Clinical Anatomy 00:00–00 (2014)

ORIGINAL COMMUNICATION

Recreation and Procreation:

A Critical View of Sex in the Human Female
ROY J LEVIN*
Sexual Physiology Laboratory, Porterbrook Clinic, Sheffield S11 9BF, England

This review deals critically with many aspects of the functional genital anatomy
of the human female in relation to inducing sexual arousal and its relevance to
procreation and recreation. Various controversial problems are discussed
including: the roles of clitorally versus coitally induced arousal and orgasm in
relation to the health of women, the various sites of induction of orgasm and
the difficulty women find in specifically identifying them because of “‘ambiguity
problems” and “genital site pareidolia,” the cervix and sexual arousal, why
there are so many sites for arousal, why multiple orgasms occur, genital
reflexes and coitus, the sites of arousal and their representation in the brain,
and identifying aspects and functions of the genitalia with appropriate new
nomenclature. Clin. Anat. 00:000–000, 2014. VC 2014 Wiley Periodicals, Inc.

Key words: G-spot; internal and external clitoris; periurethral glans; vaginal
tenting; cervix; typology of orgasms; genital orgasms; genital site
pareidolia; genital reflexes; evolution; just-so stories; urethral
emissions

INTRODUCTION GENITAL SEXUAL AROUSAL CHANGES

Some authors (Komisaruk et al., 2006) have argued
that overly stringent requirements should not be
applied to physiological processes in men and women
that are believed to play a role in reproduction and
that “much of the research on reproduction physiology
is inductive in the sense that any observed phenom-
enon is perceived as having a likely function.” Such
reasoning appears to support a scientific philosophy of
accepting mechanistic functional explanations without
empirical evidence. This leads to authors creating fan-
ciful but unscientific proposals (“just-so” stories) that
cannot be falsified (see the section below on “Does the
human female orgasm have a role in reproductive
fitness?”). This review of functional anatomy focuses
mainly, but not exclusively, on peripheral structures
involved in sexual arousal to orgasm in the human
female. It is deliberately critical and does not shy away
from focusing on and discussing many of the problems
of interpretation of published controversial studies and
asking pertinent questions. The hope is that its critical
focus will encourage more exacting investigations to be
undertaken in this most fascinating area of human
female biology. A companion review in this volume
(Georgiadis, in press) deals with the activation of
female sexual arousal and orgasm in the brain.
INVOLVED IN REPRODUCTION AND
PLEASURE
Humans undertake sexual activity for many rea-
sons; some 237 separate motivations were recorded
by Meston and Buss (2007). However, the changes
induced in the female genitalia by sexual arousal can
be conveniently categorized into those that subserve
reproduction, those that subserve pleasure, and those
that subserve reproduction and pleasure jointly. The
changes that appear to be related only to reproduction
include:
i. Increases in p02 (Wagner and Levin, 1978)
caused by enhanced blood flow to the vagina
(and probably to the uterus and fallopian
tubes). The blood flow in the vaginal microcir-
culation is controlled at the tissue level through
*Correspondence to: Roy J Levin.
E-mail: r.j.levin@sheffield.ac.uk
Received 19 November 2014; Accepted 19 November 2014
Published online in Wiley Online Library (wileyonlinelibrary.com).
DOI: 10.1002/ca.22495

C
V 2014 Wiley Periodicals, Inc.
2 Roy J Levin
“vasomotion,” an intermittent mechanism of
pseudorandom opening and closing of precapil-
lary sphincters through the relaxation and con-
traction of their smooth muscle, which is
sensitive to the levels of various vasodilator
metabolic products (viz. pCO2, lactic acid, low
pH, K1 and adenosine triphosphate). In the
basal state, vasomotion is maximal as most
capillaries are closed, but as sexual arousal
increases more and more become recruited and
open until all the vasomotion is disrupted and
all the capillaries are open and the vaginal
microcirculation is fully vasocongested (Levin
and Wylie, 2008). The enhanced pO2 facilitates
an increase in usable energy production by
spermatozoa.
ii. The increased vaginal surface pH due to
enhanced plasma transudation, brought about
by the neurogenically induced vasodilatation and
vasocongestion causing an increased production
of vaginal tissue fluid that leaks through the epi-
thelium to reside on its surface. This partially
neutralizes the acidic basal vaginal fluid (Wagner
and Levin, 1984), making the vagina more hos-
pitable to ejaculated spermatozoa.
iii. Vaginal tenting and ballooning during the late
excitation phase, as observed and filmed by
Masters and Johnson (1966) and confirmed by
others by filming, magnetic resonance imaging
(MRI), and electrical and pressure recordings
(see Levin, 2012 for references). During this
activity, the uterocervical complex is withdrawn
up into the false pelvis initially by contractions
of the pelvic striated muscles. However, these
fatigue quickly and the tenting is then main-
tained by smooth muscle contraction (Levin,
2003a). This elevates and removes the cervical
os from the path of the ejaculated semen and
its pooling in the ballooned upper part of the
vagina and critically delays any too-precipitate
transfer of spermatozoa. It also allows the coa-
gulated semen to liquefy, freeing the trapped
spermatozoa, and facilitating their contact with
various male and female genital fluids, allowing
complex precapacitation and capacitation
changes to be accomplished in the spermato-
zoa that subsequently enable them to achieve
the capacity to fertilize the ovum (Levin,
2011a, 2012). Remarkably, vaginal tenting has
been generally overlooked as a most important
reproductive function of the female sexual
response; without it, uncapacitated spermato-
zoa incapable of fertilizing an ovum would be
transferred to the fallopian tube, and thus,
wasted. A criticism of the role of tenting in
delaying sperm transport as an important
aspect of reproduction is that it is most effec-
tive only in the female knee-chest or so-called
missionary position. Coitus with ejaculation tak-
ing place by rear entry would allow semen to
fall on the cervical os. As Levin (2005a, b)
pointed out, “the answer is that no evolved bio-
logical mechanisms can be effective and ensure
reproductive fitness in every contingency.
Biological adaptation can service only one par-
ticular set of conditions; thus, there are always
selective compromises.” Another feature of
vaginal ballooning is that as it reduces penile
vaginal friction it delays ejaculation, allowing
maximal vaginal tenting to occur first.
Some further details of the role of sexual arousal in
human reproduction are described in Levin (2005a, b).
The changes relating to pleasure are:
i. Increased blood flow to the clitoris, vagina, and
labia creating vasocongestion and tumescence
in these structures, which sensitize them to
stimulation by the application of digital, lingual,
or penile friction/pressure leading to orgasm.
ii. Increased vaginal blood flow and its resultant
vasocongestion increasing the vaginal surface
temperature which, on penile penetration, is
appreciated as highly exciting by the male since
the skin surface of the erect penis is significantly
cooler (34.4 6 0.6 C, mean 6 standard error (SE)
[n 5 10]) than the vagina (37.3 6 0.1 C [n 5 6]):
Levin, unpublished results.
iii. Contractions of the female pelvic muscles at
orgasm, which can create pleasurable pressure
on the penis.
iv. Female vocalizations during sexual arousal and
orgasm that convey sexually explicit informa-
tion to the male coital partner (Levin, 2006a),
increasing his central sexual arousal and ardor
and enhancing sexual pleasure (hedonic
amplification).
Finally, the change that relates to both reproduction
and pleasure is:
i. Enhanced vaginal blood flow and congestion
(Levin and Wylie, 2008) that increases vaginal
transudation, thus, facilitating painless penile
penetration and subsequent pleasurable genital
thrusting for both males and females.
GENERATING THE FEMALE ORGASM
While the favored sites for generating orgasm in
women by friction, pressure, or vibration are the geni-
talia, many other sites and activities, some surprising,
can induce it. Early reports mention that orgasms can
be induced in some females by unusual nongenital
stimulation. Kinsey et al. (1953) describe women
being brought to orgasm by “having their eyebrows
stroked, or by having the hairs on some other part of
their bodies gently blown on, or by having pressure
applied on the teeth alone,” especially if some other
psychological stimulus is involved. However, in recent
years, a number of better-characterized studies have
revealed that the female orgasm is not linked solely to
stimulation of the genitalia or just to reproduction,
despite the claim, unsupported by evidence, that it is
the only sexual response linked by penile vaginal

