EE.
06 Sexual selection
GENDER DIFFERENCES IN RECEPTIVITY TO SEXUAL
OFFERS (Clark & Hatfield, )
- according to cultural stereotypes, men are more eager
for sex than women; women are more likely to set limits
on such activity
- three questions:
o Would you go out tonight?
o Will you come over to my apartment?
o Would you go to bed with me?
- the great majority of men were willing to have a sexual
liaison with the women who approached them; women
were not
The Sociobiological perspective
Symons, 1979: according to evolutionary biology, animals
inherit those characteristics which insure that they will
transmit as many of their genes to the next generation as
possible to both mens and womens advantage to
produce as many surviving children as possible
- humans have only moderate sex differences in structure,
but profound sex differences in psyche; among the
differences:
o men desire a variety of sex partners; women do not
o for men, sexual attractiveness equals youth; for
women, it equals political and economic power
o men have every reason to actively pursue women
(they are genetically programmed to impregnate as many
women as possible; women have every reason to be coy);
it takes time to decide if a man is a good genetic risk (is likely
to be nurturant, protective and productive); in all societies,
women copulate as a service to men, not vice versa
Kurt Freund (1983, 1986) observed that courtship normally
consists of 4 phases:
o initial appraisal and location of a potential partner
o pretactile interaction (ex: smiling at someone,
laughing, flirting, talking)
o tactile interaction (touching, embracing)
o bringing about genital union
- a woman initiates interaction, but men initiate first
sexual intercourse; afterwards, women are freer to share
initiation
The cultural-contingency perspective
- at the other end of the spectrum are those who
argue that sexual behaviour is learned (as culture expects it
to be)
- cultural rewards and punishments shape behaviour:
if men are more adventurous than women, it is simply
because the culture encourages them to be so
- feminist and Marxist scholars were speculating that
the socio-political pressure might have at least some impact
on sexual behaviour
Gender differences in interest in erotic literature and tapes
- traditionally, erotica has been written for men; the
assumption has been that women are not interested in such
things
- Kinsey(1953): 47% of men reported having been
aroused by erotic stories; only 14% of women reported
similar levels of arousal
- recent studies: although they report different levels
of interest, objective difference suggests that both are
equally aroused
- Jakobovits (1965) found that women consistently
rated hard core erotic stories as more interesting and
sexually stimulating tha men did
Gender differences in responsiveness to erotic films
- men are more likely to sign up for an experiment
involving erotica
- both are identically aroused
- both women and men were most sexually aroused
by seeing a person of the opposite sex masturbating or
having intercourse; they were least aroused by seeing
someone of their own sex engaged in these same activities
Gender differences in sexual activity
- traditionally, theorists have assumed that sex is far more
important for men than for women
Kinsey(1948, 1953) tried to asses the sexual activity of men
compared with women throughout their lives; they asked
how often they have an orgasm during a typical week; they
found that
o men did seem to engage in more sexual activity
than women
o men and women had strikingly different sexual
histories: men are sexually active starting at 15; women
begin to be more sexually active between the ages of 16 and
20;
- once the couple begins to have sexual relations, the
man has more to say about the type and frequency of sexual
activity
- by the 1980s, the differences between sexual
activity in men and women has disappeared; the double
standard turned into a single standard
How receptive are men versus women to sexual invitations?
- traditional hypothesis: women wanting love, men
wanting sex from a relationship
- androgyny hypothesis: it may be that both men and
women turn out to be more receptive to sexual
invitations than one might expect
both men and women might be reserved about sexual
intercourse; men do not want an easy to get woman
- in both studies, men and women responded as
traditionalist would expect them to: men readily
accepted a sexual invitation; women were extremely
reluctant to do so
Type of request
Sex of requester Date Home Bed
Male 50% 0% 0%
Female 50% 69% 69%
1
MHC HETEROZIGOSITY CONFERS A SELECTIVE
ADVANTAGE AGAINST MULTIPLE-STRAIN
INFECTIONS
- experiment: released mice into large population
enclosures and challenged them with multiple strains of
