Socioaffective Neuroscience & Psychology

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An evolutionary behaviorist perspective on

orgasm

Diana S. Fleischman

To cite this article: Diana S. Fleischman (2016) An evolutionary behaviorist perspective on

orgasm, Socioaffective Neuroscience & Psychology, 6:1, 32130, DOI: 10.3402/snp.v6.32130

To link to this article: https://doi.org/10.3402/snp.v6.32130

© 2016 Diana S. Fleischman

Published online: 25 Oct 2016.

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 æ

THEORETICAL ARTICLE

An evolutionary behaviorist perspective on orgasm

Diana S. Fleischman, PhD*
Department of Psychology, University of Portsmouth, Hampshire, United Kingdom

Evolutionary explanations for sexual behavior and orgasm most often posit facilitating reproduction as the
primary function (i.e. greater rate of fertilization). Other reproductive benefits of sexual pleasure and orgasm
such as improved bonding of parents have also been discussed but not thoroughly. Although sex is known to
be highly reinforcing, behaviorist principles are rarely invoked alongside evolutionary psychology in order to
account for human sexual and social behavior. In this paper, I will argue that intense sexual pleasure,
especially orgasm, can be understood as a primary reinforcer shaped by evolution to reinforce behavior that
facilitates reproductive success (i.e. conception through copulation). Next, I will describe an evolutionary
account of social shaping. In particular, I will focus on how humans evolved to use orgasm and sexual arousal
to shape the social behavior and emotional states of others through both classical and operant conditioning
and through both reproductive and non-reproductive forms of sexual behavior. Finally, I will describe how
orgasm is a signal of sensitivity to reinforcement that is itself reinforcing.
Keywords: orgasm; evolutionary psychology; behaviorism; shaping; reinforcement; punishment; classical conditioning;
operant conditioning; signaling

*Correspondence to: Diana S. Fleischman, Department of Psychology, University of Portsmouth, King Henry
Building, King Henry I Street, Portsmouth, Hampshire Po1 2DY, UK, Email: [email protected]

This paper is part of the Special Issue: Orgasm: Neurophysiological, psychological, and evolutionary perspectives.
More papers from this issue can be found at www.socioaffectiveneuroscipsychol.net

Received: 3 May 2016; Revised: 20 August 2016; Accepted: 23 September 2016; Published: 25 October 2016

magine the pleasure of seeing a cute, happy baby; (see Wheatley and Puts (2015) for a good overview and

I sitting in the warm sunshine; eating something fatty

and sugary; or getting relieved of a few minutes of low-
voltage intermittent shocks. You would likely estimate
other papers in this volume). Instead, I will consider
orgasm the pinnacle of a continuum of highly reinforcing
sexual pleasure.
the pleasure of orgasm as greater than any of these things.
Evolution has used orgasm to train us toward adaptive Orgasm and sexual behavior as primary
behavioral ends; orgasm and high sexual arousal are reinforcers
currencies that tap directly into bliss states. Reinforcement Animals have different primary reinforcers depending on
and reward are better motivators of behavior and are which cues are most likely to be statistically associated
better at shaping new behaviors than punishment (Pryor, with adaptive outcomes. A reinforcer is any stimulus that
1999; Skinner, 1938). Given these two principles, the thesis increases the frequency, duration, or intensity of a be-
in this paper is that (1) evolution has used orgasm to havior (Schultz, 2015). Organisms that evolve to experi-
promote adaptive behavior including non-reproductive ence pleasure as a result of engaging in adaptive behavior
sexual behavior, (2) we have evolved to use orgasm and will be more motivated to engage in such behavior. When
sexual arousal to shape one another’s behavior, and (3) a reward or ‘primary reinforcer’ is paired with a given
orgasm serves as a signal to another person of devotion, stimulus, that stimulus becomes a secondary reinforcer.
vulnerability, and malleability, which is, in itself, reinforcing. For example, when a dog is clicker trained, he hears a click
This paper won’t go into how orgasmic pleasure works or when he is given food for a given behavior. The click
the evidence that orgasm facilitates reproduction directly becomes a secondary reinforcer that can motivate future

