Vaginal Lubrication

Vaginal lubrication is an indicator of tissue health and increases significantly during

genital sexual arousal, facilitating sexual intercourse.

From: The Journal of Sexual Medicine, 2010

Related terms:

Dyspareunia, Sexual Behavior, Sexual Dysfunction, Orgasm, Sexuality, Estrogen,

Vagina

View all Topics

Vaginal Dryness
Myrtle Wilhite MD, MS, in Integrative Medicine (Fourth Edition), 2018

Physiology of Vaginal Lubrication

Vaginal lubrication is reliant upon healthy blood flow because vaginal lubrication is
ultrafiltrated blood.

• The vagina contains no glands.

• Blood pressure pushes fluid from the capillaries through intracellular gap
junctions between vaginal epithelial cells.7 The resultant vaginal transudate
is mainly composed of water and very small proteins, which combine at the
vaginal surface with dead epithelial cells.
• Sufficient pelvic blood flow is dependent on the bioavailability of nitric
oxide (NO).4 Gaseous NO is produced in capillary endothelia in response to
shear stress or vibration or in response to sexual arousal via parasympathetic
nitrergic nerves.9 Once produced, NO induces vasodilation through a cyclic
guanosine monophosphate (cGMP) cascade, which diminishes as phosphodi-
esterase enzymes subdue the effect.
• Therefore, vaginal lubrication production depends upon the synthesis, enzy-
matic facilitation, and bioavailability of NO (Table 59.2). The enzymatic func-
tion of NO synthase is enhanced by steroid hormones, most notably estrogen
in a rapid-action nongenomic effect.10
• Importantly, the presence of NO is not sufficient for its effect. Many bio-
logical feedback mechanisms exist to suppress the production of NO as high
production of NO in an inflammatory environment can lead to irreversible free
radical production.11 Metabolic conditions of low inflammation are supportive
to the bioavailability of NO in facilitating vaginal lubrication.

The vagina’s lubrication does not come from glands but from a transudative fluid
expressed through vaginal epithelial cells from filtered blood that requires adequate
perfusion and NO.

> Read full chapter

Sexual Behavior
J.D. Baldwin, J.I. Baldwin, in Encyclopedia of Human Behavior (Second Edition), 2012

Theoretical Interlude
This article began by stating that a developmental perspective on sex is theoretically
valuable, and we can now explain why. Tracing the emergence of sexual behavior
from infancy onward helps us understand how nature and nurture interact. Both
biological and learning processes are involved in the gradual emergence of sexual
behavior, and developmental studies reveal how nature and nurture interact and
entwine with each other.

In the past, many people tended to fall into the trap of wanting to explain sexual
activities – and many other types of behavior – as ‘completely due to biology’ or
‘completely due to learning.’ This is still seen when people begin arguing for either
‘nature’ or ‘nurture’ in the frequently held discussions about nature versus nurture.
Modern science avoids this ‘either/or’ logic by focusing instead on ways in which
biological, learning, and social inputs interact to shape our thoughts and behavior
– which in turn are mediated by multiple neural systems that operate at both the
conscious and unconscious levels.

Studies on sexual behavior allow us to intertwine data and theories that incorporate
biology, psychology, and sociology. Men and women are biologically different in
many ways and some of these differences influence their thoughts, emotions, and
behavior significantly. People's location in society also affects their thoughts,
emotions, and behaviors. The simplest way to integrate all the elements that shape
sexual behavior is to follow a developmental time line that starts with birth and traces
the following decades of maturation, learning, and socialization.
We have already given descriptive information about several features of prepubertal
sexual development. It is time to detail the key elements of the nature–nurture
interactions, beginning where this article started.

Reflexes
Both vaginal lubrication and penile erection are based on sexual reflexes, which are
biologically built into the body's neural and cardiovascular systems. From infancy,
stimulation of the genitals activates these reflexes – causing the sexual organs to
swell – and elicits pleasurable feelings in the reinforcement centers of the brain.
These positive emotions are the primary reinforcers which lead to various types of
learning.

Operant Learning
The pleasurable feelings associated with sex serve as reinforcers that can strengthen
behavior, through a process called operant learning. Reinforcers explain why an
infant's early random touching of the genitals can become more frequent and
eventually a habit. In common parlance, this type of learning is often referred to
as ‘trial-and-error’ learning, although the word ‘trial’ is a bit misleading. When
infants are first touching their genitals, they are not ‘trying’ to get sexually excited
or reach orgasm. They are merely exploring with their hands and they accidentally
discover the pleasures associated with sexual stimulation. Infants have no intentions
of discovering sexual pleasure, but when early random explorations lead to pleasing
feelings, positive reinforcement increases the chances that, on later occasions,
exploring infants will repeat those types of self-stimulation that were pleasurable
in the past.

As infants touch themselves in different ways, the more rewarding forms of

self-stimulation become ‘habits’ and the less rewarding ones gradually become less
frequent. The result is that infants become increasingly skillful at self-stimulation,
and adults begin to call the behavior masturbation. Of course, babies do not have a
word for masturbation; hence, they cannot verbalize or subjectively understand the
words adults use to describe the activity.