Salmonella and one of Listeria
- MHC heterozygotes had greater survival and weight
than homozygotes (unlike sham controls) and they were
more likely to clear chronic infection than homozygotes
- the benefits of heterozygosity were due to
resistance being dominant rather than overdominant
(heterozygotes were more resistant than the average of
parental homozygotes, but they were not more resistant
than both)
MHC-disassortative mating preference
- heterozygosity at MHC loci may enhance resistance
to infectious diseases by increasing the diversity of antigens
presented to T cells and by generating a diverse T cell
repertoire
- one study found that heterozygotes mice had higher
T cell response to a viral infection than homozygotes, and yet
heterozygotes were least likely to survive the infection
- usually, when heterozygotes show an advantage in
population studies, it is generally interpreted as
heterozygote superiority or overdominance (i.e., MHCs function in pathogen detection
heterozygotes are assumed to have higher fitness than both
parental homozygotes). Yet, heterozygotes may have a
selective advantage over homozygotes on average if
resistance is merely dominant (i.e., if heterozygotes inherit
the resistance of the most resistant parental homozygote);
and resistance is generally dominant, at least in tests with
single pathogens
- weight provided an index for the health of males
(whereas the weight of females fluctuated mainly from
pregnancy)
- in the infected populations, females had lower
survival rates
- in the infected populations, MHC heterozygotes
were twice as likely to become pregnant and had more
pregnancies than homozygotes (not significant, however)
- MHC genotype had a significant effect on pathogen
load for the secondary, but not primary infections
- MHC heterozygotes had significantly lower bacterial
loads than homozygotes in the multiple-strain, but not in the
single-strain infections
- Heterozygotes showed a survival advantage in our
control populations during the final weeks of the
experiment, which may have been due to enhancing
resistance to various uncontrolled pathogens
MAJOR HISTOCOMPATIBILITY COMPLEX GENES,
SYMMETRY, AND BODY SCENT ATTRACTIVENESS IN
MEN AND WOMEN (Thornhill, Gangestad et al.)
- scent of developmental stability (low fluctuating
asymmetry) is attractive to women who are fertile, but not
to other women or men
- the scent of dissimilarity in major histocompatibility
complex (MHC) genes may play a role in human mate choice
- men, but not women, showed a preference for t-
shirts with the scent of MHC dissimilarity
- womens scent rating was positively correlated with
the wearers MHC heterozygosity
- fertile women did not exhibit any MHC trait
preferences; womens preference for the scent of men who
were heterozygous for MHC alleles might be stronger in
women who are at infertile cycle points
- men preferred the scent of common MHC alleles,
which may function to avoid mates with rare alleles that
exhibit gestational drive; also preferred the scent of women
at fertile cycle points
- neither FA, nor facial attractiveness in either sex
correlated with MHC dissimilarity to others, MHC
heterozygosity or MHC allelic rarity
- host-pathogen coevolution is now generally thought
to explain much of MHC genetic polymorphism because of
- MHC genes influence body odour and mate choice
on body odour attractiveness
- mouse studies suggest a preference for the scent of
MHC dissimilarity
- explanations for MHC dissimilarity scent preference:
o inbreeding avoidance
o parasite-mediated sexual-selection theory
- mate choosers should choose the infrequent
individuals who are dissimilar to confer more resistance to
the offspring
- a prediction related to the the diverse-genes
hypothesis is that individuals should avoid mating with MHC
homozygotes. Homozygotes have offspring who vary less
from each other and the mate, which increases the risk of a
within-family disease epidemic
- in both sexes, low FA appears to be associated with
increased genetic, physical and mental health (including
cognitive skills and IQ)
- symmetric men are more muscular, vigorous and
socially dominant, have lower basal metabolic rate and may
be larger in body size than asymmetric men
- the scent of symmetric men is attractive to women
who are cycling normally (not using hormone-based
contraception) and are in the fertile phases of their
menstrual cycles
- men prefer the scent of women who are in the
fertile phase of their cycle, suggesting yet another
pheromone-based system of preferences in humans
- women who use contraceptive pills prefer the scent
of individuals with similar MHC
- characters measured: ear length, ear width, elbow
width, wrist width, ankle width, foot breadth, lengths of all
fingers, except the thumb
2
