conclu-
men aged 21–38 in Beijing, fathers had lower testoster- sion (Harcourt et al., 1981), the earlier work had a
one levels than both partnered childless men and single human sample size of four individuals from which tes-
men, with the testosterone levels of partnered men with ticle size was measured in contrast to over 7,000 men in
and without children similar (Gray et al., 2006). A longi- more recent analyses. Dixson summarizes data indicat-
tudinal study of several hundred men in Cebu City, Phil- ing humans have low sperm quality, ejaculate volume,
ippines, revealed that becoming a father, particularly of sperm reserves, modestly sized seminal vesicles, and
infants and associated with greater paternal care, prostate gland, and relatively small sperm midpiece vol-
caused men’s testosterone levels to decrease (Gettler ume. Human females also have relatively short oviduct
et al., 2011). Four studies have also investigated men’s length (i.e., Fallopian tube), consistent with relatively
testosterone levels with respect to polygynous marriage. low sperm competition pressures. This conclusion is at
In the first of these, conducted among Swahili men in odds with views espoused in Baker and Bellis (1994) or
Kenya, men married to two wives had higher testoster- Ryan and Jetha (2010). The evolutionary significance of
one levels than other men, whether monogamously mar- population and individual human variation in these and
ried or unmarried (Gray, 2003). The interpretation of related reproductive physiological and anatomical
these and the several other studies suggest that age and parameters, however, remains unclear. As examples, dif-
social context matter, and it is also difficult to disentan- ferences in human sperm counts (Dixson, 2009) and
gle potential confounding influences such as develop- female vaginal flora (Ravel et al., 2011) have been noted.
mental impacts of diet and physical activity. That variation helps provide a basis on which selection
A growing number of studies have investigated hormo- can act, but it is not clear if observed population differ-
nal responses to sexual behavior, with emphasis upon ences in, for example, testis size can be traced to
intercourse or solitary masturbation culminating in population-specific selective pressures rather than other
orgasm. Some of these studies have become more techni- evolutionary or developmental processes.
cally feasible with the use of minimally invasive mea- The way in which genital responses align (or not) with
surement of hormone concentrations from saliva, subjective sexual arousal tends to differ between males
whereas assessments of peptides typically rely upon and females. Through measurement of vaginal blood
blood draws. Some of the effects appear sex-specific, flow or penile tumescence in response to exposure to
whereas others, involving peptide release, appear simi- audiovisual stimuli, meta-analysis reveals that women’s
lar between females and males. Among males, a consid- responses less consistently align (r 5.26) with subjective
erable literature has tested whether men’s testosterone reports of arousal compared with men’s responses
may be temptations to downplay (e.g., affairs) or exag- TABLE 2. Frequency of sex in the past year from early 1990s
gerate (in the attempt to impress others) claims? Some U.S. national sex survey
cross-checks are feasible, such as aligning data between
A few 4 or more
partners or the sexes (e.g., regarding coital frequency Age Not times A few times 2–3 times times
within partnerships) or other sources of clinical or com- group at all per year per month per week a week
mercial information (e.g., patterns of STIs and knowl-
edge of contraception use should be consistent with Men
reports of sexual behaviors). The appropriate caution is 18–24 15 21 24 28 12
thus warranted when evaluating data on human sexual 25–29 7 15 31 36 10
30–39 8 15 37 23 6
behavior. 40–49 9 18 40 27 5
50–59 11 22 43 20 2
Sex in large U.S. surveys Women
18–24 11 16 32 9 12
From Ed Laumann and colleagues’ (1994) study of 25–29 5 10 38 37 10
U.S. sexuality, approximately 5,000 individuals aged 18– 30–39 9 16 36 33 6
59 years participated in in-person interviews. Outcomes 40–49 15 16 44 20 5
included frequency and types of various sexual behav- 50–59 30 22 35 12 2
iors, and were provided with respect to sociodemographic
variables including age group, sex, sexual orientation,
ethnicity, and educational attainment. Data on the fre- across the sample; age-related patterning in sexual
quency of sexual behavior the past year are presented behaviors showed that masturbation was relatively more
with respect to age group and sex in Table 2. As shown common early and late in adult life, while vaginal sex
in this table, the percentage of women and men report- relatively more common during the prime reproductive
ing having sex four or more times the past year tends to years; and for both men and women partnered sexual
diminish with age, while the percentage of women and behaviors declined in frequency in the 50s onward,
men not having sex the past year is highest among showing even steeper declines around the 70s.
young and old age groups.