intercourse (PVI) to reproduction (Brody, 2010) since
it is always possible for heterosexually partnered sex-
ual arousal per se to lead to PVI. Orgasm can also
arise from nipple/breast stimulation (Levin, 2006b;
Levin and Meston, 2006), anal stimulation (McBride
and Fortenberry, 2010; Komisaruk and Whipple,
2011), the mouth (Komisaruk and Whipple, 2011),
exercise (Herbenick and Fortenberry, 2011), hypno-
tism (Levin, 1992; Levin and Van Berlo, 2004), tantric
arousal (Lousad and Angel, 2011), drug side effects
(Levin, 2014a), and even childbirth (Pranzarone,
1991; Postel 2013). Some women claim to be able to
induce orgasm simply by thinking without any physi-
cal stimulation. Interestingly, such activity has a sig-
nificant history. Kinsey et al. (1953), in a footnote,
quoted some 12 authored papers on this topic, which
was given a variety of names: “idealized coitus,”
“moral or psychic masturbation,” “the mental vulva,”
and “erotic daydreaming.” It had been recorded as
early as 1903 (Bloch, 1902) from looking at nude
statues. Several of the quoted authors expressed the
curious and certainly unfounded opinion that this is
“the most noxious” of all forms of masturbation. A
more recent declaration, that clitoral stimulation to
orgasm creates “noxious consequences” (Brody,
2010), strangely echoes these past opinions. Whipple
et al., (1992) studied a number of women who
claimed that they could elicit orgasms by thinking
alone. In a later article by Komisaruk and Whipple
(2005), some brief comments were made about a few
of these subjects who underwent brain imaging during
their thinking arousal. Regions of the nucleus accum-
bens, the paraventricular nucleus of the hypothala-
mus, the hippocampus, and anterior cingulate cortex
were said to be activated by the “thought” of
orgasms. Whether these areas are truly the crucial
ones for generating orgasm (see Georgiadis, 2011, in
press) or whether such individuals fall outside the nor-
mal range are still unanswered questions.
Sexual intercourse with the penis alone does not
induce orgasm in a significant percentage of women
without additional stimulation of the clitoris (Lloyd,
2005; Levin, 2012). It is difficult to give an accurate
percentage because there is no standard metric for
comparing data as different studies have used differ-
ent measures to assess the frequency of such
orgasms (25 different metrics in 34 studies; Levin,
2012). Suggested reasons for this inability to attain
orgasm from PVI alone are contentious, controversial,
and often highly speculative, including clitorocentric
rather than vaginocentric sexual education (but see
Wade et al., 2005), early use of clitoral rather than
vaginal stimulation to obtain orgasm, not focusing
attention on the vagina during sexual arousal, too
short a duration of vaginal coitus, having partners
with small penises, and a clitoris anatomically too dis-
tant from the vaginal introitus (see Levin, 2012,
2014a for critical discussion and references). In
regard to the latter feature, two studies have investi-
gated the relationship between the clitoris and its ana-
tomical location. The most recent, by Oakley et al.
(2014), used MRI to compare the size and the location
of the clitoris in a small sample of 10 women with
anorgasmia and 20 controls. They found that the
Recreational and Procreational Sex 3
former had smaller clitorises (glans) with clitoral com-
ponents (body) further from the vaginal lumen than
women with normal orgasmic function. Wallen and
Lloyd (2011) had previously produced an interesting
and unusual statistical reworking of the possible rela-
tionship between the distance of the clitoris from the
urethral meatus (CUMD) and the generation of
orgasm from penile vaginal coitus alone using the
data of Narjani (1924) and Landis et al. (1940). Their
conclusion was that the shorter the CUMD, the more
likely the female would be to have orgasms from PVI
alone. When it came to the discussing explanatory
mechanism(s) for this connection the authors” focus
was mainly on the possible internal juxtapositions and
linkages between the clitoral complex and the anterior
vagina and the possible effects of androgens on the
development of the female genitalia in utero. How-
ever, a completely different mechanism that could
account for female orgasms induced solely by penile
vaginal thrusting is frictional/pressure stimulation of
the area of mucous membrane in the vaginal vestibule
surrounding the urethral meatus. This approximately
triangular area stretches from the upper part of the
vaginal vestibule and includes the tissue from the
underside of the clitoral glans to the upper edge of the
vaginal orifice. It was named the “female glans” by
Sevely (1987), but to make its delineation more spe-
cific and to avoid confusion with the clitoral glans, it
has been recharacterized as the “periurethral glans”
(Levin, 1991). This approximately triangular area in
the upper part of the vaginal vestibule surrounds the
urinary meatus from the Hart line junction delineating
the labia minora to the underside of the clitoral glans
and the upper edge of the vaginal orifice. It is ana-
tomically regarded as part of the female corpus spon-
giosum (van Turnhout et al., 1995), and according to
Kinsey et al. (1953) it has erotic sensitivity. During
filmed coitus, the area was observed to be pushed
into and then dragged out of the vagina by penile
thrusting so it was stimulated by friction/pressure
from the shaft of the sliding penis (Levin, 1991).
While Sevely (1987) briefly mentioned its role in the
generation of pleasure during coitus she did not make
the connection between its stimulation and the often
less-than-regular attainment of orgasm from PVI
alone. It has been proposed that orgasms from PVI
alone can be induced in women who have a highly
sensitive periurethral glans either because of the den-
sity of its innervation or because of enhanced sensitiv-
ity or both (Levin, 1991, 1992). The task now, as
proposed some years ago, is to use biothesiometric
examination to compare the sensitivity of this area in
women who can have orgasms from PVI alone with
that of women who do not. If the proposal is valid
then the former should be far more sensitive than the
latter. It would also be of some interest to examine
whether there is a relationship between the sensitivity
of the periurethral glans and the dimensions of the
CUMD.
There is little doubt that there are very contentious
issues regarding several aspects of female genital
anatomy involved in generating sexual arousal to
orgasm and in the supposed health attributes of
arousal and orgasm derived from PVI alone compared
4 Roy J Levin
to arousal and orgasms induced by clitoral stimulation
with or without PVI. The latter highly provocative and
controversial concepts, reinvoking early Freudian pro-
posals, rely mainly on correlations from cross-
sectional retrospective, self-answered questionnaires,
and as Brody and colleagues stated, “as in any corre-
lational study a universe of possible unmeasured fac-
tors could play a role in the observed findings”
(Nicholas et al., 2008). It was again acknowledged by
Brody (2010) that “In such research designs there is
always the possibility that unmeasured third variables
influence both the predictor variable and the outcome
of interest.” It should be noted that many of the con-
clusions from the studies of Brody et al. rely heavily
on correlation and causality. In this context, van Lank-
veld (2011), in his first message as incoming Editor-
in-Chief of the Annual Review of Sex Research
(ARSR), warned against inappropriate inferences of
causality in many research studies. He stated that
“most of the study designs that are used in sex
research are correlational experimental work that
allows for strong inferences regarding causing and
directionality that is, in many cases not possible or
extremely difficult to realize. This imposes major con-
straints on our scientific discourse about causes and
effects when writing about such research. Authors and
reviewers should be—and apparently are—critical of
the line of reasoning applied when reviewing such cor-
relational data.”
However, the earlier self-proclaimed caveats have
become less conspicuous in recent articles using simi-
lar survey protocols, which unfortunately have
become self-validating and alienated from less judg-
mental views of female sexual arousal. Various
aspects of the studies have now been extensively and
critically discussed by Levin (2011a, b, 2012, 2014a),
Prause (2011, 2012), Laan and Rellini (2011), Sus-
chinsky and Lalumière (2011), Suschinsky et al.,
(2013), and Suschinsky and Lalumière (2013).
FEMALE ORGASM TYPOLOGY—
FUNCTIONAL ANATOMY AT WORK
One of the more contentious aspects of the female
orgasm is its possible typology. The problem was initi-
ated when Freud (1905) pronounced that arousal
induced by clitoral stimulation was less “mature” than
that created by PVI alone. He believed that to achieve
his concept of normal femininity it was necessary to
de-eroticize the clitoris and eroticize the vagina. More-
over, he reported that this was a most important (and
difficult) transfer for women. For many years, he and
his followers held this belief with little or no empirical
evidence to support it. However, there were persistent
anecdotal accounts from many women that the
orgasms they obtained from PVI alone felt different
from those received from clitoral stimulation alone.
Fisher (1973) reported that women’s written descrip-
tions of their orgasms described the former as
“throbbing, deep, soothing and comfortable” while the
latter were “warm, sharp, ticklish and electrical.” Yet,
after their extensive studies of thousands of female
orgasms in their laboratory, Masters and Johnson
(1966) were of the firm opinion that only one type of
orgasm existed, that through clitoral stimulation;
“clitoral and vaginal orgasms were not separate bio-
logic entities.” They argued that even during PVI alone
the engorged clitoris was stimulated by penile traction
on the labia minora inducing a “to-ing and fro-ing” of
the clitoral hood over the glans. This mechanism was
later tested empirically by Alzate and Londono (1984).
They stimulated the lower third of the vaginas of
selected volunteers who were able to reach orgasm
easily from vaginal stimulation. Coital activity was
mimicked by digital stimulation. While there was trac-
tion on the labia pulling on the clitoral hood, only
slight to moderate sexual arousal was achieved, sug-
gesting that coital hood traction was not a major
cause of clitoral coital stimulation leading to orgasm.
However, it could be argued that the artificial and clin-
ical setting for such sexual stimulation was not condu-
cive to attaining a high level of arousal. Eichel et al.
(1988), originators of the coital alignment technique
to create orgasm during coitus, thought that the penis
could stimulate the clitoris and its surroundings when
the male mounted the female in a high position
(Levin, 2014a).
In support of the concept that the clitoris is stimu-
lated during PVI, but internally, other authors have
pointed out that: (i) the anterior wall of the vagina is
connected to the clitoral root internally by tendons
and that the thrusting of the penis in the vagina will
stretch these causing “internal” stimulation of the clit-
oris (Ingelman-Sundberg, 1997); (ii) the thrusting
penis stimulates the “root” of the clitoris via the ante-
rior wall (Buisson et al., 2010); and (iii) the mucous
membrane area surrounding the meatus of the ure-
thra (its external opening) in the upper part of the
vaginal vestibule, redesignated the “periurethral
glans” (Levin, 1991), will also be pushed into and
pulled out of the vagina by penile thrusting, thus,
causing sexual stimulation and, in sensitive women, a
coital orgasm.
It is accepted that the clitoris is one of the most
densely innervated organs in the female body and its
only known purpose is to create sexual arousal when
stimulated. In numerous studies (Schober et al.,
2004; Ladas et al., 2005; Waskul et al., 2007; Turn-
bull et al., 2013), the clitoris is named by females as
the most sensitive area of their genitalia and gives the
strongest orgasms and greatest sexual pleasure, while
the vagina, in this context, scores lower (Bronselaer
et al., 2013).
THE CERVIX
The putative role of cervical buffeting by the thrust-
ing penis during coitus as a trigger for sexual arousal
and orgasm is yet another contentious issue of female
sexuality. The literature falls into two quite distinct
sectors. One examines the effects of total hysterec-
tomy (favored in the United States and United King-
dom) or subtotal hysterectomy (favored in the
Scandinavian countries) on sexual arousal and
responses; the latter operation leaves the cervical
stump behind. The other sector observes and records

