COMMENTARY AND PERSPECTIVE

Origin, Function, and Effects of

Female Orgasm: All Three are
Different
GÜNTER P. WAGNER1,2,3,4
AND MIHAELA PAVLIČEV
5
1
Department of Ecology and Evolutionary Biology, Yale University, New Haven, Connecticut
2
Yale Systems Biology Institute, West Haven, Connecticut
3
Department of Obstetrics, Gynecology and Reproductive Science, Yale Medical School, New
Haven, Connecticut
4
Department of Obstetrics and Gynecology, Wayne State University, Detroit, Michigan
5
Cincinnati Children’s Hospital and Medical Center, Cincinnati, Ohio

J. Exp. Zool. How to cite this article: Wagner GP, Pavlicev M. 2017. Origin, function and effects of female
(Mol. Dev. Evol.)
00B:1–5,
orgasm: All three are different. J. Exp. Zool. (Mol. Dev. Evol.) 00B:1–5.
2017

In a commentary published in the Journal of Experimental Zo- reproductive success, and we acknowledge that fact in our pa-
ology B: Molecular and Developmental Evolution (Komisaruk, per. Yet it is important to note that the present evidence for an
2016), Dr. Komisaruk criticized our recent paper (Pavličev and adaptive function of FO (i.e. one that influences reproductive
Wagner, 2016) on the evolutionary origin of female orgasm (FO) success, fitness) ranges from weak to nonexistent. The earlier
by claiming to have identified “unfounded assumptions” and literature has been scrutinized by Elisabeth Lloyd in her influ-
“misleading” conclusions. While we acknowledge the pioneering ential book (Lloyd, 2005) with largely negative results and does
contributions of Dr. Komisaruk to the physiology of FO, we think not need to be reanalyzed here. Never the less, it is worth not-
that the disagreements expressed in Komisaruk (2016) reflect the ing that the evidence for an adaptive function of FO has not
difficulties inherent to communication across different scientific improved since 2005. Two papers that appeared in the interim
disciplines, especially between that of neurophysiology and evo- need to be mentioned, one on the question of whether FO en-
lutionary biology. In our opinion, most of the disagreements are hances sperm transport (Levin, 2011) and one that directly tests
based on two interrelated issues regarding evolutionary expla- whether there exists a correlation between the ability of women
nations in general: the nature of homology and the distinction to have orgasms during intercourse and the number of children
between the origin of a character and its subsequent evolution. these women bear (Zietsch and Santtila, 2013).
In addition, we disagree on the best way forward for this field of One of the more popular adaptive hypotheses to assign a bi-
research. We begin with the question of whether there is com- ological role to FO is the idea that the contractions caused in
pelling evidence for an adaptive function of FO in humans and the female reproductive tract by FO may aid in sperm transport
then proceed by setting our replies in the context of two broader and thus could increase the chance of fertilization. In a relatively
thematic issues: the nature of biological characters and how to recent review, Levin (2011) compared the claims about the pu-
explain the evolutionary origin of characters. tative function of FO in fertilization with the known physiology
of the female reproductive tract and concluded that “the bulk of
reported evidence favors the conclusion that the female orgasm,
Function, Effect, and Adaptive Function of FO
…, has little or no effective role in the transport of spermatozoa
There has been a long-running controversy about the func-
in natural human coitus.”
tion (i.e., the current use) of FO in humans. This is the rea-
son why philosophers, biologists, and anthropologists have
discussed FO at length for decades (reviewed in Lloyd (2005)). Reply to comments by Dr. Barry Komisaruk
The one undisputed fact is that FO is not necessary for repro- Conflict of interest: None.
duction. Women can conceive and bring a child into the world Received 9 November 2016; Revised 22 February 2017; Accepted 22
without ever having had an orgasm, in particular an orgasm dur- February 2017
DOI: 10.1002/jez.b.22737
ing heterosexual intercourse. However, this does not imply, and
Published online in Wiley Online Library (wileyonlinelibrary.com).
here we agree with Dr. Komisaruk, that FO does not influence