Operant learning derives its name from the fact that some behaviors (such as touch-
ing) operate on the world in ways that may produce consequences. There are two
kinds of consequences that affect operant learning: reinforcement and punishment.
The pleasure that a child experiences when touching the genitals provides positive
reinforcement for repeating the activity. To reinforce means to ‘strengthen,’ and
reinforcers strengthen behavior (much as steel rods can be used to strengthen
concrete, to produce reinforced concrete). The pleasures of genital stimulation are
called ‘primary reinforcers’ to indicate that they are biologically ‘wired in’ as part of
the sexual reflexes. This shows how biology and learning interact, helping to avoid an
‘either/or’ approach to the nature–nurture debate that emphasizes the importance
of one element and devalues the other.

There are ‘primary punishers,’ too; and they are also biologically ‘wired in.’ Hitting
the testicles or ovaries is painful for biological reasons: it endangers these important
reproductive organs, and we have evolved to find such experiences aversive. The
primary punishers of pain help us learn to avoid hurting our ovaries and testicles
when possible.

Not all reinforcers and punishers are ‘primary,’ meaning biological, in origin. Social
reinforcement for exploring sex with others can occur when a child's peer group
rewards taking a dare about some sexual activity. As we have already seen, social
punishment can occur when caregivers punish a child for masturbation; the stronger
the punishment, the more the inhibitions it produces.

Rule-Guided Learning
Young infants can learn how to masturbate, even though they cannot put words
on the behavior or think about it in symbolic terms. Once children begin to use
language, they can take verbal information and use it as ‘rules’ to hasten operant
learning. If a teen frequently hears about the importance of using condoms to reduce
the risks of becoming pregnant or contracting sexually transmitted diseases (STDs),
the words can serve as rules that help the teen avoid the errors that too many young
people still experience through ‘trial-and-error’ learning. Rules can be helpful in
guiding people to learn more effective bedroom techniques, relationship skills, and
much more. Having positive consequences strengthens the habits of continued rule
use.

Observational Learning
Even when no words are spoken, people can learn operant activities by watching
others, through a process known as observational learning. For example, sex ther-
apists encourage people to observe how their partner masturbates to see what they
can learn from these observations. It can help both observers increase the range
of techniques they can use together. When a girl sees her best girlfriend getting
pregnant, this observation can be a big wake-up call about the need for using
contraception. Movies, DVDs, books, and the Internet make it easy to learn about
sex, although not all the information is accurate. Much of our behavior is learned by
observing others, and sex is no exception.

Pavlovian Conditioning
While operant learning affects the frequency of behaviors that operate on people
and things, Pavlovian conditioning alters the reflexes – which evolveto respond to a
narrow range of stimuli. Pavlovian conditioning allows inborn reflexes to be linked
with new stimuli in ways that we did not evolve to experience. Because salivation
helps start the digestion of food, dogs (and many other species) have evolved to
salivate when they smell food. By ringing a bell before presenting food, Ivan Pavlov
conditioned dogs to salivate to the sound of a bell, thereby expanding the range of
stimuli that elicit salivation.

The pleasurable feelings of the sexual reflexes can be associated with all sorts of
stimuli via Pavlovian conditioning. We are biologically prepared such that touch to
the genitals triggers the sex reflex, but many other stimuli can become associated
with sex via Pavlovian conditioning. A young woman who pays attention to her
boyfriend's hairy chest while having sex may come to find hairy chests to be a sexual
‘turn-on’ or ‘erotic stimulus.’ Later she may notice that she becomes sexually aroused
when she sees other men who expose hairy chests.

Whatever people are thinking about when genital touch elicits the sex reflexes can
become a sexual turn-on. A young man who masturbates while watching pornog-
raphy is pairing new stimuli with the sex reflex, and he may end up feeling sexually
excited by seeing or thinking about such activities in the future. Even if he does
not masturbate while viewing porn, he may think about the pornographic images
when he masturbates at a later time, thereby sexualizing those stimuli. If he asks his
girlfriend to do some of the things shown by the porn videos, his girlfriend may be
happy to oblige; but some girlfriends are surprised that their boyfriends would even
think about enacting porn scenes, and this can create problems for the couple.

Masculinization of the Brain

During a critical period of fetal development, boys' testicles produce a burst of prena-
tal testosterone, which alters their brains in ways most females do not experience. As
a result, boys and men tend to prefer somewhat higher levels of rough-and-tumble
play, rowdy activities, and vigorous physical movements, compared with girls and
women. There is a great deal of variation in optimal activity levels in both males
and females, but males tend to prefer higher activity levels than females do. These
differences often persist well into adulthood. This does not mean that boys cannot
learn to control themselves and play gently with others, but many males continue
to prefer higher levels of activity and sensory stimulation than do females well
into adulthood. This contributes to a commonly observed gender difference in the
activities that males and females gravitate toward. For example, it is easier for males
than females to learn to love football, soccer, and dirt biking.
There are genes that lead some males not to secrete enough testosterone to fully
masculinize the brain, and this may predispose these males to become homosexual.
The biologically heritable component of male homosexuality is about 50%. Lesbians
appear to have about a 25% heritable component to their sexual orientation. Person-
al learning, culture, and many other variables complement the heritable influences
in the development of homosexuality.

In summary, both biology and learning are important in the development of sexual
behavior and both are intertwined starting from infancy. The following sections show
how nature and nurture continue to interact after the onset of puberty.