the behavior of the utero-cervico response during sex-
ual arousal. The clinical hysterectomy literature is
replete with conflicting studies, some showing that it
has no effect on female sexuality or is beneficial (Tha-
kar et al., 2002; Zobbe et al., 2004; Flory et al.,
2006; Kuppermann et al., 2005) while others record
negative sexual side effects (Maas et al., 2003). It
was reviewed by Grimes (1999) and by Levin (2005a,
b), who both concluded that the many observational
studies had obvious flaws and that no conclusion
could be drawn. Studies on the behavior of the utero-
cervical unit, conversely, show quite clearly that it is
elevated up into the false pelvis during the late excita-
tion phase of sexual arousal, away from the path of
the even the longest thrusting penis, and is, therefore,
not buffeted or stimulated during coitus. The tenting
and ballooning of the vagina creates an expansion
that “accounts for some loss of exteroreceptive stimu-
lation of the distal half of the fully inserted penis and
reduces sensate focus for the female. . .” the over-
distended excitement phase vagina gives many
women the sensation that the fully erect penis
(regardless of size) is “lost in the vagina” (Masters
and Johnson 1966). This expansion of the upper half
of the vagina has been confirmed by independent
studies using direct observation, filming, MRI, and
electrical and pressure recordings (see Levin, 2012,
for full references). Despite all these independent con-
firmatory studies, some authors erroneously claim
that “the penis repeatedly buffets the cervix uteri”
during coitus (Shafik et al., 2004a, b) and activates a
cervico-vaginal reflex that inhibits the tone and elec-
trical activity of the vagina. This conclusion was drawn
from their interpretation of ecologically invalid experi-
ments that used rubber-tipped steel rods to stimulate
the cervix. Brody (2012) also claimed, without empiri-
cal verification, that buffeting the cervix by the penis
during PVI alone creates a third type of orgasm and
that the cervix is able to distinguish between penile
buffeting and that induced by rods or vibrators! It
should be noted in these contexts that the sensory
innervation of the cervix is so poor that minor surgical
procedures on it can be carried out without anesthesia
(Levin, 2014a). The concept that a “long” penis is
needed to induce sexual arousal and orgasm by direct
cervical stimulation ignores the repeatedly confirmed
vaginal tenting, during which the utero-cervical unit is
elevated into the false pelvis during sexual activity
arousal away from the penile thrusting axis (see
Levin, 2012, 2014b for full critical discussion and
references).
THE G-SPOT
In the turbulent sea of controversy about the ana-
tomical sites for generating the female orgasm, the G-
spot is just one more wave, albeit a large one! Discov-
ered and then initially alluded to by Gra € fenberg
(1950) in an obscure journal ignored by most, then
“rediscovered” by Perry and Whipple (1981) and pub-
licized by Ladas et al. (2005), the G-spot has experi-
enced a checkered history. Many women claim to
have an area on the anterior wall of the vagina that
Recreational and Procreational Sex 5
when stimulated by pressure quickly creates intense
arousal leading to orgasm. Of course, it is a moot
point now as to whether it is the G-spot per se that
creates the arousal rather than the suggested
“anterior wall erogenous complex” (Hoch, 1986;
Levin, 2003a, 2012), which has also recently been
called the “clitorourethrovaginal complex” by Jannini
et al. (2014). The identity of the multiple anatomical
structure(s) creating this arousal is the problem (see
Table 1 for their possible identities). Some authors
simply argue that there is no convincing evidence that
any structure exists in and around the anterior vaginal
wall that could give rise to such responses (Hines,
2001; Burri et al., 2010; Puppo, 2012; Puppo and
Gruenwald, 2012; Jannini et al., 2014). Others dis-
agree and there have been proposals (Shafik et al.,
2004a, b), descriptions (Lenck et al., 1992) and even
published dissections (Thabet, 2009, 2013; Ostrzen-
ski, 2012) claiming to have identified its anatomy.
However, these approaches have identified different
structures at different sites. The dissection by Ostr-
zenski (2012) of a specific vaginal structure in an 83-
year-old female cadaver that he claimed to be the G-
spot complex elicited especially critical comments
concerning the lack of neural connections, histology,
and demographic details of the dead, postmenopausal
subject (Komisaruk et al., 2012; Levin and Wylie,
2012; Eichel and Ablin, 2013; Hines and Kilchevsky,
2013), to which the author replied with ill grace. The
criticisms appeared to have influenced a more recent
follow-up publication by Ostrzenski et al. (2014),
which describes similar putative “G-spot complexes”
dissected from eight female cadavers between 37 and
68 years of age (crucially, how many were premeno-
pausal is not stated). A structure found in the distal
anterior vaginal wall, internally some 4.5 cm from the
urethral meatus, averaged 7-mm long. The structure’s
basic histology and immunohistology appear to show
a neurovascular complex consisting of a number of
nerves and a claimed neuroganglion surrounded by
vascular tissue. It is unfortunate that the complex was
not studied by the modern stereological technique of
serial sectioning, which would have enabled an accu-
rate model to be constructed. Because all the studies
have been conducted on fresh cadavers who died
from violence, suicide, or drug overdose, no details of
the women’s sexual histories were available, so there
is no evidence that the females in question ever had
“G-spot” orgasms from PVI or from digital stimulation.
Indeed, it is not even known whether the women
were actually orgasmic. The authors’ highly specula-
tive suggestion that the structure is the “vaginal pace-
maker” for electrical activity (Shafik et al., 2004a, b)
involved in activating basal vaginal motility (Levin,
1980; Levin and Wagner, 1983) is obviously incorrect.
The reason is that in later papers (not cited, by Ostr-
zenski et al.), Shafik et al. (2005a, b) proposed that
as the electrical activity in other smooth muscle organ
systems was generated by the interstitial cells of Cajal
(ICC), the same should be the case for electrical activ-
ity in the vaginal smooth musculature. They identified
such cells in the muscle layers of vaginal tissue
obtained from 28 cadavers using a specific ICC
marker kit. A further study by Shafik et al. (2007)
6 Roy J Levin