Socioaffective Neuroscience & Psychology 2016. # 2016 Diana S. Fleischman. This is an Open Access article distributed under the terms of the Creative Commons 1
Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), allowing third parties to copy and redistribute the material in any medium or format and to
remix, transform, and build upon the material for any purpose, even commercially, provided the original work is properly cited and states its license.
Citation: Socioaffective Neuroscience & Psychology 2016, 6: 32130 - http://dx.doi.org/10.3402/snp.v6.32130
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Diana S. Fleischman

behavior because it has been paired with food, a primary motivated caretaker who will provision them and protect
reinforcer (even after food is no longer paired with the click). them from harm; a smile is a cue that this adaptive problem
Orgasm, in behaviorist terms, is a primary reinforcer, a is being solved by the behavior of the infant.
pleasurable unconditioned stimulus that, without any More salient to the topic of orgasm, pictures of nude
classically conditioned association, is inherently and in- adult women would be a strong primary reinforcer for
nately reinforcing. Orgasm, like other primary reinforcers, men after puberty but would probably not be a primary
can make stimuli that are paired with it (e.g. a color, reinforcer for prepubescent males. Researchers have found
someone’s face, a specific smell) reinforcing. Food, warmth, that male rhesus macaques will ‘pay’ (give up part of a juice
sex, and sleep are examples of primary reinforcers and reward) to view the genitals of female macaques or the
these are evolutionarily conserved among many organisms. faces of high status males but will not ‘pay’ to view the faces
Because evolved psychological mechanisms solve adaptive of low status conspecifics (Deaner, Khera, & Platt, 2005).
problems with biologically prepared inputs and because Another study found that peak fertility female macaques
so many cues have been recurrently associated with these preferred to view the faces of male conspecifics (Lacreuse,
inputs, there are likely many primary reinforcers that exist Martin-Malivel, Lange, & Herndon, 2007) and showed
for humans that don’t exist for animals. reduced preference during periods of lower fertility. These
Consider evolution as an agent that calibrates the sub- studies show that social information in and of itself can
jective pleasure of behavior based on how adaptive it is act as a primary reinforcer. For the sexually inexperienced
for the organism. It’s no wonder that finding and consum- male rhesus macaques, both the photos of the high status
ing food and sex are primary reinforcers as they facilitate male’s faces and the genitals of the females had not pre-
survival and reproduction. However, considering sex as viously been associated with food, sex, or other positive
reinforcing can be taken one step further. According to an reinforcement (i.e. not classically conditioned) (Michael
adaptationist account of conditioning, any cues of court-
Platt, primatologist, April 7, 2016 personal communication).
ship or reproduction that are consistently associated with
An evolutionary perspective would predict that sexual
higher reproductive success recurrently throughout evolu-
primary reinforcers do not show equipotentiality; they
tionary history can become a primary reinforcer. This can
are differentially rewarding depending on how adaptive
include practicing courtship skills (e.g. telling stories), the
they are to an organism. This may be contingent on age,
attention one gets from potential mates (e.g. eye contact,
sex, and fertility status. While stimuli like nudity may or may
smiling), initiating sexual behavior (e.g. intimate touch,
not be innate or a primary reinforcer, we should expect that
kissing) all the way up to penetration, orgasm, and seeing
for stimuli that are very salient to reproduction, there is at
one’s partner indicating intense pleasure and orgasm.
least a biological preparedness for associating sexual stimuli
Another prediction derived from an evolutionary per-
with pleasure, thereby increasing their reinforcing qualities.
spective is that primary reinforcers differ over the life-
span and that different primary reinforcers have different The sex of the organism should also be an important variable
salience depending on the adaptive problems that were influencing the valence of sexual pleasure.
recurrently faced by our ancestors during particular life
stages. For instance, parental praise will be a primary An adaptationist account of sex differences in
reinforcer for children but less so for adults. Attention sexual reinforcement
(e.g. eye contact, orienting body language) from attractive Sex differences in orgasmic capacity and sexual pleasure
members of the opposite sex should be more of a pri- can be explored within an evolutionary behaviorist frame-
mary reinforcer for reproductive-aged men or women than work. We infer from the presence of ejaculation and
children. We should also expect this with sex; sexual context-dependent facial expressions that in many other
behavior will yield the most pleasure when it is ful- species, only males consistently experience orgasm during
filling adaptive goals such as improved bonding between sexual behavior (Ferro, 2013; Lloyd, 2009). Ejaculation
parents or forming a new bond with someone of high status. could be associated with no more pleasure than urinating
For example, behaviorists would argue that a smile is a or defecating if ejaculation was not necessary for repro-
secondary reinforcer because it has been classically con- duction. An adaptationist perspective explains why males
ditioned by being paired with food or another primary consistently experience the peak pleasure of orgasm from
reinforcer. An evolutionary learning perspective might differ. intercourse more than other sexual activities; orgasm moti-
Experimental evidence shows that infants, 2
7 months old, vates men toward the most adaptively important out-
can be conditioned to prefer sounds using a smile as rein- comes. However, men also consistently reach orgasm more
forcement (Routh, 1969). This indicates a smile may be an easily when engaging in other forms of sexual behavior.
innate positive reinforcer or, at least, that infants are Men’s gametes (sperm) are much less costly to produce
biologically prepared to associate a smile with primary and men have much lower obligate parental investment
reinforcers like food or warmth. For infants, the major than women (Trivers, 1996). For men to be reproductively
adaptive problem they face is having an engaged and successful, they need, on the low end, to only engage their