This same U.S. survey provided a quantitative snap- A quantitative cross-cultural approach
shop of other features of adult sexuality. Across all age to human sexuality
groups, men engage in more frequent masturbation than
do women. As an example, at ages 18–24, 29% of men Apart from U.S. studies, we also want to know about
and 9% of women reported masturbating at least once a the wider cultural record of human sexuality. What can
week. Masturbation rates also declined with age, with be said about the sexual behavior of people around the
10% of men and 2% of women aged 55–59 reporting world? Charles Darwin himself tried to collect as much
weekly masturbation. Men reported more often thinking information on mate preferences and other features of
about sex and using autoerotic material such as watch- sexuality as he could by compiling the observations of
ing erotic videos; for example, 54% of men and 19% of travelers and others with international experience.
women reported thinking about sex at least daily. Extra- Those cross-cultural anecdotes are included in his 1871
marital affairs were noted among a minority of respond- book. As far as more rigorous cross-cultural study of
ents: “Over 90 percent of the women and over 75 percent human sexuality, the standard bearer has been Ford
of the men in every cohort report fidelity within their and Beach’s (1951) Patterns of Sexual Behavior. This is a
marriage, over its entirety.” (Laumann et al., 1994: 214). compilation of cross-cultural accounts drawn from the
Men reported a higher frequency over various time human relations area files (HRAF). In Ford and Beach’s
frames engaging in same-sex sexual behavior than did book, sections are devoted to a variety of sexual prac-
women with, for example, 2.0% of men and 0.8% of tices, from subadult sexuality to adolescence, and adult
women specifying having sex with same-sex partners sexual behaviors ranging from intercourse positions to
over the past five years. With respect to sexual identity, extramarital affairs. Variation is recognized and in some
2.8% of men and 1.4% of women reported either bisexual cases patterned; for example, longer taboos against post-
or homosexual identity. Women (22%) were much more partum female sexual behavior are found in polygynous
likely to report being forced to have sex than were men societies, consistent with the idea that a wife with cow-
(1%), with most of women’s forced sex occurring with ives can abstain longer from sex after having had a baby
known individuals such as someone with whom the when there is another sexual partner available to her
woman was in love or someone she knew well. husband. Additionally, Ford and Beach concluded that
More recently, a nationally representative probability cross-cultural standards of attractiveness varied,
sample of U.S. sexuality was conducted in 2009 by schol- although female appearance garnered more attention
ars based at Indiana University, thus continuing Kin- than male appearance, and male social success often
sey’s legacy of work there. This study relied upon mattered more than attractiveness in determining male
computer-aided survey. A sample of 5,865 men and marital and reproductive success. In this section, we
women aged 14–94 participated. Findings from the sur- move beyond Ford and Beach’s book to present more
vey were published in a special edition of the Journal of recent international studies of human sexual behavior.
Sexual Medicine in 2010. The study presented data on Some of these more recently published studies are deriv-
various sexual practices such as intercourse, masturba- atives of the HRAF data set, while others rely on nation-
tion, and oral sex with respect to age, sex, and a host of ally representative probability samples of sexual
social variables including sexual orientation, ethnicity, behavior from various countries around the world.
education, income, and geographical region. Among The most widely used quantitative measures of cross-
some of the take-home findings from the study, males cultural sexuality tend to rely upon marital patterns.