TABLE 1. Female genito-pelvic erotogenic sites capable of inducing sexual arousal and ’genital orgasms’
(see text for details)

1. Anatomical description
Site inducing arousal 2. Coital stimulation Major ’Pro’ references Major ’Contra’ references
Clitoris (external) 2. Hood traction, friction from Masters & Johnson (1966)
hood, shaft, glans penile thrusts
2. Direct stimulation Shafik et al (2005)
Puppo (2011)

Clitoris (internal) 1. Anatomical dissection (cadavers) O’Connell et al (1998) Puppo (2011)

Crus 2. Penile (and digital) pressure Foldes & Buisson (2009) Puppo (2011, 2013)
thrusts on anterior vaginal wall Buisson et al (2010)
Buisson & Jannini (2013)

Clitoral bulbs (?) 1. Anatomical dissection (cadavers) O’Connell et al (1998)

2. Pressure from penile thrusts (?) Puppo (2011)

Periurethral glans 1. Anatomical description=dissection Van Turnhout et al (1995) Puppo (2011, 2013)
(Female corpus Schober et al (2004)
spongiosum) 2. Penile shaft thrusts pushing area Sevely (1987)
in and dragging it out of vagina Levin (1991, 1992)
Urethral meatus (?) 2. Friction from penile shaft Eichel, Eichel & Kule (1988)
Eichel (2000)

Cervix (?) 2. Penile thrusting jostling the cervix Singer (1973) Gra€ fenberg (1950)
Komisaruk et al (2006) Masters & Johnson (1966)
Brody & Weiss (2010) Grimes (1999)
Brody (2010) Levin (2011, 2012, 2014)

G-spot (?) 1. Description=Anatomical € fenberg (1950)

Gra Puppo (2011, 2013)
dissection (?) Lenck et al (1992) Hines (2001)
Thabet (2009, 2013) Puppo & Gruenwald (2012)
Ostrzenski (2012)
Ostrzenski et al (2014)
2. Penile thrusts ,or digital pressure, Hoch (1986)
on anterior vaginal wall Buisson & Jannini (2013)

Halban’s fascia 1. Anatomy=innervation Krantz (1959)

(vaginal 2. Stimulation of neural receptors Minh et al (1979) Puppo (2011, 2013)
bladder septum) by penile thrusts on anterior Hoang et al (1991)
vaginal wall Battaglia et al (2010)

Urethra 2. Stretching by penile thrusts on Levin (1992)

anterior vaginal wall
Labia (minora) 2 Friction from penile shaft
Masters & Johnson (1966)
Martin-Alguacil et al (2006)
Levin (2011)
Ginger et al (2011)

The Major ’Pro’ references are those that recognise, describe and support the site’s activity while the Major ’Contra’
references are those that are against the naming of, or the existence of, or activation of, the site(s) during coitus to
create sexual arousal. The added question marks in brackets indicate sites or activity that have been proposed but
need confirmation. Table modified from Levin (2001, 2011).

investigated the location of the neural vaginal pace-
maker by immunohistochemical and morphometric
methods and found that it was positioned in the circu-
lar muscle of the upper posterior vagina wall and not
in its upper anterior wall as claimed by Ostrzenski
et al. (2014). Another function claimed for the puta-
tive G-spot complex by Ostrzenski (2014) is its
involvement in the “genesis of anterior wall
ballooning.” This appears to be based on his descrip-
tion that the expansion of the structure “elevated the
anterior vaginal walls in each of the subjects.” The
article did not cite or discuss any previous studies that
related this vaginal “ballooning” and “tenting” to
contractions of the pelvic striated and subsequently
smooth muscles (Masters and Johnson, 1966; Levin,
2003c). This retraction of the upper part of the vagina
and elevation of the utero-cervical complex during
female sexual arousal was extensively described and
documented by Masters and Johnson (1966), who
created the term “tenting.” It is well-illustrated in the
film of the phenomenon by Wagner (1974). The sud-
den and rapid movements of vaginal tenting and bal-
looning as filmed cannot be induced by a small, less
than approximately 1 cm long, structure filling up with
blood; they are obviously of muscular origin. Inferring
physiological mechanisms of tenting and ballooning