> Read full chapter

Sexual Behavior☆
John D. Baldwin, Janice I. Baldwin, in Reference Module in Neuroscience and
Biobehavioral Psychology, 2018

Theoretical Interlude
This chapter began by stating that developmental perspectives on sex are theo-
retically valuable, and we can now explain why. Tracing the emergence of sexual
behavior from infancy onward helps us understand how nature and nurture interact.
Both biological and learning processes are involved in the gradual emergence of
sexual behavior, and developmental studies reveal how nature and nurture interact,
naturally entwining with each other.

In the past, many people tended to fall into the trap of wanting to explain sexual
activities—and many other types of behavior—as “completely due to biology” or
“completely due to learning.” This is still seen when people begin arguing for
either “nature” or “nurture” in the frequently held discussions about nature versus
nurture. Modern science avoids this “either/or” logic by focusing instead on ways
in which biological, learning and social inputs interact to shape our thoughts and
activities—which in turn are mediated by multiple neural systems that operate at
both the conscious and unconscious levels.

Studies on sexual behavior allow us to intertwine data and theories that incorporate
biology, psychology and sociology. Men and women are biologically different in
multiple ways, and some of these differences influence their thoughts, emotions
and behavior significantly. People's location in society also affects their thoughts,
emotions and behaviors. The simplest way to integrate all the elements that shape
sexual behavior is to follow a developmental time-line that starts with birth and traces
the following decades of maturation, learning and socialization.

We have already given descriptive information about several features of pre-pubertal

sexual development. It is time for a theoretical interlude to explain the key elements
of the nature-nurture interactions, beginning where this chapter started.

Reflexes
Both vaginal lubrication and penile erection are based on sexual reflexes which are
biologically built into the body's neural and cardiovascular systems. From infancy,
stimulation of the genitals activates these reflexes, causing the sexual organs to swell
and eliciting pleasurable feelings in the reward centers of the brain. These positive
emotions are the primary reinforcers which lead to various types of learning.

Operant Learning
The pleasurable feelings associated with sex serve as reinforcers that can strengthen
behavior, through a process called operant learning. Reinforcers explain why an
infant's early random touching of the genitals can become more frequent and
eventually a habit. In common parlance, this type of learning is often referred to
as “trial-and-error” learning, though the word “trial” is a bit misleading. When
infants are first touching their genitals, they are not “trying” to get sexually excited
or reach orgasm. They are merely exploring with their hands and they accidentally
discover the pleasures associated with sexual stimulation. Infants have no intentions
of discovering sexual pleasure, but when early random explorations lead to pleasing
feelings, positive reinforcement increases the chances that, on later occasions,
exploring infants will repeat those types of self-stimulation that were pleasurable
in the past.

As infants touch themselves in different ways, the more rewarding forms of

self-stimulation become habits and the less rewarding ones gradually become
less frequent, due to the lack of reinforcement. The result is that infants become
increasingly skillful at self-stimulation, and adults begin to label the behavior as
masturbation. Of course, babies do not have a word for masturbation, hence they
cannot verbalize or subjectively understand the words adults use to describe the
activity.

Operant learning derives its name from the fact that some behaviors (such as touch-
ing) operate on the world in ways that may produce consequences. There are two
kinds of consequences that affect operant learning: reinforcement and punishment.
The pleasure that a child experiences when touching the genitals provides positive
reinforcement for repeating the activity. To reinforce means to “strengthen,” and
reinforcers strengthen behavior (much as steel rods can be used to strengthen
concrete, to produce reinforced concrete). The pleasures of genital stimulation are
called “primary reinforcers” to indicate that they are biologically “wired in” as part
of the sexual reflexes. This shows how biology and learning interact, helping avoid
an “either/or” approach to the nature-nurture debate, hence reducing the chances
of exaggerating the importance of one element and devaluing the other.

There are “primary punishers,” too; and they are also biologically “wired in.” Hitting
the testicles or ovaries is painful for biological reasons: It endangers these important
reproductive organs, and we have evolved to find such experiences aversive. The
primary punishers of pain help us learn to avoid hurting our ovaries and testicles
when possible.

Not all reinforcers and punishers are “primary,” meaning biological in origin. Social
reinforcement for exploring sex with others can occur when a child's peer group
rewards “taking a dare” about doing some sexual activity. Social attention is often
a reinforcer. As we have already seen, social punishment can occur when caregivers
punish a child for masturbation; and the stronger the punishment, the more inhi-
bitions it produces.

Rule-Guided Learning
Young infants can learn how to masturbate, even though they cannot put words
on the behavior or think about it in symbolic terms. Once children begin to use
language, they can take verbal information and use it as “rules” or “guidelines” to
hasten operant learning. If a teen frequently hears about the importance of using
condoms to reduce the risks of becoming pregnant or contracting sexually transmit-
ted diseases, the words can serve as rules that help the teen avoid the errors that too
many young people still experience through “trial-and-error” learning. The Internet,
sexually-oriented magazines and books provide many instructions for exploring sex.
Rules can be helpful in guiding people to learn more effective bedroom techniques,
relationship skills and much more. If the rules lead to rewarding experiences, the
positive outcomes strengthen the habits of continued rule use.

Observational Learning
Even when no words are spoken, people can learn operant activities by watching
others, through a process known as observational learning. For example, sex ther-
apists encourage people to observe how their partner masturbates to see what they
can learn from these observations. Such learning can help both partners increase the
range of sexual techniques they can use together. When a girl sees her best girlfriend
get pregnant, this observation can be a big wake-up call about the need for using
contraception. Movies, DVDs, books and the Internet make it easy to learn about
sex through observation, though not all such information is accurate. Much of our
behavior is learned by observing others, and sex is no exception.