engaged in higher rates of masturbation than females Societies are variously described as monogamous, with
years) and Mali (13.2 years). The age of first intercourse with respect to sex (male/female), sexual orientation and
varies globally. In some countries, women have an ear- age, among other factors, while also providing a window
lier sexual debut and higher intercourse rates compared into cross-cultural variation. Sex differences in behav-
with men; examples include a number of west African ioral measures of desire such as masturbation or use of
countries and Bangladesh, in which women marry at prostitution are quite robust across these samples, con-
younger ages and the pattern reflects having sex within sistent with theory that males have evolved a more pro-
marriages. The prevalence of premarital sex among ceptive sexuality. Males engage in higher rates of same-
women is highest among countries such as the U.S. and sex sexual behavior than females in these large U.S. and
in Europe, but quite low in other countries such as Mali, international data sets. Age-related declines in various
Nepal, and Indonesia. In the latter cases, less than 25% measures of sexual behavior, such as coital frequency,
of women report engaging in intercourse before mar- also conform to waning selective pressures to maintain
riage. In other countries, such as Brazil, men tend to sexual function, a topic we will also turn to in the next
have younger ages of sexual debut than women, with section on human life course variation in sexuality. Most
this reflecting higher rates of male premarital sex, sexual behavior occurs within the context of long-term
including in some cases with prostitutes. Data compiled bonds, even when humans today face an unprecedented
by Wellings and colleagues only tabulate use of prosti- array of novel potential mating partners compared to
tutes by men, a recognition that prostitution is largely hunter-gatherer forebears. Yet the international data
an industry servicing men; most male prostitutes service also indicate that, at the societal level, slight polygyny is
other men rather than women. Yet the percentage of a better descriptor of human marital patterns than
men who say they paid for sex in the past year—a mea- monogamy. The international data also underscore the
sure of use of prostitutes—varies around the globe. importance of contextual variables such as fertility rate
Higher rates are found in China and East and West and OSR. As an example, the societies with higher fertil-
Africa (around 10%) than in Latin America, North Amer- ity rates tend to have more restrictive attitudes toward
ica, Europe, and North Africa (less than 3%). female sexuality. The Internet provides a new window
into human sexuality, showing that some of the long-
Sex on the Internet standing patterns such as sex differences in sexual
desire manifest in this new arena. The collective weight
New sources of data on human sexuality continue to
of this body of evidence argues for the continued rele-
arise. The Internet is one opening novel avenues by
vance of evolutionary theory to understanding human
which measures of human sexuality can be obtained and
sexuality today, and presents a standard against which
interpreted. This is reflected in multiple ways, including
competing explanations are challenged to explain.
the diversification of websites catering to all kinds of sex-
ual and romantic interest groups. Online dating has
become a burgeoning industry and an increasingly com- LIFE COURSE PERSPECTIVE ON HUMAN
mon means by which partners in countries such as the SEXUALITY
U.S. meet. The quantitative analysis of search contents
The vast amount of theoretical and empirical attention
has also opened up fresh ways to explore human sexual
to human sexuality from an evolutionary perspective
expression. The most ambitious and current account to
has focused on prime adult years. A large literature
quantitatively explore sexuality related Internet search
investigating and adaptively interpreting sex differences
contents was published in Ogas and Gaddam’s (2011) A
in mating strategies has, for example, tended to empha-
Billion Wicked Thoughts. They report the results of over
size young adults who do not have children. In this sec-
400 million web searches from more than 2 million users,
tion, a life course perspective is applied to human
primarily from the U.S., but also other countries such as
sexuality. This is important for a number of reasons: sex
the U.K., India, Nigeria, and Canada. They analyzed
differentiation of brains and genitals begins before birth;
search data from 2009–2010 as well as a supplementary
juveniles learn and practice sociosexual behaviors (in
AOL search database on more than 650,000 users. Much
addition to economic and social skills emphasized by evo-
of their effort honed in on potential differences in search
lutionary thinkers) that may be suited to their context
content according to sex and sexual orientation.
and future reproductive success; puberty and adoles-
Summarizing what they found, “On the Web, men pre-
cence transition individuals into the raised stakes of
fer images. Women prefer stories. Men prefer graphic
potential reproduction; features of human sexuality fluc-
sex. Women prefer relationships and romance. This is
tuate across the prime adult years, including across peri-
also reflected in the divergent responses of men and
partum transitions; and features of human sexuality
women when asked about what sexual activities they per-
change with age in ways that can be understood with
form on the Internet.” (p. 19) Men were much more likely
respect to evolutionary-based models of reproductive
(37%) than women (6%) to list viewing erotic images and
senescence.
movies as preferred Internet sex-related activities. Con-
Life history theory provides a theoretical and empiri-
versely, women (21%) were more likely than men (8%) to
cal toolkit to make sense of the patterning of human sex-
report sex-related activities on the Internet involving
uality across the life course (Stearns, 1992; Geary, 2002;
attempts to maintain contact with love or sex partners.
Hawkes and Paine, 2006; Muehlenbein, 2010). Individu-
Of other patterns in Internet sex searches, younger
als have limited time and energy to devote to competing
women (age 16–18) draw the most interest, consistent
allocations of growth, maintenance, and reproduction.
with male orientation toward cues of high reproductive
The specific allocations to these agendas can exhibit age-
value and pursuit of a long-term mate.