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 Evolutionary behaviorist perspective on orgasm

time and resources as much as is necessary to have sex with been found to be a greater driver of behavior that is
a woman. resistant to extinction than consistent reinforcement (e.g.
Consider men’s rate of orgasm in light of Error gambling on a slot machine is more resistant to extinction
Management Theory (Haselton & Buss, 2000) also known than pulling a lever with a consistent payout) (Pryor,
as the Smoke Detector Principle (Nesse, 2001). If there are 1999). For women, sex is much more costly both in terms
two kinds of errors an organism can make, evolution of potential parental investment and sexually transmitted
will bias behavior toward the less costly of these errors. disease risk and thus we should expect evolution to be
If you design a smoke alarm, it’s better for it to be more selective about the sort of sexual behavior that
calibrated to go off in error when there is no fire than it is should be reinforced with extreme pleasure.
for it to remain silent when there is a fire. For men, sexual In order for a woman to be reproductively successful,
behavior has low potential costs compared with great she must carry a child for 9 months and, until recent
potential reproductive benefits. This is why men are history, breastfeed for another 3 years. Women can have
calibrated to over-perceive sexual interest on the part of far fewer offspring than men over their lifetimes, have less
women (Perilloux, Easton, & Buss, 2012). For the same variable rates of reproduction, and have little reproductive
reason, we should expect men to experience orgasm in a incentive to have sex with more men. A woman who has
variety of conditions, even if they do not lead directly intercourse with 100 men in a year will, on average,
to fertilization. Experiencing orgasm when engaging in have no more offspring than a woman who has regular sex
an act that cannot result in reproduction is a less costly with one fertile man. Moreover, women are much more
error than not experiencing orgasm during an act that can susceptible to sexually transmitted infections than men
lead to conception (Figueredo et al., 2005). and have much greater disease burden (e.g. sterility) as
There is evidence for this both in humans and animals. a result of these infections (Madkan, Giancola, Sra, &
Semen can be collected easily in many birds and mammals, Tyring, 2006). For these reasons, copulation isn’t always
by providing a male with a crude facsimile of a female or adaptive and a woman who was easily orgasmic in a variety
vagina (Rouge & Bowen, 2002). Men, more than women, of conditions could be making a more costly error: being
are prone to fetishes and paraphilias, and easily pair sexual motivated to engage in behavior that is unlikely to result
arousal with a given stimulus through classical condi- in the optimal reproductive outcome of conceiving with a
tioning (Rachman, 1966). However, just because men are genetically fit male who is free of disease (Miller, 2000).
more likely to reach orgasm with myriad forms of sexual Thus, it is useful to consider what kinds of sexual
behavior and with different sexual partners does not mean activity evolution would want to orgasmically reinforce in
that orgasm or sexual pleasure will be equal across these women for fertilization and conception versus for bond-
contexts. We should expect more adaptive conditions to ing. I speculate that this disease risk and the concomitant
lead to greater sexual pleasure and more intense orgasm focus of sexual pleasure away from the vaginal mucous
(more on this later). However, we should also expect membranes (which are more likely to transmit disease
plasticity built into the sexual motivation system through upon contact than the clitoris and external vulva) might
reinforcement. Men have preferences for young, fertile have been a driver toward the variety of sexual practices
women and various forms of attractiveness that signal (e.g. frottage) humans engage in that do not involve
health and reproductive value (Sugiyama, 2005). However, copulation. Further evidence for this is that most women
men will experience orgasm and sexual pleasure with cannot achieve orgasm through vaginal intercourse alone
women that are available to them even if they do not evince (Lehmiller, 2013; Lloyd, 2009) and most women do not
these cues. In terms of fertilization, evolution should consider the vagina to be the most important erogenous
maximize orgasmic pleasure for men who are with zone compared with the clitoris (S. Fisher, 1973). Later,
partners most likely to conceive, that is, women of we will explore how orgasm may be adaptively calibrated
reproductive age who are healthy enough to carry a for bonding between parents and social benefits that
pregnancy, but men will also be motivated with a history translate into increased reproductive success (e.g. forma-
of orgasmic reinforcement toward women who will be tion of alliances that increase status).
more likely to choose them repeatedly as mates.
Many scientists have been puzzled by women’s orgasmic Sexual pleasure calibrated to maximize
frequency. Why don’t women consistently experience orgasm fertilization
and, conversely, why do women have orgasms at all? The Pleasure and primary reinforcement are not binary
fact that women experience orgasm, at least some of the phenomena. We should expect pleasure to be gradient
time, is one piece of evidence that orgasm is a primary and calibrated to the scarcity or adaptive value of the
reinforcer for some kinds of adaptive behavior. Some stimulus. Drinking water is more pleasurable when one is
(e.g. Prause, 2011) have speculated that it is precisely thirsty and eating high-calorie foods
especially those that
because orgasm is variable in women that it may be more have nutrients that were scarce throughout evolutionary
reinforcing than it is for men; variable reinforcement has history (e.g. salt, sugar, fat)
feels more pleasurable than