Pavlovian Conditioning
While operant learning affects the frequency of behaviors that operate on people and
things, Pavlovian conditioning alters the reflexes and emotions that are involved with
sex and intimacy. Pavlovian conditioning allows inborn reflexes to be linked with new
stimuli in ways that we did not evolve to experience. Because salivation helps start
the digestion of food, dogs (and many other species) have evolved to salivate when
they smell food. By ringing a bell before presenting food, Ivan Pavlov conditioned
dogs to salivate to the sound of a bell, thereby expanding the range of stimuli that
elicit salivation.

The pleasurable feelings of the sexual and intimate activities can be associated with
all sorts of new stimuli via Pavlovian conditioning. We are biologically prepared such
that touch to the genitals triggers the sex reflex, but many other stimuli can become
associated with sex via Pavlovian conditioning. A young woman who pays attention
to her boyfriend's hairy chest while having sex with him may come to find hairy
chests to be a sexual “turn-on” or “erotic stimulus.” Later she may notice that she
becomes sexually aroused when she sees other men who expose their hairy chests.

Whatever people are thinking about when genital touch elicits the sex reflexes can
become a sexual turn-on. A young man who masturbates while watching pornog-
raphy is pairing new stimuli with the sex reflex, and he may end up feeling sexually
excited by seeing or thinking about such activities in the future. Even if he does
not masturbate while viewing porn, he may think about the pornographic images
when he masturbates at a later time, thereby sexualizing those stimuli. If he asks
his girlfriend to do some of the things shown by the porn videos, his girlfriend may
be happy to oblige; but some girlfriends are surprised that their boyfriends want
to try some of the aggressive activities depicted in porn scenes, and this can create
problems for the couple.

Masculinization of the Brain

During a critical period of fetal development, boys' testicles release a short burst
of prenatal testosterone, which alters their brains in ways most females do not
experience. As a result, boys and men tend to prefer somewhat higher levels of
rough-and-tumble play, rowdy activities and vigorous physical movements, com-
pared with girls and women. There is a great deal of variation in optimal activity levels
in both males and females, but males tend to prefer higher levels of stimulation
and activity than females do. These differences often persist well into adulthood.
This does not mean that boys cannot learn to control themselves and play gently
with others, but many males continue to prefer higher levels of activity and sensory
stimulation than do females well into adulthood. This contributes to a commonly
observed gender difference in the activities that males and females gravitate toward.
For example, it is easier for males than females to learn to love rugby and downhill
mountain-bike racing, and they may also prefer more active sexual activities than do
females.

There are genes that can affect the sexual orientation of females and males, leading
some individuals to become lesbians, gay males or other variations on “gender
queer.” Male homosexuality is about 50% genetically heritable. The sexual orienta-
tion of lesbians appears to be about 25% heritable. Personal learning, culture and
many other variables interact with heritable influences to produce many different
sexual and gender preferences.

In summary, both biology and learning are important in the development of sexual
behavior and both intertwine, starting in infancy. The following sections show how
nature and nurture continue to interact after the onset of puberty.

> Read full chapter

Addressing Sexual Activity of Elders

Occupational Therapy with Elders (Fourth Edition), 2019

Women
1. Decrease in rate and amount of vaginal lubrication may possibly lead to painful
intercourse.8,12,71
2. Orgasmic phase decrease may occur in elder women, resulting in a decrease
in orgasm intensity.8,11,12,13,71
3. Structural changes or atrophy may occur in the labia or uterus, in addition to
a reduction in the expansion of the vagina width and length.13,71,72
4. Thinning of the lining of the vagina can result in irritation and painful inter-
course.8,13,71
5. Sexual stimulation from the nipples, clitoris, and vulva may decrease with age
due to a decrease in sensation.72

> Read full chapter

Female Sexual Function

Lesley Marson, Ursula Wesselmann, in Principles of Gender-Specific Medicine
(Third Edition), 2017

4.3.2 Genital Arousal

In women genital arousal comprises an increase in vaginal lubrication and blood
flow to the vagina and clitoris, which facilitates clitoral erection; these changes can be
clearly mimicked in animal models. Understanding the peripheral and spinal mech-
anisms mediating genital sensations and arousal is not only important for helping
understand decreased genital arousal disorders, but is essential for development
of treatments to aid women with persistent genital arousal that can cause severe
distress in women.2,136,137

Vaginal dryness can occur concomitant with chronic disease; e.g., with diabetes
mellitus, hypertension, and after spinal cord injury. Vaginal dryness is common
in postmenopausal women, and can be reduced by the use of topical or systemic
estrogens in women, who have no medical contraindications to gonadal hormonal
treatment—i.e., women with estrogen receptor-positive breast cancer would need
to be treated with nonhormonal agents. A clinical trial reported that topical lidocaine
applied to the vulvar vestibule before vaginal penetration improved sexual function
and decreased sexual distress, suggesting an alternative treatment to hormones.138
Vaginal dryness often leads to decreased sexual desire, impaired arousal, personal
distress, and vaginal pain with intercourse (dyspareunia). Female sexual dysfunction
in diabetic individuals, e.g., manifests in inadequate vaginal lubrication which is
independent of depression.139,140