, sex-, and context-specific variation in ways that may
Summary make adaptive sense. There may also be tradeoffs across
these allocations, meaning that allocation to one out-
A variety of large U.S. and international quantitative come comes at expense to another. To illustrate, based
sexuality data are available. These data reveal patterns on the theory reviewed above, female mammals tend to
Life course stage Key adaptive and physiological processes Key patterns of sexuality
Perinatal Early organizing effects on brain and Sex-specific developmental trajectories begin
differentiation of primary reproductive to unfold
characteristics Sex-specific and plastic mechanisms orient
toward social stimuli to enhance survival
Early Childhood Dependency on mothers and other Interest in own and others’ genitals, includ-
caregivers for social and energetic ing auto-stimulation
support Some degree of sex differentiation in social
behavior
Middle Childhood Greater independence and interaction with Expanded distinctions in sex differentiation,
other adults and juveniles with females more oriented toward care-
giving contexts and males more oriented
toward physical play
Increasing frequency of sex play, especially
for males, in same-sex and between-sex
pair-bonding contexts
Puberty and Adolescence A second phase of organizing effects of Shift into potential reproductive realm
steroid hormones on the brain, and Females tend to be 1–2 years ahead of males
development of secondary sexual Marking transition toward social, economic,
characteristics and political activities conducive to repro-
ductive success, with female transitions
highlighting aspects of fertility and male
transitions highlight social achievement
Increased frequency of intercourse, but also
quite variable by individual, sex,
population
Adulthood Age-related changes in reproductive Age-related changes in sexuality during
physiology adulthood can be related to reproductive
partnerships
Subtle physiological changes across the Cyclical female shifts across the ovarian
ovulatory cycle and dramatic changes cycle have at best subtle influences on
across the reproductive cycle sexuality, whereas shifts across pregnancy
and postpartum phases have pronounced
impacts on female sexuality
Male sexuality is typically intertwined with
that of a long-term partner, although sex
differences in desire are amplified during
late pregnancy and early postpartum
Senescence Age-related declines in reproductive The menopausal transition in women is
function, including decreases in sex associated with some declines in sexual
steroids function
Measures of sexual behavior decline with
advancing age in both women and men,
although in sex- and context-specific ways
devote more of their reproductive effort to parenting children have a limited sexual capacity, but a capacity
effort, whereas male mammals tend to devote more of that can foster engaging in the kinds of sociosexual play
their reproductive effort to mating effort. The shifts that may eventually attune their sex- and context-
from growth to reproductive effort at puberty entail specific reproductive investment. To help capture some
development of secondary sexual characteristics; by of the key processes and patterns of human sexuality
waiting until puberty to develop these characteristics, across the life course, from the perinatal period onwards,
organisms delay the potential social and energetic costs these are summarized in Table 3. I consider some of
of their maintenance. Peripartum shifts in female socio- these major processes and patterns from early to late in
sexuality, which can also impact the sexuality of a long- the life course, touching on some of the most common
term partner, can be viewed from the standpoint of life changes in sociosexual behavior, underlying physiologi-
history tradeoffs between investment in current and cal processes, and with some reference to life history
future reproduction. theory.
Rigorous, quantitative studies of childhood sexuality
The early development of human sexuality are fraught with ethical and methodological challenges,
even more than research on adults. Thus, the available
As an unusual anecdotal insight into the development data are quite limited, and the studies conducted by
of human sexuality, an ultrasound taken of a male fetus Alfred Kinsey and colleagues in the mid-20th century
appeared to show him engaging in autostimulation (mas- still stand as pillars in the field. What Kinsey et al.