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Diana S. Fleischman

eating foods that were abundant or offer less nutritional with health, e.g. Miller, 2015). Thus, orgasm may serve to
value. Orgasm may be similarly calibrated and sensitive to reinforce engaging in repeated sex with men who are likely
contexts of scarcity or abundance. For both men and to produce such sexy sons, regardless of whether these
women, we should expect sex and orgasm to feel more traits signal health or quality.
pleasurable when mates or the opportunity for sexual
contact are scarce. We should predict that sexual contact Orgasm as reinforcement to facilitate pair
with healthy and attractive conspecifics to yield more bonding and social bonds
pleasure. Aspects of health and attractiveness are normally Thus far, we have mostly considered the idea that orgasm
distributed. Thus, we can expect that health and attrac- reinforces sexual behavior toward conception. This may
tiveness increase the pleasure and reinforcing quality of have been the reason why orgasm initially evolved to
sexual behavior not only because they are statistically be pleasurable and seems to be the motivating force for
associated with greater fertility and therefore greater most males of other sexually reproducing species. It has
reproductive success but also because those sexual part- long been theorized that the aseasonal (i.e. no distinct
ners are necessarily more scarce than partners more period or season of fertility and no estrus) and high sex
average on genetically endowed qualities. Evolution has drive of humans (i.e. not strongly tethered to fertility)
used orgasm to reinforce behaviors that are directly related is indicative of its evolution as a mechanism for rein-
to fertilization such as an orgasm associated with ejacula- forcing bonds between two people. These bonds have
tion during intercourse. However, it’s also clear that been discussed especially with regard to their importance
orgasm facilitates the motivation to engage in other non- between parents provisioning for offspring (Marlowe,
reproductive sexual behaviors such as oral sex, masturba- 2000). Although this discussion will focus on parental
tion, and same-sex sexual behavior. bonding, ‘pair bond’ in this paper can indicate any attach-
Let us first consider how orgasm could be calibrated ment between two people whether monogamous or poly-
to help women gain genetic benefits that would make gynous, reproductive or non-reproductive, heterosexual
each of their costly offspring more likely to be healthy and or homosexual.
reproductively successful. There is controversial evidence If evolution designed a woman to have sex with a man
(Wheatley & Puts, 2015) that female orgasm facilitates for his superior genetic complement or a man to have sex
conception (e.g. by dipping the cervix into the seminal in order to conceive offspring, they need not be motivated
pool); however, orgasm need not facilitate conception to engage in sex longer than is needed for fertilization
directly for the purposes of increasing reproduction with and certainly would not be motivated to engage in non-
certain males but instead could reinforce repeated sexual reproductive sex. Indeed, for many researchers, bonding is
behavior with those males. How would evolution optimally seen as the primary function of female orgasm; ‘orgasm
calibrate female orgasm to motivate women to have con- serves as a secondary reinforcer linking sexual behaviors
ceptive sex? Some evidence has shown that women are and partner affiliation’ (Prause, 2011). However, others
more likely to have orgasms with men who show good see sire choice as its primary mechanism (Puts, Dawood,
genetic quality. Attractiveness, masculinity, and symmetry & Welling, 2012). In practice, because the characteristics
have been associated with health as they are expensive of good genes, good mates, and good social partners often
signals to produce (Rantala et al., 2012; Rhodes Chan, overlap (e.g. strength, empathy, health), it is often difficult
Zebrowitz, & Simmons, 2003; Zahavi, 1975); Some studies to disentangle these influences.
have shown that women are more likely to have orgasms If orgasm functions primarily to form and maintain
with men who are masculine and symmetrical (Puts, pair bonds, especially between parents, this leads us to
Welling, Burriss, & Dawood, 2012; Thornhill, Gangestad, expect somewhat different design features of orgasm than
& Comer, 1995). The vigor, health, coordination, and theory those for the benefit of merely facilitating reproductive
of mind that aid in facilitating orgasm in a partner are all sex. In the case of parents, there isn’t much information
costly signals of health and fitness as well (Miller, 2000). about how sexual behavior connects men and women in
The ‘sexy sons’ hypothesis (R. A. Fisher, 1930) posits the service of provisioning for offspring. One can imagine
that females should choose males who are likely to pro- that feelings of pleasure and well-being would lead to
duce male offspring that other females will find sexually more positive associations in a couple caring for a child or
attractive. Traits that are attractive to females are often but children; this would enable them to better allocate effort
not necessarily associated with health. For instance, as well as forgive one another for errors, asymmetries in
women who prefer longer penises are more likely to have effort, or other indiscretions that would, in the case of a
vaginal orgasms (Costa, Miller, & Brody, 2012) demon- less reinforced pair bond, cause one or both parties to
strating a drive to mate with males who may produce sons abandon the relationship. In the case of pair bonding
that other females prefer but not with a trait that has between parents, a man’s pleasure at sexual contact with the
(thus far) been shown to correlate with health or quality mother of his children may prevent him from alloca-
(But see also evidence that erectile function correlates ting his effort or resources to other mating opportunities