During sexual arousal fluid is secreted through the vaginal wall. Several mechanisms
have been proposed to regulate vaginal vasocongestion, including increased vaginal
blood flow. The vaginal epithelium transports Na+ ions from the lumen to the
serosa. Neurotransmitters such as nitric oxide, vasoactive intestinal peptide (VIP),
neuropeptide Y, and calcitonin gene-related peptide have been shown to be involved
in modulating vaginal fluid secretion.8,88,141–143 Vaginal lubrication is also regulated
by aquaporins (AQPs), which regulate water transport in and out of smooth muscle
cells.144 Studies examining AQP expression in both rat and human vaginal tissue,
and the changes that occur with menopause and with diabetes have provided a
working hypothesis that decreased vaginal fluid secretion occurs concomitant with
an increase in AQP expression.144–148

Animal models examining changes in vaginal blood flow have been used to de-
termine the neural pathways and the influence of putative neurotransmitters in
modulating the blood flow to the vagina during genital arousal.149–151 Stimulation of
the pudendal sensory nerve (to mimic genital stimulation/arousal) increases vaginal
blood flow primarily via parasympathetic (pelvic nerve) pathways.151 Activation of the
pelvic nerve or exogenous infusions of VIP, can also be used to induce increases
in vaginal and clitoral blood flow.149 Inhibition of neutral endopeptidase (which
degrade VIP) potentiates the vaginal and clitoral blood flow induced by pelvic nerve
stimulation in anesthetized rabbits, suggesting that neutral endopeptidase may be
used to restore sexual arousal in women who have female sexual arousal disorder.

Sympathetic pathways also play a major role in regulating female genital arousal
by modulating the tone of vascular and nonvascular smooth muscle (via 1/2--
adrenergic receptors). In women, administration of -adrenergic agonists facilitate
subjective and physiological sexual arousal152–154 and activation of -adrenergic re-
ceptors, via adrenaline, or stimulation of the pelvic nerve induces vaginal lubrication
in vivo by stimulating vaginal epithelial Cl− secretion.155 These studies suggest that
both - and -adrenergic mechanisms mediate physiological sexual arousal. Women
engaging in physical activity have decreased prevalence for sexual dysfunction,
further suggesting that activation of sympathetic pathways enhance libido.

Changes in specific oscillations of blood flow have been recorded in both rat and
human vagina.156,157 In women, the oscillatory patterns correlated with subjective
arousal and not genital arousal. In rats, slow oscillatory vaginal blood flow patterns
were modulated by both autonomic and central mechanisms. Furthermore the
vaginal blood flow dynamics were regulated independent of other areas of the
body. These studies provide a translational opportunity for studying female sexual
dysfunction.

Isolated tissue bath studies in which myogenic and neural factors can be modulated
are useful for understanding the neural and pharmacological mechanisms medi-
ating tissue contraction and relaxation. For example, hypothalamic neuropeptides
-melanocyte-stimulating hormone ( -MSH), VIP, oxytocin, and neuropeptide Y
were all found to have differential roles in contracting or relaxing vaginal smooth
muscle and arterioles and provide further support for peptides, such as MCs, in
regulating peripheral genital arousal.158

> Read full chapter

Sexual Behavior
John D. Baldwin, Janice I. Baldwin, in Encyclopedia of the Human Brain, 2002

II.A Before Conditioning

The basic sex reflex occurs when touch to the genitals triggers penile erection and
vaginal lubrication. With prolonged tactile stimulation of appropriate intensities,
the reflex mechanisms eventually produce all of the genital changes that lead up
to orgasm and resolution (which is the end of the sex reflex).

Starting from birth, the sexual reflexes in the lower spinal cord send signals to and
receive signals from the brain. The brain can (1) sense inputs from the genitals and
sexual reflexes and (2) cause sexual responses in the genitals. The connections be-
tween the brain and the spinal reflex centers allow sexual responses to influence and
be influenced by cortical and subcortical processes, including learning, memory,
emotions, thoughts, and fantasies. Nerve fibers from the autonomic nervous system
connect the internal and external sexual organs to the central nervous system.

Reflexes are neural mechanisms that allow stimulus–response (S–R) activities to

occur without any prior learning. Touch-sensitive nerves in the genitals send afferent
signals to the reflex centers located in the lower spinal cord. Tactile stimulation of the
genitals serves as the unconditioned stimulus (US) that activates the reflexes; the word
“unconditioned” indicates that no prior conditioning is needed for these stimuli to
activate the reflex. All of the sexual responses are called unconditional responses (URs)
to indicate that no prior learning is needed for them to occur.

From birth, the sexual reflexes can be activated by gentle touch to the genitals,
without inputs from the brain. By adolescence, males tend to prefer stronger tactile
stimulation of the genitals than do females, but both males and females have
thresholds beyond which additional stimulation elicits pain. The most sensitive parts
of the female body are the glans and shaft of the clitoris (even though the shaft is
hidden beneath the skin), the minor lips (surrounding the outside of the vaginal
opening), and the frenulum of the clitoris (located where the minor lips connect
with the glans of the clitoris). The most sensitive parts of the male body are the glans
and shaft of the penis, along with the frenulum (the small strip of skin that extends
along the lower side of the penis, where the glans and shaft join). Touch of these
structures, with the appropriate pressure and patterns of movement, activates the
spinal reflex centers, and continued stimulation produces a long chain of several
distinct responses––called excitement, plateau, orgasm, and resolution––that often
take from 2 to 20 min for a person to experience all four of these phases. Hence,
the sex reflexes are quite different from such quick S–R reflexes as the patellar knee
jerk or jerking away from electric shock. The following paragraphs present a simple
summary of our current knowledge about the ways in which the sexual reflexes
function.