turbation), an indication that sexuality can precede birth learned, primarily through interviews conducted with
(Meizner, 1987). Female infants are capable of vaginal adults who recalled their childhood sexual experiences,
lubrication, and male infants of erections. Infants of was that children’s sexual behavior was patterned with
both sexes are capable of orgasm, though without ejacu- respect to age and sex. By around age 5, approximately
lation. These observations indicate that even young 10% of girls and boys reported having engaged in any
sex play. Across aged 5 through 13, however, that fre- societies, women and men had similar regulations gov-
quency remained comparable for girls, but increased erning sexual behavior, whereas among more restrictive
steadily to around 35% for boys aged 11–13. A fraction societies men’s sexual activities were sometimes less con-
of these experiences were same-sex: while 8% of girls strained than women’s. This yields a sexual double
aged 5–7 and 3% of girls aged 11 reported engaging in standard among some societies. A variety of factors
homosexual play, 5% of boys aged 5–7 and about 30% of underlie cross-cultural differences in these aspects of
boys aged 11 reported in homosexual play. More recent adolescent sexuality (Schlegel and Barry, 1991). Some of
quantitative studies of children’s sexuality have been the most important factors are residence (matrilocality
conducted in the U.S. and Europe, but typically rely on associated with more permissiveness, patrilocality with
parent or doctor reports rather than reports by children less) and inheritance and descent (greater permissive-
or adult retrospective reports. These latter studies ness associated with bilateral descent and little inheri-
underestimate frequencies of childhood sexual behaviors tance, but more restriction in patrilineal descent).
that children intentionally shield from adult view, but Schlegel and Barry (1991) also discuss cross-cultural
nonetheless can offer some useful insights. For example, patterns of same-sex adolescent sexual behavior. If
one study found that masturbation frequencies in the restrictions govern same-sex sexual behavior of one sex,
U.S. increased from around 10% by age 7 to around 80% those are often found in the other sex too. If adolescent
by age 13 (Mallants and Casteels, 2008). males are subject to same-sex sexual restriction, they
As a complement to quantitative studies, ethnographic are often subject to similar restrictions through adult-
reports of childhood sociosexuality provide insight into hood. While a fraction of men hold lifelong same-sex pri-
age-, sex-, and context-specific patterns. Eleanor Mac- mary attractions, adolescent male same-sex sexual
coby (1998) summarizes research on U.S. children’s sex- behavior is often a life history phase resulting from a
uality, noting that by around ages 7–9 children begin to lack of mating access to females.
conceptualize cross-sex interactions with romantic or Quantitative studies of adolescent sexuality are avail-
sexual overtones. Herdt and McClintock (2000) suggest able for the U.S. and many other countries, complement-
that the timing in the U.S., New Guinea, and elsewhere ing the qualitative cross-cultural compilations. In the
of children developing sexual attractions to same- and U.S., about 50% of secondary school students report hav-
other-sex partners around age 10 may be related to ris- ing engaged in intercourse, with no notable sex differen-
ing concentrations of adrenal androgens as part of adre- ces, although frequencies vary across reported ethnicity.
narche. Ford and Beach (1951) parse cross-cultural Approximately 25% of these students also report more
variation in childhood sexuality into three broad catego- than one sexual partner. To gain insight into quantita-
ries: restrictive, semirestrictive, and permissive societies. tive international adolescent sexual behavior, we can
They note that the varied degree of proscriptions guiding return to the Wellings et al. (2006) review previously
childhood sexuality tend to be aligned with adult sexual- noted. From this review, one of the central findings is
ity; in other words, children are variably granted sexual that in countries with young age of female marriage,
latitude or constraint in ways consistent with the adult females typically engage in earlier sexual intercourse
sex lives they will later have. Several hunter-gatherer than males, often within marriage. As another pattern,
ethnographic anecdotes recognize that sex play also in developing countries the duration between sexual ini-
occurs, as among the Hadza of Tanzania in which Frank tiation and marriage is shorter among females than in
Marlowe (2010) noted boys and girls “playing house” and males, meaning that female sexual debuts tend to be
engaging in sex play by around age 7–8. more closely tied to marriage than are male sexual
debuts. However, this is not the case in developed coun-
Puberty and adolescence tries such as the U.S. or Australia, in which both males
and females tend to engage in premarital sex that is not
As childhood wanes, puberty and adolescence begin closely tied to marriage.