from cadaveric dissections of women who were vic-
tims of violent or non-natural deaths is fraught with
complications and the risk of misinterpretation.
Despite the apparent confirming study of a putative
“G-spot structural complex,” we still have no empirical
knowledge that this structure is actually functional
and how/whether it is connected neurally to the spinal
cord/brain despite the ready, but clearly premature,
acceptance of such functionality by Miller (2014). It is
not unknown for human anatomical structures to exist
without any currently identifiable function, for exam-
ple, the vermiform appendix (Theobald, 2007) and
the vomeronasal organ (Levin, 2004). Independent
confirmation of the structural complex and its loca-
tion, its imaging in life and its functionality during sex-
ual arousal are all still clearly essential.
PUTATIVE FEMALE GENITAL REFLEXES
The neural control of genital function by the brain
and spinal cord was briefly described in a previous
section as mediated by the hypogastric, pudendal,
and pelvic nerves with the possible inclusion of the
vagus. There has also been a series of publications
describing nervous reflexes involving changes mainly
of the pelvic/genital musculature or blood flow acti-
vated in the laboratory in volunteer female control
subjects in their basal or sexually unstimulated state.
These have been reviewed previously (Levin, 2003a,
2007a) but none have been confirmed by independent
studies and they have been largely ignored in the lit-
erature on female sexual arousal. By far, the most
extensive series of papers are those from Shafik and
his coworkers (Shafik, 1993, 1995a, b; Shafik and El-
Sibai, 2001; Shafik et al., 2004a, b, 2005a, b). They
repeatedly claimed that these putative reflexes would
be useful in the clinical diagnosis of female sexual
dysfunctions (FSD), but none have been applied in
this way to date, viz. comparing the reflexes in FSD
patients with those in normal controls. This could be
because their assessment involves time-consuming
invasive procedures using skilled electrode insertion
placement techniques coupled with expensive elec-
tronic recording equipment. In fact, because they
were activated by artificial stimuli such as distending
the vagina with balloons, electrical stimulation, or
stimulation by foam-tipped steel rods and were,
therefore, not ecologically valid, it is not clear whether
they are actually activated and involved in influencing
genital functions during normal coitus, and therefore,
have true physiological utility.
OBNUBILATION, GENITAL AROUSAL
PAREIDOLIA, AND THE “AMBIGUITY
PROBLEM” DURING COITUS
Coitus, unlike external clitoral masturbation, is
clearly a multisite stimulus; there are now at least 10
possible genital sites that, when stimulated together
(or possibly even separately), can create sexual
arousal to orgasm during PVI (Table 1). Moreover,
during PVI, it is not possible for women in the throes
Recreational and Procreational Sex 7
of sexual arousal, especially if possessed by obnubila-
tion (clouding of consciousness), to discern and spec-
ify exactly which of these sites will create (are
creating) the arousal that will lead to orgasm, espe-
cially if they are asked long after sexual activity has
occurred. It is not unknown for subjects to make mis-
takes about their body reactions at orgasm in the lab-
oratory (Bohlen et al., 1982) and many become
unaware of their surroundings (obnubilation) during
sexual arousal (Kinsey et al., 1953). They even
greatly underestimate the duration of their orgasm
immediately after its occurrence (Levin and Wagner,
1985). This difficulty in delineating the activation
site(s) has been called the “ambiguity problem”
(Levin, 2011a, 2012), which Brody et al. have yet to
acknowledge; they continually ignore the problem,
making it more than problematical to interpret and
draw conclusions from their profuse, cross-sectional
questionnaire studies of women’s self-reports of geni-
tal sexual arousal. In a number of their studies, they
put much faith in the crucial use of a lie scale
(Eysenck, 1974) or questionnaire (Ballard, 1992) to
check whether their subjects are not simply answering
with sociably desirable responses or misrepresenting
behavior regarded as socially undesirable without
acknowledging that astute “dishonest” subjects could
well discern the scale or questionnaire’s purpose and
answer even this dishonestly! Strangely, while they
have taken care to check the validity of their
responses, they have never checked the core validity
of their studies; namely, whether women can actually
delineate their anatomical genital responses inducing
orgasm during sexual arousal with any certainty (see
also Prause, 2012 for discussion of this problem).
Singer (1973) raised this very same problem when he
stated that “Tactile stimulation by the male genitals or
body pressing against the labia minora, the clitoris, or
the vestibule of the vagina alone would provide suffi-
cient stimulation to bring most females to orgasm.
The location of this stimulation may be correctly rec-
ognized or it may be incorrectly attributed to the inte-
rior of the vagina.” Notice that the vestibule of the
vagina was included, and we now can specifically
delineate it as the periurethral glans. In some
respects, women’s attribution of arousal to a specific
site despite multiple-site activation during PVI is akin
to a form of pareidolia, a known behavior that occurs
in situations where humans try to interpret unclear/
vague signals as clear and distinct. It is most often
applied when people think they can see images in
clouds, rock faces, burnt toast, and so forth. In the
female sexual context, it can be described as “genital
site arousal pareidolia,” that is, ascribing their arousal
to a particular anatomical site. The women are not
delusional, just mistaken. It is impossible to know the
percentage of women in the many studies of Brody
et al. who were unconsciously, and thus, inadvertently
involved in such “genital site arousal pareidolia” when
making their responses. However, just as there is a
questionnaire to assess whether participants in such
surveys are using socially acceptable answers, there
is also a critical need, since the condition is now
spelled out, to develop a questionnaire that can
assess the possible role of “genital site pareidolia” in
8 Roy J Levin
responses by women to the question of what genital
site(s) cause(s) their arousal/orgasm during PVI.
Interestingly, the condition is known to be especially
amplified by any clear bias, in their case, unbalanced
leading questions that assume that individuals can
isolate the focus of their coital sexual arousal to a spe-
cific anatomical location.
THE CONCEPT OF A FEMALE “GENITAL
ORGASM” FROM PVI REPLACING THAT
OF THE NOW-OUTDATED “VAGINAL
ORGASM”
Given the complexity of the known genital struc-
tures that could be involved in creating coital sexual
arousal, describing orgasm during PVI as “purely”
vaginal is now clearly anomalous. Interestingly, when
Jannini collected the opinions of five researchers
about the female orgasm during PVI (Jannini et al.,
2012), he did not describe it as the “vaginal orgasm”
but used the more circumspect “vaginal activated
orgasm (VAO).” Twenty-eight years previously, Hoch
(1986) had proposed that because PVI activated a
number of “trigger sites” (then far fewer than are now
listed in Table 1), instead of looking for a PVI orgasm
different from a clitoral one, the continued stimulation
of these multiple genital “trigger” sites by PVI created
a “genital orgasm.” A very similar conclusion was
reached by Gillespie (1993) who argued that the
“evidence seems overwhelming—it is a physical
impossibility to separate the clitoral from the vaginal
orgasm as demanded by psychoanalytical theory.” He
proposed that “vaginal orgasm” should be used to
describe an orgasm that could be generated from
both vaginal and clitoral sites simultaneously by PVI.
Similar ideas were presented in a previous review
(Levin, 2012). The time has now come to accept fully
the concept and abandon the use of “vaginal
orgasms” and even VAOs for those generated by PVI
as—first and foremost—they are in reality “genital
orgasms.”
DOES THE HUMAN FEMALE ORGASM
HAVE A ROLE IN REPRODUCTIVE
FITNESS?
Since Dobzhansky’s (1973) famous dictum that
“nothing in biology makes sense except in the light of
evolution,” developments in studies of human female
sexuality have seen attempts by authors to interpret
and explain their findings, both behavioral and physio-
logical, in terms of evolutionary concepts (see for
references Symons, 1979; Lloyd, 2005; Georgiadis
and Kringelbach, 2012; Gray, 2013; Gray and Garcia,
2013). However, there is a distinct danger of too-
eager authors making up fanciful “just-so” stories, as
referred to in the Introduction. It should be noted that
a few praise such stories as “goads to further thought”
despite their lack of supportive data and empirical val-
idation (Barash and Lipton, 2009). The great difficulty
with such stories is to winnow the “good goads” from
the profusion of published chaff. Levin (1992) listed
nine speculative explanations for the female orgasm
culled from the literature, which included ending coi-
tus to redistributing the potentials in the brain to cre-
ate psychological resuscitation. Rancour-Laferriere
(1983) created four, Barash and Lipton (2009) listed
12, Lloyd (2005) dealt with 21 relevant papers while
Brody et al. created three. While it has been said “Of
the making of books there is no end” (Ecclesiastes
12:12), the same can be said of female orgasm just-
so stories!
Early proposals for a role of the female orgasm in
reproduction argued that it was an adaptation to
enhance the transport of spermatozoa by increasing
both the rate and number moved into the uterus and
fallopian tube (see Lloyd, 2005 for references). This
claimed evolutionary adaptation was supposed to
facilitate the female’s reproductive fitness (success).
The enhanced transport was said to be mediated by
contractions of the uterus induced by the oxytocin
released at orgasm, which supposedly created a
sucking-up of the semen—the inelegantly termed
“upsuck mechanism” (see Levin, 2011a for referen-
ces). Linquist (2006) wrote that “to show that the
female orgasm is most likely not an adaptation would
require good evidence contradicting the ‘upsuck’
hypothesis, which is altogether absent.” That position
cannot now be sustained. In reviews entitled “The
human female orgasm: a critical evaluation of its pro-
posed reproductive functions” (Levin, 2011a) and
“Can the controversy about the putative role of the
human female orgasm in sperm transport be settled
with our current knowledge of coitus?” (Levin,
2011b), enough good factual and experimental evi-
dence is presented and referenced to demonstrate
clearly and explicitly that the oxytocin released sys-
temically during the human female orgasm does not
feature in the so-called “upsuck mechanisms”
involved in sperm transport by inducing contractions
of the uterus (see also Leyendecker et al., 2004), and
that the arousal phenomenon of vaginal tenting
removes the cervix from the ejaculated pool of
semen. Moreover, there is no empirical (or even theo-
retical) evidence for the need, in the sexually aroused
human female, for an increase in either the rate or
the number of spermatozoa transported to the fallo-
pian tube during orgasm from natural coitus. Females
actually try to reduce the numbers of spermatozoa
transported because of the problems of polyploidy
(more than one sperm entering the ovum) and degen-
eration of the ovum through the release of sperm
enzymes during their acrosome reactions; both fea-
tures reduce fertility (Levin, 2011b).
Some authors are diehard protagonists for the role
of the female orgasm as an adaptation to facilitate
reproductive success through the postulated “sire-
selection mechanism,” also known as “mate choice
selection” (Puts et al., 2012). In their extensive analy-
sis of the possible reproductive role(s) of the female
orgasm in the section headed “Female orgasm may
promote conception,” these authors still interpret the
results of a number of studies, especially including
those in animals, to indicate that oxytocin is involved
in sperm transport. These latest interpretations were