Appropriate tactile stimulation to the genitals first activates parts of the sacral reflex
mechanisms (in the lower vertebrae of the spinal cord). Early sacral activities relax
the muscles in the arterial walls, allowing the arteries to dilate and carry blood into
the genitals faster than usual. The excess blood in the arteries compresses the veins,
slowing the flow of the blood from the genitals. As a result, blood accumulates in
the pelvic area, creating vasocongestion of the genitals. This vascular engorgement of
the genitals produces the earliest phase of the sexual response, called the excitement
phase. Vasocongestion causes the male's penis to become larger and erect. It also
causes the female's clitoris to enlarge by a small amount and fluids to exude through
the vaginal wall, lubricating the inside of the vagina (making it more receptive to
penetration).

After the sacral reflexes are activated in the excitement phase, continued genital
stimulation transfers reflex control to the thoracic and lumbar sections of the spinal
cord (located above the sacral area). This initiates the plateau phase of the sexual
response, when the thoracic and lumbar reflexes cause even stronger vasoconges-
tion, resulting in additional vaginal lubrication for females and stronger erections
for males.

Continued sexual stimulation eventually causes the thoracic and lumbar reflexes
to initiate a several-second transition period that leads to orgasm. In males, the
thoracic and lumbar reflexes cause a 2- or 3-sec-long response called “ejaculatory
inevitability” during the first several seconds of the orgasm phase. At this time,
men can feel pleasurable genital sensations that come from rhythmic muscular
contractions in the walls of the male internal reproductive organs—the vas deferens,
seminal vesicles, and prostate gland—as these organs push their contents into the
tubes from which they will be ejaculated. Then reflex control is transferred to the
sacral reflexes, which initiate ejaculation and orgasm. Most men learn they cannot do
anything to stop ejaculation from occurring once the 2- to 3-sec period of ejaculatory
inevitability starts, hence its name. After this brief preejaculatory period, reflexive
control is transferred to the sacral reflexes, which now cause rhythmic muscle
contractions around the base of the penis (inside the body), starting at intervals of
0.8 sec and then gradually slowing to longer intervals. These muscle contractions
force the ejaculate out of the male's body at the time of orgasm, and they cause
orgasm to be experienced as rhythmic pulses or surges of pleasure.

In females, the transition from plateau to orgasm is similar to that seen in males,
but more subtle. During the plateau phase of the sexual response, the thoracic
and lumbar reflexes produce increasing vasocongestion, which augments vaginal
lubrication. Then, for about 2 or 3 sec before the sacral reflexes cause contractions
in the pelvic muscles at 0.8-sec intervals, women report the onset of pleasurable
experiences (at the time when males feel the pleasurable sensation of ejaculatory
inevitability). After this 2- to 3-sec period, females begin to experience the rhythmic
surges and pulses of orgasmic pleasure that occur as their genital muscles begin
to contract at intervals of 0.8-sec, gradually slowing to longer intervals. Thus, both
females and males have 2–3 sec of pleasure before reflexive control is transferred
to the sacral reflexes (which cause the muscle contractions that begin at 0.8-sec
intervals). Even though females do not ejaculate semen, the contractions of the pelvic
muscles are experienced as rhythmic pulses of pleasure. Many females and males
report that orgasm lasts for about 15 sec, though this timing shows considerable
variability.

After orgasm, the final phase of the sexual reflex is resolution, in which all of the
genital responses seen in the prior three phases subside. There is a reversal of
vasocongestion, and blood flows out of the genitals faster than it flows in. Most
muscle contractions slow and weaken, though small movements may continue for a
while. Gradually, the genitals return to the relaxed condition seen before the onset of
sexual stimulation. After orgasm, most males experience a refractory period during
which no additional stimulation can activate the sexual reflexes. In adult men, the
refractory period tends to last 30–90 min, but the period becomes longer with age.
At this time, the fluids that will be ejaculated at the time of the next orgasm flow into
the internal male reproductive organs. Females do not have an equivalent process
and can have multiple orgasms without several-minute pauses between them.

Although the sexual reflexes usually function rather automatically, as described

earlier, they can be partially or completely inhibited by pain, fear, depression, fatigue,
exhaustion, illness, biomedical problems, and certain drugs, even when effective
tactile stimulation is present. Before sexual therapy, a medical exam is usually
conducted to determine whether a sexual problem is the result of medical or drug
problems. If these causes are ruled out, psychological treatment is in order.

> Read full chapter

Spironolactone
In Meyler's Side Effects of Drugs (Sixteenth Edition), 2016

Breasts
Spironolactone causes breast tenderness and enlargement, mastodynia, infertility,
chloasma, altered vaginal lubrication, and reduced libido in women, probably be-
cause of estrogenic effects on target tissue. Menstrual irregularities were experi-
enced by almost all women taking spironolactone 400 mg/day and most developed
amenorrhea at doses of 100–200 mg/day. Normal menstruation was resumed within
2 months of withdrawal.