C 2017 WILEY PERIODICALS, INC.
2 WAGNER AND PAVLIČEV

But of course the most important question is whether women Functions and Origins
that have higher rate of coital orgasms have higher reproductive This train of thought leads us to a frequent source of misun-
success than women with a lower rate of coital orgasm. This derstandings about evolutionary explanations that concerns
question was explicitly tested by Zietsch and Santtila (2013). In the association between the function or biological role of a
a sample of 8,000 twins and siblings, the authors tested for a trait and its evolutionary origin. In one school of thought,
correlation between coital orgasm rate (COR) and the number of the function of a trait also implies a model for its origin. The
children, and found a near zero correlation (r = –0.01 in iden- reasoning goes like this: a trait comes into existence because of
tical twins, and r = 0.04 among nonidentical twins or siblings). natural selection. Natural selection fixes heritable phenotypic
Interestingly among identical twins, there is a moderate intra- variation if these variants have a positive impact on fitness,
class correlation in orgasm rate (r = 0.25) and in the number that is the average reproductive success of individuals that
of children (r = 0.41), which suggests a genetic component to carry this trait. Based on this reasoning, the function of a
the likelihood of coital orgasm for women, and a genetic com- trait is, at the same time, the reason for its existence. Many
ponent of fertility (for details, please see Zietsch and Santtila decades of evolutionary research, however, have shown that
(2013)). Even though some of the correlations between orgasm this view involves a problematic assumption. Any counterex-
rate and reproductive success are statistically significant, due ample suffices to demonstrate the poverty of this approach in
to the large sample size, their effect size leaves ample room for evolutionary biology. For instance, it is clear that feathers of
skepticism about their biological importance. birds are useful and even necessary for flight in birds. Does
Dr. Komisaruk cites an older paper by Sam Baxter (Baxter, ’74) this fact allow us to conclude that feathers originated because
that neither we nor Elisabeth Lloyd have cited and which Dr. they are good for flight? A large amount of paleontological and
Komisaruk claims provides evidence for a biological function of developmental evidence shows that flight was not the reason
FO. The claim that Dr. Komisaruk attaches to this paper is that for the origin of feathers, and that feathers originated long
women who had no orgasm had a longer second stage of partu- before any theropod dinosaur even thought of flying (Prum,
rition, that is, the critical extrusion phase. First, we need to note ’99; Prum and Brush, 2002). By the same token, the search for
that this paper did not set out to test the hypothesis for which a current biological role of FO in humans does not guarantee
Dr. Komisaruk cites the paper. The question pursued in Baxter that we will find the explanation for its origin. Functional roles
(’74) is whether childbirth leads to a decrease in sexual desire of biological traits can change; characters can originate for
and responsiveness in women. Never the less, from the tables one purpose and then acquire a new one, or lose its function
presented in this paper one might come to the conclusion that completely.
Dr. Komisaruk derives from that data. In Table 2 of Baxter (’74), Given the paucity of evidence for an adaptive role of FO in
COR is related to the duration of the second phase of parturition humans, Dr. Komisaruk and co-authors propose that we should
(SPP). If the data area aggregated, then one can find an apparent not give up hope and ignore alternative explanations (Chapter
difference in SPP duration, where women classified as “orgasmic 2 in Komisaruk et al. (2006)). We think that there are plausible
prepregnancy” have, in this sample, an average SPP of 41 min alternatives, one of which is the side effect hypothesis supported
while women classified as “nonorgasmic” women have a SPP by Symons (’79) and Lloyd (2005); the other is the hypothesis
of 70 min. That may look like a significant difference, but the that FO is homologous with and derived from a functional trait in
data provided do not allow for a statistical assessment, because the deep evolutionary past of mammals: male-induced ovulation
the within-class variation is not documented. Never the less one (Pavlicev and Wagner, 2016). We will not summarize this model
can estimate a standard deviation for the sample of “orgasmic” here again. We only want to emphasize a few points that may
women that comes to about 36 min, and 46 min for the “nonor- have been lost in the conversation.
gasmic” women. Hence, the estimated standard deviations are of First, we have explicitly kept the door open for the possibil-
the same magnitude as the difference in SPP duration. Further- ity that what in humans is now called FO has acquired a non-
more, in clinical practice, the expected duration of SPP is about reproductive role in human biology and psychology, for instance
1 hr with a 95% percentile (considered normal) of 140 min, i.e. in pair bonding or other functions. Second, we cannot empha-
2 hr and 20 min (Beckmann et al., 2014). This means that SPP size enough that hypotheses about the origin of a character do
duration is highly variable and even a seemingly large differ- not depend on the presence or the type of current function for
ence may not be significant. Finally, Baxter also showed in the that character. Even though we eagerly anticipate additional re-
same paper that the SPP duration is correlated with the num- search about this question, and even if a positive result is found
ber of psychiatric symptoms exhibited by the women, such that in the future, these functions would not necessarily explain the
the difference between orgasmic and nonorgasmic women in the evolutionary origin of the physiological machinery on which
duration of SPP could be confounded by other variables. Taken FO was grafted. We maintain that evolutionary derivation from
together the data published by Baxter give little if any support copulation-induced ovulation is a plausible model with stronger
for an adaptive role of FO in humans. support than the adaptive models like sperm transport and sperm