(Bogin, 1999; Weisfeld, 1999). This is a time of shifting From an evolutionary perspective, some of the most
life history priorities. Investment in growth wanes and profound impacts on adolescent sexuality entail a
maintenance efforts can be compromised as allocations lengthened duration of adolescence, especially for
to reproductive effort increase. Females typically under- females. Among hunter-gatherer societies, females tend
take the pubertal transition 1–2 years before males, a to experience menarche around age 15–16, with mar-
sex difference consistent with slight polygyny (Dixson, riage and reproduction following a few years later (Kelly,
2012). Sex steroids exert a second stage of organizational 1995; Ellison, 2001; Howell, 2010). The brief phase of
effects on the brain, serving as physiological mecha- adolescence among female foragers entails infertility or
nisms shaping these shifting priorities. Sex steroids also subfecundity as ovarian cycles are less regular and often
regulate the expression of secondary sexual characteris- nonovulatory, helping make possible sexual experimen-
tics. Effectively, the increased estrogen released by the tation (if not socially proscribed) (Montague, 1946). For
ovary and androgen by the testis shape various anatomi- hunter-gatherer males, pubertal milestones are more dif-
cal and physiological characteristics in ways that ficult to measure (such as “nocturnal emissions” or
enhance reproductive success in sex-specific ways. As changes in testis and penis size), although male pubertal
examples, females develop greater gluteofemoral fat, transitions (like in other societies) typically begin later
which can serve as a metabolic reservoir on which to and extend considerably longer. Males must demonstrate
draw during gestation and lactation, and males put on their social and foraging prowess, perhaps through bride
more upper body musculature, thought to aid ancestral service, to show they are worthy of marriage. A conse-
male success in mating competition. quence is that in foraging societies males are commonly
The impacts of the pubertal transition on human sex- in their mid-20s before marrying and becoming fathers.
uality are context-specific. Returning to Ford and Beach Against a hunter-gatherer backdrop, however, in
(1951), their scheme recognized restrictive, semirestric- many contemporary contexts females and males are
tive, and permissive societies. Among permissive undergoing earlier puberty and postponing the
documented cross-culturally (e.g., Ford and Beach, 1951) erning women’s sexual behavior vary in length; in one
and in many other studies. A parsimonious comparative compilation, taboos lasting 1–5 months were typical in
interpretation of these cycle-related patterns in human Eurasia and the Mediterranean region, but taboos
female sexuality is that they are consistent with the lack around 2 years in length more common across many
of selection on female multimale mating (unlike chimpan- sub-Saharan African societies. A key factor in variable
zees, for example: Nunn, 1999), while also providing sup- postpartum female sexual taboo lengths is whether a
port for subtle increases in female proceptivity, receptivity mother’s partner—typically the infant’s father—has
and attractivity when fertilization is most likely. another sexual outlet. In polygynous societies, the father
of an infant may have another partner (e.g. a mother’s
Peripartum fluctuations in human sexuality cowife), allowing for longer taboos. In the same kinds of
demographic reports on women’s recent sexual behavior,
Any ovulatory cycle-related changes in female sexual- Brewis and Meyer (2005b) summarized the data across
ity pale in comparison to those occurring across the 19 countries such as the Philippines and Kenya, showing
reproductive cycle, or transition from life history states dramatically lower frequencies of female intercourse
of cycling to pregnancy to postpartum amenorrhea (not during the early postpartum phase. Similarly, Von
menstruating) and eventually cycling. From the stand- Sydow (1999) observed, in the same review as noted
point of life history theory, women’s changes in sexuality above for gestational sex, that, “Compared with the pre-
across the reproductive cycle can be viewed as tradeoffs pregnancy period, coital frequency is reduced in most
between current and future reproduction (Escasa-Dorne couples during the first year after birth.” (p. 36) In this
et al., 2013). Sexual behavior may be high during times same review, a fraction of men’s sexual desire remained
cycling, possibly leading to conception and successful lower during the postpartum period, though postpartum
reproduction. Sexual behavior may occur at variable paternal sexuality has been poorly studied relative to
rates across pregnancy, and for functions such as rela- that of postpartum women.