presented despite acknowledgment that the published
major criticisms are valid: (i) all the human studies
trying to mimic the actions of orgasm-liberated oxyto-
cin used doses of exogenous oxytocin far higher than
those experienced during female orgasms (acting on
the uterus for periods of 40 min instead of the few
seconds at orgasm); and (ii) all were undertaken on
women in their basal, nonsexually aroused state
(Levin, 2011b). Such features invalidate any conclu-
sions from studies of the physiological role of systemi-
cally released oxytocin at orgasm in sperm transport.
Strangely, however, the authors claimed that these
criticisms did not collectively undermine the brain
stimulation research (they) reviewed, which sug-
gested that the “female orgasm increases sperm
transport.” Unfortunately, the only human study that
they quoted in this context was that of Komisaruk
et al. (2004), a study of five women with complete
spinal transections. Not only is this condition known to
change physiological sensitivities both in the brain and
at the periphery (Henderson et al., 2011; Levin,
2011b), its authors also applied supraphysiological
stimuli to the cervices of the subjects using custom-
made rods. This stimulation is not ecologically valid as
in natural coitus, in the sexually aroused woman, the
penis does not normally stimulate the cervix owing to
vaginal tenting (Levin, 2011a, b, 2012) activated
according to Shafik et al. (2005a, b) by the clitoro-
uterine reflex through penile clitoral stimulation. Geor-
giadis (2011) also questioned the study because of its
technical shortcomings. Furthermore, while animal
studies are of interest, because species differences
are particularly frequent in the field of reproduction
(Zuk, 2002; Baum, 2006; Dixson, 2009), results from
them may only illustrate mechanisms for that species
and can never be the final arbiter of support for
human physiological mechanisms. For example, in the
female rabbit, ovulation is induced by coitus; coital
locking of the penis in the vagina occurs in dogs; in
rats, the semen is ejaculated into the uterus; while
during coitus in pigs, the penis enters and is locked
into the uterus and then deposits the semen; in rats,
stimulating the cervix causes extreme immobilization;
all fascinating mechanisms of reproduction but none
of these occur in human females during normal coitus.
Last, we cannot be sure that animals experience
human-type orgasm because we cannot ask them
what they feel.
An even more bizarre extension of the “mate
choice mechanism” is surely that of Costa et al.
(2012). They propose that it evolved during the late
Pleistocene period (some 2,588,000–11,700 years
ago), when the fertility from coitus of girls and women
was compromised because they were either young
and subfertile, or adult and lactating, or in the nono-
vulating part of their cycle. This, they speculate,
allowed them to undertake coital activity, sampling
the penis sizes of various sexual partners, which
“open the way for differential rates of female orgasm
with different male lovers to influence male choice
and fertility.” Such behavior would have needed the
women of that time to know that ovulation occurs in
midmenstrual cycle and to have had a remarkable
understanding of safe periods, orgasm functions,
Recreational and Procreational Sex 9
inhibition of ovulation by lactation, and so forth. It has
apparently not occurred to the authors (or unfortu-
nately the referees!) that correct understanding of
such physiological mechanisms began to appear only
200 years ago and was finalized only over the last
100 years (Cobb, 2007; Gray and Garcia, 2013). This
is an excellent example of the classic error of
“colonizing the past,” that is, using the knowledge of
today to explain the behavior of humans hundreds of
thousands of years ago. There is no evidence for
these behavioral claims. It is no wonder that such pro-
posals are widely and pejoratively considered “just-
so” stories, like the fairy tales created by Rudyard Kip-
ling (1902) for little children’s amusement (Schlinger,
1996).
As previously discussed, direct sexual arousal to
orgasm in the human female can arise from a number
of sites with high erotic potential when they are
stimulated, usually by friction (stroking) or pressure,
as listed in Table 1. The literature fails to explain the
abundance of sites in the female at which direct physi-
cal sexual arousal can putatively lead to orgasm if the
stimulus is sustained and is facilitated when such
stimulation is desired. Interestingly, it can still occur
even if the stimulation is nonconsensual (Levin and
Van Berlo, 2004). The maintenance of so many sites
(blood flows, nervous innervations, tissue cellular
metabolism) incurs a significant metabolic energy
cost, so in view of the usual parsimony of evolution,
there must be a reason(s) for maintaining this prolif-
eration. Contemporary views of biological processes
almost always interpret their functioning in terms of
evolutionary benefit-cost. Structures or mechanisms
are maintained because they offer reproductive or life
advantages to their possessors, and thus, allow them
to breed more successfully. Bearing these concepts in
mind, the burden of multiple sites and mechanisms
for inducing sexual arousal in the human female
should clearly carry a reproductive advantage (Levin,
2007a, b). However, this is incompatible with the sur-
prising claim that only PVI and its orgasm convey any
health benefits and advantages; all other methods for
activating sexual arousal are claimed to cause
“noxious outcomes” (Brody, 2010). The outstanding
unanswered question is: why would evolution make
the most sensitive structure in the female genitalia,
viz. the clitoris (Bronselaer et al., 2013), which has
the highest density of innervation, and therefore,
needs the least energy to activate orgasms of the
greatest intensity (Schober et al., 2004) and is
actively potent from young girls to aged, postmeno-
pausal females, become the source of “noxious out-
comes” (Levin, 2014a)?
Interestingly, of all the genital abnormalities found
in human female neonates, the lack of clitoral devel-
opment is exceptionally rare. According to Gold-
schmidt et al. (2009), only a few cases have been
reported in the world literature. If the clitoris was not
useful and could easily be dispensed with then it
would probably have a high incidence of absence or
nondevelopment among female newborns. Interest-
ingly, conditions exist where even such basic genital
organs as the uterus and upper vagina are absent, for
example, the Meyer–Rokitansky–Ku € ster–Hauser
10 Roy J Levin
syndrome, which occurs in one of every of 4,000–
10,000 women, caused by embryonic growth failure
of the Mu € llerian ducts (Guerrier et al., 2006). Why
should there be such a remarkably low incidence of
clitoral nondevelopment? Is it just fortuitous or does it
indicate that the mechanisms for the development of
the clitoris are carefully preserved, even “ring fenced,”
because the organ is of major importance (Levin,
2014a)? Yet no one has shown it to have any function
other than creating sexual pleasure. It is more than
strange that there are conditions/syndromes in which
the uterus/vagina is dispensed with but not the clito-
ris. Yet another question that has never been
answered by those who repeatedly demonize the clit-
oris (see Levin, 2011a, 2012, 2014a for references) is
“Why should this organ’s function be so highly con-
served if its activation leads to such ‘noxious out-
comes’?” In fact, to isolate and pathologize clitoral
stimulation as leading to “noxious outcomes” is non-
sensical, as has been pointed out previously (Levin,
2012): every source of sexual pleasure comes with a
cost/benefit tariff. Even consensual PVI, for example,
can create female genital injuries leading to pain and
infection, various venereal diseases (HIV, gonorrhea,
syphilis, trichomoniasis, chlamydia, herpes simplex
virus, infertility from a number of these, genital warts,
cervical cancer through papilloma viral infections,
hepatitis) and last, unwanted pregnancies, with 44
million abortions carried out worldwide resulting in
some 79,000 related maternal deaths (Aitken, 2014).
All have far greater consequences for the individual
female than any of the putative nonlethal conditions
listed from the papers of Brody et al. by Levin
(2014a); so much for their conclusion that “evolution”
rewards those who undertake PVI rather than clitoral
stimulation.
DO THE DIFFERENT PERIPHERAL
SITES OF SEXUAL STIMULATION
CREATE DIFFERENT BRAIN
ACTIVITIES IN RELATION TO
PLEASURE AND REPRODUCTION?
The various pelvic sites that can generate sexual
arousal (Table 1) have different neural innervations
connecting them to the spinal cord and brain (see also
Georgiadis, in press). According to Komisaruk et al.
(2006), the major organs are innervated via three
main nerves, with the possible inclusion of the vagus
as a fourth. The hypogastric nerve conveys sensory
information from the uterus and cervix, the pelvic
nerves innervate the vagina and cervix, and the
somatosensory pudendal nerve the clitoris. The vagus
nerve, not previously thought to innervate the female
genitalia, has also been claimed to mediate the sen-
sory innervation of the cervix. This has been con-
cluded from interpretations of brain images obtained
during self-stimulation in totally spinalized women
(Komisaruk et al., 1997, 2004). However, Georgiadis
(2011) was critical of these studies and suggested
that independent verification is needed before definite
acceptance of the innervation is warranted. Georgiadis
and Kringelbach (2012) detailed that great care
should be taken in evaluating and interpreting brain
imaging studies, especially given the lack of ecological
validity of the scanner environment. While brain imag-
ing has allowed real advances to be made in uncover-
ing the workings of the brain, scanning has been
likened to discovering what is going on in the office by
looking at whether the lights are on. We are a long
way from being able to identify the brain areas really
involved during arousal and especially during orgasm.
It has been claimed from another brain imaging study
that different but overlapping areas of the somatosen-
sory cortex are activated by separate stimulations of
the cervix, anterior vaginal wall, and clitoris, using
inanimate rods for the former two sites and self-
masturbation for the latter (Komisaruk et al., 2011).
As this is one of the few empirical studies that Brody
et al. herald as firm objective evidence that vaginal
and clitoral orgasms are physically/mentally different,
and thus, can have different effects on the woman’s
mental and physical health, the possible functional
significance of the overlapping of sites has been con-
veniently ignored, for example, that they could
actually be interconnected (also see Georgiadis, in
press for a critical discussion of cortical–genital repre-
sentation). However, Levin (2012) pointed out that
the study had no ecological validity as the cervix is
not stimulated by the thrusting penis during normal
coitus. Moreover, the mechanical jostling/buffeting of
the cervix by the rods used, in the study, would create
stimuli of supranormal physiological intensity. Accord-
ing to the criterion set by Brody (2010), such genital
stimulation with inanimate objects used “non-living
and emotionally unresponsive dildos,” again consist-
ent with this lack of ecological validity. Costa and
Brody (2011) also argued that “sex with a non-living
object” was most associated with their proposed
immature defense mechanisms and insecure attach-
ment. Strangely inconsistent, given this and his previ-
ous claims about other noxious effects of clitoral
masturbation, Brody obviously acquiesced to the
recruitment of the female subjects who would have to
perform cervical and clitoral masturbation as arousal
stimuli during the experiments. If these recruited sub-
jects habitually used clitoral stimulation to obtain
arousal and orgasm in their private sexual behavior
(not reported), the site(s) on the somatosensory cor-
tex could well have been influenced/induced (?) by
such behavior. No women who had never previously
used clitoral/cervical stimulation for arousal were
involved as controls. One further and obvious problem
with the interpretation of these experiments is that
the results from separate, independent stimulations of
the three pelvic sites were used to mimic brain condi-
tions at orgasm during PVI with its multiple simultane-
ous stimuli (viz. the sites in Table 1). Yet Komisaruk
and Whipple (2011) reported with their illustrative
“Tapestry brain scan” that practically the whole brain
becomes activated and involved during arousal to
orgasm. Komisaruk (2012), himself has admitted that
“all major brain systems evidently contribute to wom-
en’s orgasm.” Such evidence clearly indicates that the
claimed “separate site representations” of cervix,
anterior vaginal wall, and clitoris cannot tell us