J. Exp. Zool. (Mol. Dev. Evol.)

ORIGIN, FUNCTION, AND EFFECTS OF FEMALE ORGASM 3

competition in spite of a much longer history of research on Here, we need to briefly explain our vocabulary using a sim-
these models. ple example. Let us consider the case of forelimbs, a widespread
Finally, we want to point out that the discussion of function morphological characteristic present in quadrupeds. The fore-
and evolutionary origins pertains to a deeper issue called the limb itself is a character, a developmental module, and it is easy
“logic of research questions” by Lloyd (2015). The way a research to see corresponding homologues are present in humans, dogs,
agenda defines its research questions has serious and far reach- mice, whales, frogs, and birds. However, the forelimb character
ing consequences for how research is actually pursued. Lloyd ar- was modified in different lineages in different ways, and thus a
gues that adapationism asks the question: “What is the function wing represents the character state of the forelimb that is present
of this trait?” or “What adaptive explanation can account for this in birds. As a consequence, we would not require wing-specific
trait?” But this framing of the questions precludes the search for characteristics when asking whether a dog’s foreleg is homolo-
and even the validity of other (nonadaptive) explanations. In gous to the bird or bat wing.
fact, it has been noted by many proponents of this approach This distinction between the origin of a new character, and
that nonadaptive explanations can only be considered when re- its lineage specific modifications is not semantic, but material
peated and exhaustive searches for adaptive/functional expla- and has important consequences for the evolutionary biology of
nations have failed (Mayr, ’83; Alcock, ’87; Sherman, ’89; Reeve FO. In our paper (Pavličev and Wagner, 2016), we assume, and
and Sherman, ’93). This approach denies that there can be pos- empirically support, that human FO, as generally described, is a
itive empirical evidence for nonadaptive models, which clearly character state of a much older physiological system that was
is not the case. We believe that the position that Dr. Komisaruk responsible for male-induced ovulation. The exact extent of this
and colleagues have defended in 2006 is akin to methodologi- character, which parts of the brain, peripheral nervous system,
cal adaptationism and inherits its shortcomings. Lloyd proposes and endocrine organs are part if the character, is not known at
an alternative approach she calls the “evolutionary factors ap- this point.
proach.” In our original paper (Pavličev and Wagner, 2016), we The methodology we used to empirically test this assumption
use a similar approach called “homology thinking” (Ereshefsky, will be discussed further below. The reason why we thought that
2007; Ereshefsky, 2012, Wagner, 2016). We turn to this perspec- FO is more likely to be a character state than the character itself
tive in the next session. is that it seems unlikely that the limited effect of FO on repro-
ductive success can explain the origin of the complex sensory–
Homology, Character Identity, and Character States neuroendocrine machinery underlying the surface phenomenon
In his commentary Dr. Komisaruk starts out by noting that we of FO.
did not “provide a definition of female orgasm” and ignore the A frequent misunderstanding of evolutionary biology is the
evidence that “orgasm in women involved activation of all ma- assumption that a statement of homology between characters
jor brain systems.” We are well aware that there are many at- in different species implies even a remote degree of similarity.
tempts to define orgasm, all of which heavily rely on subjective Homology is the hypothesis that there is a morphological, de-
experiences and to some degree on peripheral signs of excite- velopmental or functional module that is derived from the same
ment (muscle spasm and accelerated heart rate and breathing), module in the most recent common ancestor of the two species
tension, and its release (Meston et al., 2004). However, none of compared. Note that all we ask for here is historical continuity
the published “definitions” amount to anything like a definition of an individualized part of the body (Wagner, 2014); no spe-
in the technical sense. At most they can be seen as “descriptive cific degree or kind of similarity is implied. For physiological
characterizations” of what one calls an orgasm in women. None functions, a similar reasoning applies (Love, 2007). Hence, our
of these definitions are suitable for a broader comparative study hypothesis that FO in humans is homologous to some part of fe-
of the evolutionary roots of FO. We thus started our investigation male physiology found in other mammals, copulation-induced
by noting objective neuroendocrine events triggered by FO and ovulation, does NOT carry with it any a priori expectation of
proceeded by testing the evolutionary predictions of our model. an easily recognizable similarity. The very root of the homology
More importantly, these “definitions” of FO raise two unan- concept clearly states that homology is about “the same organ in
swered questions: different species regardless of form and function” (Owen, 1848).
This has been accepted biological thinking since over 150 years.
(1) Is the so “defined” orgasm expected to identify a unitary Against this background, it is also clear that we do not claim
physiological or developmental module (character) that that any other mammal experiences FO in the same way as a
originated in the human lineage and is therefore uniquely human female does. They may, but we remain neutral, due to a
present only in humans? Or, lack of positive evidence, with respect to whether that in fact is
(2) Is what is called FO a derived character state of a much the case.
older physiological system with, potentially, completely Parenthetically, and contrary to Dr. Komisaruk’s claim, we
different biological role in other mammals? do provide empirical evidence for our hypothesis. Regardless of

J. Exp. Zool. (Mol. Dev. Evol.)

4 WAGNER AND PAVLIČEV

whether phylogenetic evidence is familiar in neurophysiology, origin of FO. The evolutionary history of female reproductive
it is never the less an accepted form of scientific evidence biology in mammals is too deep and complicated that a sim-
(Sober, 2008). We were focusing on two predictions of our ple unitary explanation based on (largely undocumented) adap-
model. First, we tested whether male-induced ovulation is tive advantages is biologically plausible. We do acknowledge the
ancestral to spontaneous ovulation in the sense that ovula- point, raised by Dr. Komisaruk, that much is not known which
tion proceeds independently of external stimuli. Second, we would be important to further test our scenario and anticipate
tested the hypothesis that the configuration of female external incorporating new information into future tests.
genitalia, as presented in primates, is a derived condition
that evolved coincidentally with the evolution of spontaneous ACKNOWLEDGMENT
ovulation (for details, see Pavličev and Wagner, 2016). The authors thank Professor Alan Love for helpful comments on
the draft of this paper.
Anatomical Details
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