 - the scent of men who had been sick tended to be
less attractive than other mens scent; sick women did not
smell less attractive
- women who have longer cycles, on average, ovulate
later in the cycle than women who have shorter cycles
- womens probability of conception across the cycle
had a near-zero correlation with their preference for MHC
dissimilarity
- sex did not interact with heterozygosity to predict
scent attractiveness
- mens scent attractiveness was not associated with
the commonness of their alleles
- womens facial attractiveness predict their scent
attractiveness
- womens estimated probability of conception at the
time of wearing the t-shirt was correlated with their mean
scent attractiveness
- older men particularly preferred the scent of
symmetrical women
- older and more facially attractive men particularly
preferred the scent of women near ovulation
- mens preference for the scent of symmetrical
women was negatively associated with their preference for
MHC heterozygotic women
- Our results address four adaptive hypotheses that
have been proposed for scent-based, MHC disassortative
mate preferences: outbreeding, produc- tion of MHC-
heterozygous offspring or of offspring with rare MHC alleles,
and increased within-brood MHC diversity
- women with zero conception risk exhibited a
significant preference for the scent of symmetry
- Manning et al. have shown a positive correlation
between ejaculate size, sperm quality and body symmetry in
men
- We found no evidence that either sex prefers scents
associated with rare MHC alleles. Indeed, men in our study
preferred scents associated with common MHC alleles. No
association between womens conception risk and
preference for the scent of rare MHC alleles was detected. A
rare-allele preference is predicted from the hypothesis that
MHC-dependent mating preferences function to generate
offspring that have reduced autoimmune problems
(associated with common MHC alleles) or that cannot be
invaded by parasites as a result of possessing alleles to which
parasites are not well adapted.
- men exhibit significant scent preferences for
women who are at high conception risk in the menstrual
cycle
HUMAN FEMALE ORGASM AND MATE
FLUCTUATING ASYMMETRY
- human female orgasm is not necessary for
conception; hence, it seems reasonable to hypothesize that
orgasm is an adaptation for manipulating the outcome of
sperm competition resulting from facultative polyandry
- if heritable differences in male viability existed in
the evolutionary past, selection could have favoured female
adaptations (orgasm) that biased sperm competition in
favour of males possessing heritable fitness indicators
- possible hypothesis for the orgasm evolution:
o orgasm has been claimed to create and
maintain the pair bond between male and female by
promoting female intimacy through sexual pleasure
o a number of evolutionists have suggested
that it works in selective bonding by promoting affiliation
primarily with males who are willing to invest time or
resources in the female and/ or males of genotypic quality
o motivates female to pursue multiple males
to prevent male infanticide of the females offspring and/or
to gain material benefits from multiple males
o functions to induce fatigue, sleep and a
prone position, and thereby passively acts to retain sperm;
o the upsuck hypothesis: orgasm actively
retains sperm by sucking sperm into the uterus
- orgasm occurring near the time (or up to 45 minutes
after) of male ejaculation results in greater sperm retention
- some theorists claim that female orgasm is an
incidental by-product of male orgasm, which is an adaptation
- female orgasm is not necessary for conception;
hence, its function could not have arisen simply to ensure
conception
- a woman might have evolved to have sex with
multiple males because she gains resources from one and
uses better genes from the other; she needs a way of
selecting the desired sperm
- sexual selection favours preferences in mates who
posses honest indicator of pathogen resistance
- a marker of male mating advantage in a variety of
species might be fluctuating asymmetry (bilateral
characters); because their developments is controlled by a
single gene, asymmetrical development can be an indicator
of developmental instability (pathogens)
- in a range of species, individuals fluctuating
asymmetry negatively predicts their fecundity, growth rate,
survival
in order for female orgasm to be correlated with mate
fluctuating asymmetry, it should occur in the estimated
time, more often with symmetrical mates
- women with men who possessed lower fluctuating
asymmetry were reported by both partners as having more
orgasms during copulation
- as a group, male investment behaviour measures
had particularly low predictive utility
3