anything about possible mental/behavioral differences
between clitoral and cervical/vaginal orgasms from
natural coitus. Indeed, in regard to the cervix, what
has been completely overlooked is that the cervical
neural links to the brain probably have no involvement
in sexuality per se but rather involvement in the phys-
iological mechanisms of parturition. Oxytocin is
released when the neural circuits of the cervix are
stretch-activated by the ejecting fetal head. The
supraphysiological stimulus of the rods on the non-
pregnant cervix could cause neural stimulus
“spillover” into the sexual areas of the brain. There
has been no comment from the various authors about
such difficulties of interpretation of their laboratory
experiments compared to ecologically valid, real-life
coital experiences.
WHY CAN WOMEN BUT NOT MEN HAVE
MULTIPLE SERIAL ORGASMS?
It has been known for a long time that women are
able to have repeated, serial orgasms within very
short intervals of time, but this is not normally possi-
ble for men. The postejaculation refractory time
(PERT) in men occurs immediately after ejaculation
with its concomitant orgasm; males cannot have
another erection, ejaculation, and orgasm for a finite
time. This period varies, being short (minutes) in
young men but becoming longer and longer with
aging until it lasts for many hours. While we have
data for the duration in young men there are none for
old males (see Levin, 2009 for details).
All current possible central and peripheral explana-
tions for PERT were reviewed critically (Levin, 2009);
the conclusion was that none satisfactorily explained
its mechanism(s). Two recent flawed attempts to
explain the activity are no more than unsupported
speculations. One implicated stretch receptors in the
seminal vesicles as the culprits (Turley and Rowland,
2013) but did not report that the possible involvement
of these structures was discussed in a previous review
and that animal experiments in which the seminal
vesicles were removed failed to show any evidence
that the procedure affected PERT. The other (Pazhoohi
and Salehi, 2013) speculated that the peptide gonad-
otropin inhibitory hormone causes PERT by activating
the secretion of prolactin. The possible role of prolac-
tin was discussed in previous reviews and the hor-
mone was found to be highly unlikely to cause PERT
(Levin, 2003b, 2009, 2011a, b).
In women, both prolactin and oxytocin are released
into the systemic circulation at orgasm but clearly
they do not prevent multiple serial orgasms (see Levin
2003b, 2011a, b for references). It has been sug-
gested that the difference between men and women
in regard to PERT is that women do not normally have
a urethral emission of fluid at orgasm (often referred
to as female ejaculation). However, some women do
have such an emission of fluid, which is not urine
(Pastor, 2013). The orgasms that occur with urethral
fluid emission are claimed to be more intense that
those without (Davidson et al., 1989). Unfortunately,
while a few authors report anecdotally that women
Recreational and Procreational Sex 11
who exhibit orgasmic urethral emissions appear to
have a refractory period, there are few if any pub-
lished empirical studies of whether such women expe-
rience PERT routinely (Levin, 2009). We do not know
what causes sexual satiation in women.
HYPNOTICALLY INDUCED
ORGASMS—REALITY OR FAKES?
Female orgasms induced hypnotically have been
described in only a few published reports (Hoenig and
Hamilton, 1960; McVaugh, 1979). However, numerous
web sites show female subjects being hypnotized and
then induced to have “orgasms.” The subjects display
the obvious physical behavioral signs of orgasm, viz.
extensive vocalizations together with multiple pelvic
thrustings and general myotonia, which can continue
for up to a few minutes. On being questioned after the
hypnosis they report having mentally experienced
high levels of orgasmic sensations. It is clearly not
possible to confirm that these subjects are experienc-
ing a genuine physiological orgasm; some could be
faking the behavior. However, in one hypnotized sub-
ject, it was possible to count the contractions of the
anus and vulva (n 5 13) during on 15-s orgasm
induced under hypnosis. Using a vibrator on her clito-
ris, the same subject exhibited 13 such contractions
during her 16 seconds of clitorally induced orgasm,
indicating near-identical duration, and muscle activa-
tion for both stimuli. Another subject who had an 80-s
orgasm induced by hypnotism reported afterward that
“it was different from those obtained from my ‘private
parts’ as there was more vibration that coursed
through everything.” A third female had her electro-
cardiogram (ECG) recorded to assess her heart rate.
In her basal state, this was 65–70 beats/min while
during her 15-s hypnotically induced orgasm, it regis-
tered 99 beats/min. While these examples suggest
that a number of the aspects of orgasm appear to be
inducible in females by hypnosis, critical experiments
would be to examine such subjects during fMRI and
compare the imaging obtained with that induced by
clitoral vibration, and to record the vaginal and anal
contractions induced during the two stimuli.
IN DEFENSE OF DESCRIBING AND
NAMING “NEW” FUNCTIONAL
ANATOMICAL ASPECTS OF THE HUMAN
FEMALE GENITALIA
Over the years, a number of authors have
reported on, or referred to, newly described
“functional anatomical” aspects of the human female
genitalia and have often labeled both the structures
that create these functions and the functions them-
selves with new, distinct names (Table 1). These
have included the G-spot, the periurethral glans, Hal-
ban’s fascia, female ejaculation (emission), vagal
innervation of the cervix, internal clitoris, and persis-
tent genital arousal disorder (PGAD). These designa-
tions have repeatedly been the targets of near-
12 Roy J Levin
identical dismissive accounts and letters to journals,
unfortunately often expressed intemperately, assert-
ing that such descriptions have no “scientific basis”
and should not be used by “gynecologists, sexolo-
gists, sexual medical experts, women, and mass
media” (Puppo, 2011a, b, 2013; Puppo and Puppo, in
press). Most of the terminology and designations
criticized were developed because their authors
found that the accepted descriptions of female
arousal by classical anatomists were lacking in spe-
cific areas (where new appearance meets the eye,
new suppositions thereupon arise); and despite the
claim by Puppo, they normally included an empirical
or observational study in support of their new
description. The concept of a “scientific basis” simply
means that a proposal must, by the generally
accepted Popperian criterion of “scientific,” be falsifi-
able or refutable (Popper, 2004). In fact, all of the
quoted designations have at least one possible falsifi-
able aspect, making them “scientific.” In brief, the
evidence against the so-called G-spot as an anatomi-
cal entity is not as clear-cut as Puppo would have us
believe, as a number of studies have claimed to iden-
tify it anatomically (see the section on the G-spot).
The periurethral glans (the triangular area surround-
ing the urethral meatus from the anterior edge of the
introitus to under the clitoral glans) has been filmed
being pushed into and out of the vagina by the
thrusting penis during PVI (Levin, 1991, 2012) and
has been identified as a sexually arousing area (Kin-
sey et al., 1953). This structure has always been
there but has been overlooked as a potential source
of erotic excitement. It now has a putative described
function and a designated label (see also the section
above on generating female orgasm). Halban’s fascia
(lying in the urethrovaginal septum) was described
by Minh et al. (1979), identified surgically during
prolapse operations (Hoang et al., 1991), and
imaged with its inclusions using ultrasound (Battaglia
et al., 2010). Female ejaculation (better described as
female urethral emission) has been described and
observed by a number of researchers (Darling et al.,
1990; Wimpissinger et al., 2007; Rubio-Casillas and
Jannini, 2011). The vagal innervation of the cervix
has been inferred from experiments on women with
spinal transection where mechanical stimulation of
the cervix by plastic rods has activated a site in the
brain as observed by imaging (Komisaruk et al.,
2004, 2006). The internal aspects of the clitoris have
been dissected in cadavers (O’Connell et al., 1998)
and imaged in vivo using ultrasound during coitus
(Foldes and Buisson, 2009; Buisson et al., 2010;
Buisson and Jannini, 2013). PGAD is now a defined
clinical entity, first described in 2001 (Leiblum and
Nathan, 2001) and recognized and diagnosed by
numerous clinicians who treat patients (Facelle et al.,
2013), and its vaginal activity has been observed in
laboratory recordings (Levin and Wylie, 2008). It is
not the same as clitoral priapism (Puppo, 2014;
Puppo and Puppo, in press). The clitoris does not
become erect like the penis because its structure is
not built to create rigidity (see van Turnhout et al.,
1995). The correct description of vasocongestion of
the clitoris is “tumescence,” not erection. In PGAD,
there are feelings of sexual arousal, often close to
orgasm. Furthermore, women with PGAD often
attempt masturbation to relieve themselves of the
orgasmic urge; this does not occur with a clitoris that
is painfully engorged—so-called clitoral priapism.
Thus, there is sexual arousal, notwithstanding the
claim in a letter by Puppo (2014) that if genital
arousal is unwanted the term “arousal” is inappropri-
ate. It is used because PGAD sufferers describe such
feelings. Puppo has no right to devalue what PGAD
patients tell us in clinics during history-taking. It has
been associated with overactive bladder and restless
leg syndrome (Waldinger and Schweitzer, 2009). A
severe case of PGAD, treated by electrocortical treat-
ment (ECT), caused the immediate disappearance of
the condition but it slowly relapsed until the next ECT
(Korda et al., 2009). PGAD is indeed a separate
entity from priapism.
Against the above brief, descriptive resume of sup-
porting—and importantly refutable—empirical studies,
the criticisms leveled by Puppo (2011a, b, 2012),
when examined, are mainly of semantics and rely on
a few repeated stock phrases, some out-of-date ana-
tomical text book references and inaccurate physio-
logical descriptions. Moreover, he presents no quoted,
definitive empirical studies that invalidate the func-
tional interpretations by their proponents. Puppo
(2011b) even resorts to sophistry. He declaims that
“The whole clitoris (glans, body, roots, or crura) is an
external genital organ; the glans and body are visible
while the roots are hidden, therefore they are not
internal.” If an anatomical structure is “hidden,” that
is totally underneath the skin and can only be reached
by dissecting away the skin and muscle barrier, then
ipso facto, it is internal compared to the observable
external glans and hood, so it is perfectly valid to
describe the clitoris as having internal and external
parts.
As has been pointed out, changes or additions to
nomenclature in a particular subject area usually indi-
cate dynamic developments and focus attention on
these. Moreover, by their very nature, they will always
be ahead of conservative anatomy committees and
dated textbooks (Levin, 2011a). If such changes help
to delineate specific aspects of female sexual function
in better detail and are thus, found useful they will be
adopted and survive; if not, they will simply fade
away. Examples of such adoption are the prescient
diagrams of the extent of the internal aspects of the
clitoris in the book “A Review of a Woman’s Body”
published in 1981 (Federation of Feminist Women’s
Health Centres, 1981), which initially had no impact
on the anatomy textbooks of the period, while exam-
ples of those ignored include Singer (1973), whose
characterization of the female orgasm as uterine,
vulval, and mixed (utero-vulval) was never fully
adopted by others, and the amazing claim of Puppo
(2011a) that “To describe the cluster of erectile tis-
sues (i.e., clitoris, vestibular bulbs, and pars interme-
dia, labia minora, corpus spongiosum of the female
urethra), the correct anatomical term should be the
female penis’! Despite chastising authors for using
neologisms for anatomical structures and functions,
Puppo (2013) himself creates new terminology for the