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 Evolutionary behaviorist perspective on orgasm

or other children. For a woman, continued sexual plea- We might expect orgasm would reinforce the attachment of
sure with the father of her children could prevent two people after a period of divestment. For example, if
her from abandoning a relationship that is provisioning orgasm serves to reinforce closeness and positive associa-
and securing her offspring and could act as a signal of tion between two people, we might expect intense sexual
assurance of paternity to a mate provisioning offspring pleasure, orgasm, and intense orgasm to occur after re-
and offering protection. Thus, orgasm may act as a safe- lationship stress (e.g. threat from a rival, absence from a
guard and mechanism for forgiveness of defection. The partner) rather than during every sexual episode. This
outcome of having healthy offspring that survive to could be a complementary reason why men find cues of
adulthood is distal but the pleasure in the pair bond other males mating with their mate sexually arousing
facilitates it in the short term. These kinds of factors (Pound, 2002), not just to engage in sperm competition
maintaining parental bonds may not be that important but to strengthen the association between proximity and
in the modern age, but in many hunter gatherer groups sexual pleasure to their mate to reignite the pair bond.
thought to resemble ancestral human societies, children Thus, we should predict that strong pair bonds should
without paternal provisioning have lower survivability not exhibit regular orgasmic frequency but orgasm may
(Hurtado & Hill, 1992). instead track the degree to which strong reinforcement
Oxytocin is thought to be important for romantic would be adaptive.
bonding and is released during close affectionate and Another piece of evidence for orgasm as a pair bond
sexual behavior and most importantly during orgasm. mechanism is that foreplay increases the rate of orgasm in
Oxytocin is positively reinforcing (László et al., 2016), women (Singh, Meyer, Zambarano, & Hurlbert, 1998).
increases the sensitivity of non-human primates to positive This may indicate that foreplay classically conditions
reinforcement, and increases the reinforcing properties women to be secondarily reinforced by increased proximity
of seeing another monkey reinforced (Chang, Barter, to their partner as well as their time and attention. This
Ebitz, Watson, & Platt, 2012). Oxytocin may increase the could also indicate that orgasm as a signal is a reinforce-
strength of orgasm and is released in amounts based on the ment, a consequence that operantly conditions devoting
quality of orgasm (Behnia et al., 2014) and increases attention and time to the partner (but more on that later).
the sensitivity to socially reinforced (e.g. smiling faces) Thus far, the evidence indicates that an attractive and
learning (Hurlemann et al., 2010). However, the fact dominant male may increase the rate of orgasm in women.
that sexual excitement and satisfaction decreases over the Dominant men may also have the time, experience, and
course of a relationship and are greater with novel partners confidence to engage in more extensive foreplay without
implies that orgasm is not intended to pair bond couples being interrupted by rivals. It seems that foreplay can be
indefinitely but may only improve pair bonding for the used by dominant men to cultivate the pair bond with an
duration in which children need intensive care and pro- especially attractive partner (as this will entail fewer
visioning (H. E. Fisher, 1989). opportunity costs for copulating with others), but that
The pair bonding hypothesis of orgasm has been lower status males may also be able to use the same
supported and contradicted by evidence for and against strategy: time, attention, and knowledge of a partner to
the association of orgasmic frequency and relationship increase attachment and reinforce the pair bond.
satisfaction. Coital (but not non-coital) orgasm frequency Let us now briefly consider orgasm and sexual pleasure
has been found in some studies to correspond to good from the perspective of forming and maintaining bonds
dimensions of relationship quality such as intimacy, passion, between men and women for various benefits other than
love, and satisfaction (Costa & Brody, 2007). However, directly provisioning offspring. One might predict that the
other studies have found that attractiveness but not pleasure of sex and orgasm would be more associated with
positive dimensions of relationship quality account for relationships that optimally conferred status, resources, or
orgasm (orgasm during last coitus) (Shackelford et al., safety as a way of reinforcing association with that social
2000). If one considers attachment and affection to partner and as a way of sexually reinforcing that social
another person as a behavior that can be reinforced with partner’s association with you. It’s difficult to disentangle
orgasm, we might not expect consistent orgasmic and orgasm in response to ‘good genes’ (e.g. characteristics
sexual pleasure. With animals, a behavior that is being like attractiveness and masculinity), as these character-
successfully and consistently produced need not be re- istics also are likely to confer high status in social groups
inforced consistently (e.g. a dog sitting), but behavior and lead to social and resource benefits of association
that is decreasing in frequency may need to be carefully regardless of whether sex causes conception with these
shaped when any subtle cue of that behavior reappears. A men. One example of using sexual pleasure and orgasm
waning behavior like affection or attachment may be to garner social benefits is the case of female baboons
more effectively strengthened by, for example, rewarding forming sexual friendships with males (Smuts, 1985).
the desired cues with variable reinforcement (e.g. the slot The female allows sexual access to a male while she is
machine example used earlier) (Pryor, 1999; Skinner, 1938). in estrus and that male protects her and her offspring.