tionship maintenance (to recruit an investing partner From an evolutionary and life history perspective, it is
such as a child’s father) or as a byproduct of physiologi- during late pregnancy and the early postpartum phase
cal shifts arising during pregnancy (such as elevated when sex differences in sexual desire are at their maxi-
estrogen concentrations). Sexual behavior tends to mum during the reproductive years. That is an impor-
diminish after having had a child, as a mother focuses tant consideration because it is rarely considered in
her reproductive investment on keeping alive and aiding discussions of sex differences in sexuality, including sex-
the current reproductive product (baby) rather than ual desire (see Baumeister et al., 2001). It is also an
attempts to beget the next one. Because of the typical important contributor to potential sexual conflicts of
links between male and female sexuality in long-term interest, whereby mothers may be adaptively shifting
partnerships, many of the shifts in female sociosexuality life history investments toward a current reproductive
across the reproductive cycle impact male sexuality too. bout (infant), but fathers’ investments represent a
Cross-cultural attitudinal and international quantita- greater mixture of parenting and mating effort. Other
tive studies reveal that many women report being sexu- considerations can be at play too during this postpartum
ally active during pregnancy, but with additional period: the duration of female lactational amenorrhea
patterns across pregnancy and social context. In Ford varies considerably across and within populations, and
and Beach’s (1951) cross-cultural survey, they found that in relation to central energetic influences; that variation
among small-scale societies 70% approved of female sex- can impact a women’s physiology in relation to orienting
ual behavior during early pregnancy, but that those toward investment in current or future reproduction
rates of approval diminished across gestation. A primary (e.g., shorter durations of amenorrhea and more rapid
reason for disapproving of pregnant women’s engaging resumption of ultimately functional cycles can promote
in sex, especially during late gestation, was concern over female sexuality). The availability of a partner and other
the well-being of the fetus. Among more polygynous soci- allocaregivers can also be important; mothers without a
eties, sex during pregnancy was disapproved of earlier desirable partner may be motivated to more rapidly shift
in gestation; this is consistent with the sexual availabil- toward enhanced mating effort. Above all, from an evo-
ity of an alternative sexual outlet (cowife) to a husband lutionary perspective, studies of nonhuman primates
and would-be father, also lessening any pressures for and human hunter-gatherers recognize that the bulk of
pregnant women to engage in sex for a relationship female reproductive years were spent pregnant or in a
maintenance function. In Brewis and Meyer’s (2005b) state of lactational amenorrhea. The reproductive cycle
compilation of recent sexual data from more than 90,000 shifts in female (and male) sexuality summarized here
women from 19 countries such as Bolivia and Cameroon, can be viewed as central to ancestral adult sexuality in
they found that pregnancy impacted sexuality: addition to being of relevance to society today. In other
“Pregnancy has a negative effect for the most part on words, the sexual lives of our recent ancestors were
sexual frequencies. . .[G]enerally, coital frequencies are likely characterized by shifts across pregnancy and post-
significantly lower in pregnancy compared with the non- partum phases throughout the primary adult reproduc-
pregnant state. . .[O]verall. . .sexual frequencies decline in tive years.
the first trimester, decrease further in the second, and
still further in the third.” (p. 508) Similar patterns were Aging and human sexuality
observed in a meta-analysis of 59 sexuality studies
largely conducted in North America or Europe, with Human reproductive senescence proceeds, marking
measures of sexual frequency initially little different the the last major stage from a life course perspective in
first trimester from a nonpregnant state, but decreasing which to situate human sexuality. While considerable
across gestation (von Sydow, 1999). attention has been given to the evolution of human
Many studies find that measures of women’s postpar- senescence (loss of function with age) (e.g., Williams,
tum sexuality are diminished. Cross-cultural taboos gov- 1957), with discussions regarding grandmothering
pressures. The phylogenetic origins of these long-term example, postpartum women’s sex taboos, may still be
bonds, by most accounts, appear to trace to the genus drawn upon. Biological anthropologists are well-suited to
Homo, with ongoing debate about the most likely timing conceptualizing specific facets or cases of human sexual-
of shifts with earlier Homo vs. Homo heidelbergensis vs. ity within evolutionary and life history theory. Just as
Homo sapiens. At the same time, the expression of Charles Darwin long-ago showed us, the most integra-
human sexuality exhibits considerable within- and tive and robust understanding of human sexuality is
between-population variation. Rates of polygynous mar- rooted in an evolutionary perspective.
riage, affairs, masturbation and other features of sexual-
ity vary, which underscores the point that sexuality ACKNOWLEDGMENTS
needs to be placed within its social context. Such an For comments on an earlier draft, I thank Kali Bertel-
observation argues against any simplistic and determin- sen, Bill Jankowiak, Matthew Martinez, Timothy McHale,
istic view of the interaction between the heritable basis Michael Moncrieff, David Puts, Shelly Volsche, and Sharon
of sexuality and socioecological context. Furthermore, a Young.
life course perspective recognizes that age-related
aspects of human sexuality have a structure to them
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