erectile cycle of the male, abandoning the internation-
ally accepted Excitation, Plateau, Orgasm, Resolution
model terms created by Masters and Johnson (1966)
and replacing them with his own inventions—latent,
tumescence, rigid, or muscolar (?) and detumescence
phases. The utricle of the prostate gland is renamed
“the male vagina”! He characterizes the tumescence
of the clitoris erroneously as “erection”—this term is
inaccurate as there is no rigidity—and erroneously
asserts that the tonic contractions of the male and
female perineal muscles “squeeze blood into the cor-
pora cavernosa from the roots.” Gerstenberg et al.
(1990) showed with their electromyographic activity
recordings of the bulbocavernosus and ischiocaverno-
sus that these muscles in the male are not involuntar-
ily involved in erection. Does the old proverb “People
in glass houses should not throw stones” apply here?
UNANSWERED QUESTIONS—THE
PROBLEM OF SUPERSEDING THE
ADULT FEMALE’S GENITAL SEXUAL
PLEASURE PATHWAYS
Freud, later supported by Brody et al., proposed
that the clitoral orgasm is immature and needs to be
replaced by the “PVI-induced orgasm” for mature
femininity. According to the latter group, avoidance of
its putatively noxious actions raises many important
and unanswered questions about human female sex-
uality (for discussion see Laan and Rellini, 2011;
Levin, 2011a, 2012, 2014 a, b; Prause, 2011, 2012).
Are the neural mechanisms/pathways for female sex-
ual pleasure “hard wired” in the brain/spinal cord at
birth waiting to be activated later? If they are not
“hard wired” then what causes their initiation and sub-
sequent development? Do they have to be generated
specifically by genital stimulation? Is the pleasure of
stimulating the genital/clitoris area discovered acci-
dentally by random hand exploration, which is then
repeated to gain pleasurable rewards (see Clifford,
1978; Wade et al., 2005; Waskul et al., 2007)? As
there is anecdotal evidence that initial orgasms are
not as pleasurable or as satisfying as those of later
years (Levin, 1981), there appear to be elements of
practice, experience, and learning in perceiving and
appreciating sexual pleasure. In fact, this is not a fea-
ture unique to sexual pleasure, as many of the other
pleasures of life—food, music, humor, and art—are all
enhanced with age and concomitant increases in
experience (Georgiadis and Kringelbach, 2012: see
companion review, Georgiadis, in press). Sex in this
company is no different, but what makes sex appa-
rently distinct is that the neural path(s) that have
been used or created during youthful clitoral mastur-
bation, often the first initiator of female sexual pleas-
ure, have to be either bypassed or superseded by
conscious effort using vaginal stimulation, but only by
PVI, with no other sexual stimulation (Levin, 2014b).
This suggests that the original “clitoral” pathway(s)
must be exceptionally strong and functionally domi-
nant because, according to Brody and Costa (2008)
and Brody (2010), if clitoral stimulation occurs during
Recreational and Procreational Sex 13
PVI, and even sometime before, the claimed benefits
of PVI do not occur; clitoral stimulation simply trumps
PVI. This is more than surprising considering that PVI
is claimed by them to be the only sexual activity that
can lead to gene propagation, and thus, should be
more highly rewarded (Brody, 2010). It would have
been thought that through evolution, the PVI neural
path(s) would have been of extreme durability and
resilience. However, this does not seem to be the case
as the clitoral neural pleasure pathways in the spinal
cord and input to the brain appear to be markedly
more resilient and more powerful, even according to
their most ardent critics! They have offered no expla-
nation of this remarkable anomaly. Clearly, many
aspects of the human female’s sexual pleasure path-
ways have been significantly glossed over. Despite the
numerous studies, publications and commentaries on
human female sexual arousal and orgasm, it remains
a fascinating and contentious topic where there is still
much to study and understand.
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