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This isn’t a perfect example because the friendships are to reason that bonds between people of the same sex
platonic between estrus periods and maintained by (homosocial) can be strengthened with sexual behavior.
proximity and grooming. However, we could say that Affiliating with others and engaging in cooperative
the female’s sexual pleasure reinforces proximity with exchange activates neural reward centers in the brain
a male who protects her and the male’s sexual pleasure (Bora, Yucel, & Allen, 2009). These interactions can be
reinforces attention and care for the female. Among even more reinforcing if they include associations of sexual
human opposite sex friendships, both men and women pleasure.
want opposite sex friends who are agreeable and depend- The affiliation hypothesis of homosexual behavior
able, but men tend to prefer opposite sex friends who (Fleischman, Fessler, & Cholakians, 2015) proposes that
are attractive, and women tend to prefer opposite sex selection used the pleasure of sexual behavior to promote
friends who have resources and physical strength (Lewis, same-sex social bonds. In many societies that are thought
Conroy-Beam, Raja, Dekay, & Buss, 2011). Sexual plea- to resemble social groups throughout human evolutionary
sure and orgasm, but also mere proximity to attractive history, social bonding and alliances play a critical role
cues, may maintain such relationships. Furthermore, in survival. Some examples include defense in violent
when one gives sexual pleasure to a social partner, this conflict (Van Vugt, 2009), surviving food shortfalls (Hill &
increases his or her preference for you over other friends Hurtado, 2009), care during illness or injury (Sugiyama,
when there is competition for his or her status, attention, 2004), and cooperative child rearing (alloparenting) (Hrdy,
or other resources. 2009). Among humans and non-human primates, social
Orgasm is highly complex and occurs in a variety of bonds translate into better survival and reproductive
sexual contexts. Although sire choice can encompass success (Silk, Alberts, & Altmann, 2003; Silk et al., 2010).
many aspects of reproduction (e.g. orgasm with a symme- In the toolkit of means of reinforcement, sexual pleasure and
trical male), the so-called ‘sire choice’ hypothesis of orgasm would have been very useful both with members of
orgasm posits that orgasm causes women to ‘preferentially the same sex, and members of the opposite sex, for cementing
retain’ sperm of higher genetic quality (Sela et al., 2015). and maintaining social bonds. Evolution maintains sexual
However, orgasm, seems overly complex in our species to pleasure for a diversity of sexual behavior among humans
have evolved to mainly solve this particular problem. To and facilitates motivation to engage sexually to different
summarize, I tend to agree with Hrdy (1996): ‘why solve adaptive ends.
a (presumably relatively simple?) cell transport problem
through the evolution of a complex psychophysiological
phenomenon that requires selection pressures to elaborate Classical and operant conditioning and orgasm
and link up various organs’. Orgasm seems well-designed Classical conditioning, as discussed earlier, is a process
as a mechanism to increase copulation among genetically by which a neutral stimulus is paired with a primary
desirable mates. However, we would also expect that evo- reinforcer, like sexual pleasure or orgasm, which thereby
lution would select against orgasm and sexual pleasure in makes the neutral stimulus in itself reinforcing. When
other contexts that didn’t facilitate direct fertilization if two people engage in sexual behavior and have orgasms
they were maladaptive. Orgasm occurs in many different they are associating sexual pleasure with the characteristics
contexts and with people both of the same sex and the of the other including proximity, smell, taste, and form;
opposite sex; this may indicate that some non-reproductive these all become secondary rewards/reinforcers. When two
sexual behavior is adaptive. people have repeated erotic contact, they become classi-
cally conditioned to perceive one another as secondary
Same-sex sexual behavior and orgasm reinforcers and can better shape one another toward their
For many species, there is no motivation for sex or pleasure own strategic goals.
in sex if there is no chance for reproduction; evolution is If one person in a dyad is associated with sexual
capable of eliminating the pleasure of behaviors, including pleasure, he or she can very easily engage in what is called
sexual behaviors, if there is no adaptive benefit. This is one ‘negative punishment’, withholding positive consequences
reason why same-sex sexual behavior has been seen as a for a behavior as a means of punishment and an adaptive
mystery to biology and psychology; how can a behavior tactic for extinguishing the behavior. Take, for example,
that offers no possibility of direct reproductive success ‘silent treatment’; If an individual values social interaction
be maintained in the population? However many people with a sexual partner, then taking away this reinforcer
feel the answer to the following question is obvious: Why is a way of simultaneously punishing the behavior and
might opposite sex partners engage in sexual behavior that no longer supplying positive reinforcement toward that
has no possibility of reproduction (e.g. the woman is behavior. Silent treatment will be much more painful if
already pregnant or one partner is past fertile age)? It’s someone’s mere presence, eye contact, or voice is strongly
likely we would say they engaged in the act to increase their secondarily reinforced by sexual pleasure. Furthermore,
intimacy or to give one another pleasure. It also stands withdrawal of other stimuli often associated with orgasm