- mens physical attractiveness had a non-significant
tendency to predict womens HSR orgasms
- mens FAI remains significantly predictive
- women who tend to act in less exclusive ways to
their mate tended to fake orgasms more often than other
women
- some evidence that mens weight and physical
attractiveness predict copulatory female orgasm frequency
independent of male body symmetry
- Smith suggested that female orgasm has 2 design
features:
it preferentially retains sperm of males of high
phenotypic quality
it promotes differential bonding with such males
- circulating oxytocin during female sexual response,
peaking at orgasm, and is believed to facilitate orgasmic
contractions of the smooth muscles of the uterus and vagina
which may lead to sperm retention
- no evidence that female orgasm is associated with
greater investment by women in their romantic relationship
or greater love for the male partner
- recent theory and empirical results suggest that
females will have evolved to trade off male investment for
male genetic quality
- the occurrence of female orgasm across mammal
species is predicted to be positively related to the
development of sperm competition adaptations in males
THE EFFECTS OF FACIAL HAIR MANIPULATION ON
FEMALE PERCEPTIONS OF ATTRACTIVENESS,
MASCULINITY AND DOMINANCE IN MALE FACES
(Neave, Shields)
- facial hair condition incorporated five levels: clean-
shaven, light-stubble, heavy stubble, light beard and full
beard
- Darwin referred to intersexual selection as female
choice because in many species, the female is the choosy
one due to different parental investment
- the features preferred by women are assumed to be
under the control of the gonadal sex-steroids beginning at
puberty when the individual enters the reproductive market
- testosterone is assumed to affect a number of facial
features that determine perceived masculinity: lateral
growth of cheekbones, jawbone and chin, the forward
growth of the eyebrow ridges and the lengthening of the
lower face, leading to a more robust face shape
- males with facial hair were perceived as being more
aggressive, older, less attractive and were subject to reduced
perception of social maturity in comparison to clean-shaven
faces
- clean-shaven faces were regarded as being
significantly more youthful, attractive and sociable than
bearded faces (Wogalte&Hosie)
- on one hand, a female might desire a male who is
strong, mature and dominant, but is also perhaps seeking a
faithful, cooperative and caring partner; it has been shown
that male faces combining elements for both mature and
childlike features are perceived as being the most attractive
- the faces were rated for: masculinity, aggression,
dominance, attractiveness, social maturity; and how
desirable the male would be for a romantic partner in a
short-term and a long-term relationship; how old she
thought the male was
expected to be linearly correlated: masculinity,
dominance, aggression, social maturity and age; for the rest
(attractiveness, preference for a short-term//long-term
partner) were expected to be U-shaped
- most attractive: light stubble; least attractive: full
beard;
- short term relationship: light stubble (most
favoured) and full beard (least favoured);
- long term relationship: light stubble (most favoured)
vs clean shaven (least favoured)
4
CROSS-CULTURAL CONSENSUS FOR WAIST-HIP
RATIO AND WOMENs ATTRACTIVENESS (Singh)
- a gynoid body fat distribution as measured by the
size of waist-hip ratio is a reliable indicator of their sex
hormone profile, greater success in pregnancy and less risk
for major diseases
- in each culture participants selected women with
low WHR as attractive, regardless of increases or decreases
in BMI
- womens mate value is determined by: hormonal
status, reproductive age, fecundity, parity and resistance to
diseases (cannot be directly observed)
- WHR becomes distinctively sexually dimorphic
during pubertal development, with a low WHR triggering
menarche in adolescent girls;
- WHR is shown to be between 0.67-0.80 in healthy,
premenopausal, Caucasian women
- low WHR is correlated with an optimal sex hormone
profile and endocervical pH, which affects sperm transport;
low WHR is also an independent predictor of successful
pregnancy in women attending an artificial insemination
clinic and in women undergoing in vitro fertilization and
embryo transfer
- women with low WHR are at a lower risk of
developing cardiovascular disorders, adult onset diabetes
and various cancers (breast, ovarian, endometrial)
- exceeding these loads (baby+food) in the hot dry
season could lead to risks of death or injury for the mother,
perhaps from heat stress; if, as seems more likely, she took
care to avoid such risks by limitating her load and her choice
of days on which to forage, there might be a nutritional
shortage for the family;
- it might be reasonable to expect an impairment of
the mothers lactation at high levels of work, heat or water
stress, and lowered food intake; the last one might increase
the risks of infection
- much longer intervals would be less effective simply
because fewer births could be fitted into the reproductive
span
- there is much evidence that the death of a baby
leads rapidly to a pregnancy in many populations
(replacement births)
- groups of children: lived to 10 years of age or more
(survivor), or died and whether it died after the subsequent
birth, or clearly before the next pregnancy (12 months or
more before the next birth)
- chimpanzees have longer IBIs than do humans;
since chimps do no provision for their families, mothers
work as a provider cannot be the explanation of the long IBI,
although her work at carrying her offspring can be