In patients with hepatic cirrhosis, gynecomastia was twice as common during

treatment with spironolactone than with potassium canrenoate (42% versus 20%)
at equiactive doses [59]. This difference may be related to structural characteristics
unique to metabolites of spironolactone, namely a thiol group at the 7-alpha posi-
tion [60].

In a man with chronic heart failure and gynecomastia attributed to spironolactone

there was bilaterally increased uptake of 67Ga citrate and 18F FDG in the breast
tissues [61].

Bilateral painful gynecomastia that occurred after treatment with spironolactone

100 mg tds for 2 months in a 54-year-old Caucasian man resolved within 1 month
after treatment was switched to eplerenone 25 mg tds [62]. Eplerenone is 370-fold
less potent than spironolactone in blocking dihydrotestosterone activation of an-
drogen receptors [63], which probably explains this. In the EPHESUS study [64] the
incidence of gynecomastia was 0.5% compared with 9% with spironolactone in the
RALES study [1].

In 175 patients with resistant hypertension who took spironolactone 25–100 mg/day

adverse reactions that were attributed to spironolactone included gynecomastia or
breast discomfort in seven; as a result spironolactone was withdrawn in six (3.4%);
one man had reduced libido [25].

In a prospective study of the effect of spironolactone on atrial remodeling, 110 pa-

tients with preserved left ventricular function who had an acute myocardial infarction
were randomized into two groups: conventional therapy (n = 55) and additional
spironolactone 25 mg/day with standard conventional therapy (n = 55) [65]. Serum
creatinine and potassium concentrations were unchanged during the 6-month
study. Three patients stopped taking spironolactone because of gynecomastia.

Of 53 patients on hemodialysis who were randomized to spironolactone 50 mg or

placebo thrice weekly after dialysis, two had slight breast tenderness [66].

Of 168 patients with mild to moderate heart failure (NYHA class I–II) and left
ventricular ejection fraction 40% or less who were randomized to spironolactone
mean dose 52 mg/day or placebo, spironolactone was withdrawn in two patients
with gynecomastia and breast tenderness [30].

The effect of spironolactone, in combination with other medications, on recurrent

atrial fibrillation has been evaluated in 164 consecutive patients, mean age 66 years,
in a randomized 12-month trial; there was gynecomastia in three patients who took
spironolactone; the plasma potassium concentration did not change significantly
[67].

When spironolactone 25–50 mg/day was added to an existing regimen of three anti-

hypertensive agents for the treatment of uncontrolled hypertension in 344 patients,
mean age 62 years, of whom 43% had cardiovascular co-morbidity, eight men (5.2%)
developed gynecomastia and two (1.3%) had impotence leading to withdrawal of
spironolactone [31].
In a randomized controlled comparison of the antihypertensive effects of spirono-
lactone and eplerenone in 141 patients with primary hyperaldosteronism, 21% of
the men who took spironolactone developed gynecomastia compared with 4.5% of
those who took eplerenone [68]. In women there was mastodynia in 21% of those
who took spironolactone compared with none of those who took eplerenone. Those
who took eplerenone were more likely to report general and non-specific complaints
than those who took spironolactone. Withdrawal rates because of adverse events
were similar. A potential source of bias in these results came from the higher rates
of withdrawal due to treatment failure in those taking eplerenone (20%) compared
with spironolactone (4.2%).

In 19 patients with hepatic cirrhosis who had taken spironolactone and developed
painful gynecomastia, substitution of spironolactone with eplerenone for 3 months
resulted in reduced breast pain [69]. Although this suggests that eplerenone does not
cause painful gynecomastia, it does not rule out the possibility, in the absence of a
control group, of tolerance to the adverse reaction during continued administration
of an aldosterone receptor antagonist.

> Read full chapter

Sexual arousal and response — the psy-

chosomatic circle
John Bancroft MD FRCP FRCPE FRCPsych, in Human Sexuality and Its Problems
(Third Edition), 2009

Oestrogens and female sexuality

There is long-standing and consistent evidence of the importance of E for normal
vaginal lubrication, with post-menopausal decline in E commonly resulting in vagi-
nal dryness. However, whether E has a direct effect on sexual interest and arousability
has been less certain. In a recent review of the literature, Dennerstein et al (2003)
concluded that there is a dramatic decline in sexual functioning with the natural
menopausal transition, with changes in most aspects of sexual function correlating
with levels of oestradiol but not androgens.

A number of studies of the symptoms of either natural or surgical menopause,

reviewed above, have either taken women on E replacement, who continue to have
loss of sexual interest or response, and evaluated the effect of adding T, or have
compared E replacement with a combination of E and T, or T alone. Much less
attention has focused on the effects of E itself on sexual interest and arousal in
post-menopausal women.

Dennerstein et al (1980) studied 49 surgically menopausal women, all of whom

had stable, satisfying sexual relationships, using a double-blind crossover design
in which they spent 3 months on each of four treatments: ethinylestradiol (EE),
l-norgestrel, combination of EE and l-norgestrel, and placebo. The EE only regime
was significantly better than the others in improving sexual interest, enjoyment,
orgasmic frequency and mood. Also found were correlations between measures of
sexual desire and of mood, particularly ‘feelings of well-being’.

Nathorst-Böös et al (1993b) compared transdermal E (Estraderm 50 μg/24 h) with

placebo in 242 naturally menopausal women seeking treatment for menopausal
symptoms who had not previously received hormone replacement. After 12 weeks,
in comparison with the placebo group, the E group reported significantly more
sexual fantasies, better lubrication, less coital pain and more sexual enjoyment.