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 Evolutionary behaviorist perspective on orgasm

like touch, eye contact, close proximity, and smell can be emotions are evoked to punish the romantic partner. In
used more subtly (than silent treatment) to shape behavior the case of men, jealousy is more often evoked in the case
both consciously and unconsciously. of sexual infidelity as it indicates that men may have heavily
Orgasm can also be used more directly to operantly invested in offspring that are not genetically related to
condition behavior with or without classical conditioning. them (Buss, Larsen, Westen, & Semmelroth, 1992). In this
Operant conditioning is a process by which a behavior is case, a man may punish a woman in a variety of ways for
followed by positive or negative consequences that increase having sex outside of the pair bond because it interferes
or decrease the frequency of the behavior. An animal will with his adaptive strategy of investing in offspring that
engage in a behavior for a food reward and increase the are genetically related to him. Women are slightly more
frequency of that behavior if food rewards continue. likely to experience emotional jealousy indicating the
possible divestment of resources away from her or her
The evolutionary psychology of dyadic social offspring. She may punish the man for divesting resources
shaping others’ behavior as an extended to another woman as she wants to dissuade him from
phenotype interfering with her strategy of obtaining resources and
In ‘The Extended Phenotype’, Dawkins advances the hy- security from her romantic partner. When individuals
pothesis that those things that are proximal to an organ- engage in sexual behavior, they increase the leverage they
ism, their physiological characteristics, behavior patterns, have to both reinforce and punish behaviors to promote
and individual differences are not the only things that strategic facilitation and interfere with strategic interference.
make up their phenotype (Dawkins, 1999). A phenotype A variety of tactics can be used to reinforce and punish
also consists of an organism’s effects on the environment, others, especially if you are a secondary reinforcer to
including its effects on other organisms. One good example someone because of classical sexual conditioning. While
of this is parasitic manipulation. Toxoplasma gondii has this hasn’t been explored directly in the literature before,
an extended phenotype that includes rat psychology. This there is one paper that details possible ‘manipulation tactics’
protozoa causes rats to increase their attraction to the that can be used between romantic partners (Buss, 1989):
smell of cat urine thus facilitating being eaten by cats, charm, silent treatment, coercion, reasoning, regression,
Toxoplasma’s primary host (Berdoy, Webster, & Macdonald, and debasement. Each of these can be viewed from a social
2000). In a more sexually relevant example, the ‘Bruce shaping perspective and most could be augmented with the
effect’ is a pregnancy disruption that occurs when a female association of sexual pleasure. The ‘charm’ tactic includes
mouse smells an unfamiliar male mouse’s pheromones acts of love and affection, compliments, gifts, and promising
(Dawkins, 1999). In this case, her pregnancy is immedi- a reciprocal favor in exchange for a behavior in the mate,
ately terminated and she becomes sexually receptive and other means of operantly and classically conditioning a
fertile much more quickly than if she had carried her partner than sexual behavior. We already discussed silent
litter to term. Dawkins argues that one does not have to treatment above as an example of negative punishment.
view the abortion and early receptivity of the female Here, I want to stress that positive association between a
mouse as only an adaptation on her part but can also behavior and a reinforcing outcome like sexual pleasure is
view this as an adaptation on the part of the male mouse. unlikely to be overt and, in fact, may work better for
Manipulation of anything external, including the manip- strategic goals if the manipulation is covert or uncon-
ulation of other organisms and conspecifics, is under just scious. Self-deception is likely at play here; people are
as intense selection pressure as those phenotypic charac- rarely conscious of the ways they manipulate others, and
teristics that enable the organism to adapt to its immediate consciously verbalizing how you are manipulating some-
environment. Some of the physical and social environment one else through reinforcement, would likely be viewed
represents an organism’s phenotype, and humans are an as unethical or sociopathic. The tactics of manipulation
example par excellence of a species that shapes its social people use both sexually and non-sexually very likely
environment in myriad ways (e.g. niche construction). exceed those they can verbalize. It doesn’t seem that people
Orgasm and sexual arousal can be used to extend one’s are either often consciously aware of how they have sex to
phenotype into the minds of others in order to manipulate reinforce behavior or that they are willing to admit this in
them to one’s own strategic goals. questionnaires. For example, individuals report engaging
Between any two people, interests will sometimes align in sex for attraction or pleasure or to say ‘I’ve missed you’
and sometimes diverge. The adaptive interests of indivi- very frequently, but very infrequently report that they have
duals can be furthered with a variety of strategies; other sex to manipulate another person into a given behavior
individuals can engage in strategic interference or strategic (Meston & Buss, 2007). Perhaps, there has been selection
facilitation (Buss, 1989). One of the most fraught kinds of for ignorance of one’s own manipulation tactics so that
relationships where there is the most at stake is between they have greater effectiveness.
romantic partners whose interests can diverge in impor- This is an area that could be important for further
tant ways. For example, in the case of jealousy, negative study. For example, having sex because you’ve missed