Singh: 3 body weight categories (under-weight, normal,

overweight) and 4 levels of WHR (0.7, 0.8, 0.9, 1.0); for
normal category, WHR=0.7 was found most attractive
- women were also tested for preference because
they might be subject to even more social-media pressure
- similar preference for low WHR is evident in non-
Western and Western groups
- body weight has a negligible effect on how
attractive the body looks (postsurgically); rather, the critical
variable is the proportionality of the waist and the size of the
buttocks

BUSHMAN BIRTH SPACING: A TEST FOR OPTIMAL

INTERBIRTH INTERVALS (Blurton Jones,, 1986)

- interbirth intervals (IBI) for !kung women average

around 4 years (long)
- given the observed features of !kung ecology,
shorter IBIs require a mother to carry much greater loads
(backload) of baby and food on her foraging trips;
- reproductive success would seem to be maximised
by frequent births, particularly if offspring mortality is not
taken into account; if mortality is high enough when there a
frequent, closely spaced births, then fewer, more widely
spaced births may leave more descendants
- explanations that have been offered for so long
birth intervals:
- children are not valued highly in the !kung society
- restraint on population size in order to conserve
resources and a leisurely life-style
- the work entailed by shorter interbirth intervals is
simply too much (Lee, 1972)
5
GENETIC EVIDENCE FOR UNEQUAL EFFECTIVE - As an explanation for the observed differences in
POPULATION SIZES OF HUMAN FEMALES AND the genealogies of the human NRY and mtDNA, we favor a
MALES model in which the human effective population size is
skewed toward an excess of females by sex-biased
- the time to the most recent common ancestor
demographic processes. The human mating system has often
(TMRCA) of the human mitochondria (171 to 238 kya)
=mtDNA is estimated to be older than that of the been considered to be moderately polygynous, based on
nonrecombining portion of the Y chromosome (46-110 kya) both surveys of world populations and on characteristics of
=NRY human reproductive physiology
hypothesis: natural selection has acted on the NRY,
- Another sex-specific demographic process that may
leading to a recent selective sweep
sex-biased demographic processes are cause the observed differences in the genealogies of mtDNA
responsible and the NRY is a sex bias in rates of migration among human
populations. For instance, the widespread phenomenon of
- NRY has extraordinarily low levels of sequence patrilocality (defined anthropologically as the tendency for a
diversity which has made characterization of variation
difficult and labour intense wife to move to her husbands natal domicile) could
- mtDNA has proven to be a prolific source of DNA contribute to the observed pattern if it resulted in higher
variation, even among very local populations rates of mtDNA than NRY gene flow between genetically
- because the TMRCA of a selectively neutral locus is distinct populations
influenced primarily by its effective population size, the
- we favor a hypothesis whereby sex-specific
observed disparity between mtDNA and NRY is somewhat
unexpected demographic processes act to reduce the male breeding
- the genealogy of the studied African population is population size
much deeper than those of non-African populations at both
mtDNA and the NRY
hypothesis: positive selection has reduced variation
on the human NRY; all else being equal, a recent
selective sweep on the NRY would indeed cause it to
have a more recent TMRCA than that of mtDNA in
accordance with the results; however, no population
sampled exhibits a significant excess of rare variants in
NRY, which is thought to be one of the genetic
signatures of a recent selective sweep

- purify- ing selection. For both genetic systems,

diversity (and, hence, the TMRCA) may be reduced by
selection acting against deleterious mutations at linked sites
- Whereas all mtDNA genes are single copy (and play
a vital role in cell metabolism), nearly all genes on the NRY
are multicopy or have closely related autosomal or X-linked
homologs (Skaletsky et al. 2003). Consequently, we do not
expect purifying selection to differentially reduce the
effective size of the NRY relative to that of mtDNA
- These lines of reasoning lead us to postulate that
sex- specific demographic processes are the most likely
causes of the observed discrepancy in the TMRCAs of mtDNA
and the NRY
- Based on one of the few direct comparisons of
patterns of mtDNA and NRY sequence variability, Tang et al.
(2002) suggested that the difference in coalescence times for
mtDNA and the NRY (which were given in numbers of
generations) could be minimized to some degree (and hence
explained) by taking account of evidence (see below) that
males have a longer generation time than females (i.e., 30
versus 25 years)
6