Only one study of hormone replacement has incorporated different doses of E. Sher-
win (1991) randomly assigned ‘perimenopausal’ women to four treatment regimes,
involving either a low (0.625 mg) or high (1.25 mg) dose of Premarin for 25 days
out of each month, and either Provera (5 mg) or placebo for days 15–25 of each
month. Each woman took the assigned regime for 12 months and was assessed
with daily ratings during the 3rd, 6th, 9th and 12th months. The main purpose of
the study was to evaluate the effects of the progestagen on mood and sexuality and
to assess whether the dose of E would modify any mood effect. Results showed that
the progestagen had a negative effect on mood, but not on sexuality. The mood
effect was attenuated by the higher E dose. However, it is the comparison of the
low and high E dose (combined with placebo), which is of most relevance here.
Apparent from the graphed sexual interest data (Figure 3, Sherwin 1991), women
in the high E+ placebo group had substantially lower levels of sexual interest during
the pre-treatment month than the low E+ placebo group, yet by the 6th month of
treatment, and continuing through the 12th month, were showing noticeably higher
levels of sexual interest. The results were analyzed using a four groups × five time
periods ANOVA, so the implicit assumption is that the comparison of the two E
groups was not significant. But this was not commented on in the paper.

The need to establish with more certainty the role of E in women's sexuality is rein-
forced by uncertainties about the ways the T effects, reviewed above, are mediated.
Wallen has strongly advocated that the sexual effects of T in women result from the
consequent increase in free E, supporting this conclusion with experimental data
from Rhesus monkeys (Wallen & Parsons 1998).

> Read full chapter

Neurology of Sexual and Bladder Disor-
ders
Alessandra Graziottin, ... Laura Bertolasi, in Handbook of Clinical Neurology, 2015

Vaginismus and dyspareunia

Vaginismus indicates the persistent or recurrent difficulties of the woman to allow
vaginal entry of a penis, a finger, and/or any object, despite the woman's expressed
wish to do so (Basson et al., 2004; Graziottin et al., 2004; Graziottin, 2006a; Graziottin
and Rovei, 2007).

There is often (phobic) avoidance and anticipation, fear, and experience of pain,
along with variable involuntary pelvic muscle contraction. The disorder may be
lifelong or acquired, generalized or contextual, biologic and/or psychogenic and may
(or may not) cause personal distress. In the vast majority of cases, however, coital pain
is a powerful trigger of personal and relational distress. When severe, vaginismus is
the leading female etiology of unconsummated marriages and relationships. When
mild, it may allow difficult, painful penetration, becoming the most frequent
etiology of lifelong dyspareunia, i.e., of coital pain present from the first sexual
intercourse. A lifelong hyperactive pelvic floor (“myogenic hyperactivity,” sometimes
associated with phobia of penetration, i.e., “true” vaginismus) anatomically reduces
the entrance of the vagina and predisposes the introital vestibular mucosa to mi-
croabrasions mechanically provoked by any attempt at intercourse.

The contributing factor is inadequate genital arousal, due to the reflex inhibition pain
has on vaginal lubrication and vulvar congestion and/or fear of pain. The mechanical
mucosal damage immediately activates the MC response (Graziottin and Murina,
2011; Theoharides et al., 2001) when attempts at intercourse are recurrent, and/or
coital damage is persistent, and/or concomitant factors such as Candida vaginitis
further contribute to the vestibular inflammatory state, and localized vestibular pain,
contributing to vulvodynia. Three key consequences are involved:

1. the MC is hyperactivated, with hyperproduction of inflammatory molecules

and neurotrophins such as nerve growth factor, which induces
2. the proliferation of pain nerve fibers, responsible for introital hyperalgesia, and
allodynia, and induces or worsens
3. hyperactivity of the pelvic floor.

In summary, vulvodynia can trigger dyspareunia, and painful intercourse (due to

vaginismus or dyspareunia from any etiology) may worsen or precipitate vulvar pain,
 and maintain it. The only exception is vulvodynia in children or virgin adolescents,
or in women of any age who do not have penetrative sex for a number of reasons.

> Read full chapter

Neurological Rehabilitation
Mindy Lipson Aisen, in Handbook of Clinical Neurology, 2013

Females
Women with MS report decreased libido, orgasmic difficulty, severe external
dysesthesias, and lack of vaginal lubrication. The majority (72%) report a change in
sexual function, and 39% an unsatisfactory sex life (Lundberg, 1981). Approximately
30% experience decreased genital sensation and an equivalent number report im-
paired lubrication (Dachille et al., 2008). As with men, women with MS experience
sexual impairment directly related to the location and severity of lesions. Symptoms
often develop abruptly, and are often associated with bowel and bladder dysfunction.
Women with lesions involving the sacral spinal segments may maintain the capacity
for psychogenic lubrication while losing the capacity for reflex lubrication (Sipski,
1995b). Alternative stimulation techniques and vaginal lubricants are recommended.

As with SCI, hypersensitivity or dysesthetic pain does not respond well to medica-
tions, though anecdotal reports support the use of tricyclics or venlafaxine. Sildenafil,
in doses of 50−100 mg, has been reported to improve sexual satisfaction in women
with MS (Dachille et al., 2008).

> Read full chapter

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