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Diana S. Fleischman

another person or to make up after a fight may be a way to experience emotional jealousy, we should expect that the
to orgasmically reinforce their proximity, the dissolution intensity of sexual pleasure and orgasm of a woman’s mate
of their anger, or their investment in the relationship with another partner should elicit acute jealousy because
(whether this takes the form of resources or security). of its likelihood of eliciting greater attachment through
However, it is not only the pleasure of orgasm that rein- classical conditioning.
forces behaviors; seeing another person have an orgasm
can be in itself reinforcing and a signal of sensitivity to Conclusion
reinforcement and punishment. This paper has been an overview of a potentially over-
looked framework for elucidating sexual behavior. One
Orgasm as a signal of sensitivity to shaping reason for this may be that self-deception is involved in
Given what has been discussed so far in this paper, it shaping others. This framework helps explain why social
would be surprising if cues of intense sexual pleasure bonds and influence can be amplified with the addition of
and orgasm in another person were not themselves pri- sexual pleasure and, especially, orgasm. It also explains
mary reinforcers. Perceiving that you have given someone why we, as humans, have so much attention and memory
else intense sexual pleasure can be a signal of trainability. for the kinds of pleasure our partners find most reinfor-
Seeing intense sexual pleasure in another person is an cing and intense. Many of these ideas have yet to be
indication of the strength of the extension of your pheno- explored through scientific investigation. Do people find
type into their minds, like altering a landscape to bear fruit. ‘silent treatment’ or distance most painful from those that
This may be yet another reason that orgasm is less frequent give them the most sexual pleasure? Does one find the
in women than it is in men. If men have, over evolutionary signal of orgasm most reinforcing from those that are the
history, experienced paternity uncertainty, they would most strategically important to one’s adaptive goals?
not only be sensitive to women’s fidelity but also sensitive How does sexual pleasure reinforce bonds between
to the extent that their partner was specifically sensitive people of the same sex in different contexts? Explicitly
to receiving pleasure from them as a sexual partner and using the psychology of learning and social shaping can
thus more reinforced by their company. If one had a pre- expand an evolutionary account of sexual behavior and
cious resource in a lockbox, one would feel more secure the myriad reasons it is adaptive both for conception,
if that lockbox had fewer combinations capable of opening stronger social bonds, and strategic facilitation.
it. Similarly, if one’s sexual partner is intensely reinforced
by one’s sexual stimulation, this indicates that they are,
potentially, uniquely reinforced to the exclusion of others Conflict of interest and funding
and therefore one’s investment is secure. This also indicates The authors have not received any funding or benefits
that they needn’t try to maintain as intense a monopoly from industry or elsewhere to conduct this study.
on sexual access through ‘mate guarding’. Indeed, men’s
relationship satisfaction is positively predicted by their References
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