Sex

Lecture 23; December 3, 2013
n e u r1
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Sex
SEX!!!
Now that I have your attention, lets learn some neuroscience
and endocrinology.
Sexual behavior is interesting, because while it is highly
essential to the survival of the species, it doesnt do a great
deal to the individuals lifespan.
Nevertheless, motivation for sex is tremendously high.
All vertebrate, terrestrial animals go about sex roughly the

same way.
Humans may be the most sex-obsessed of all animals, but not by
as much as youd guess.
NEUR%1201%%Fall%2013%%Harry%MacKay%
Sex determiniation
Females and males share almost all of their genes. The only real
dierence between them, genetically speaking, is that females have two
X chromosomes, and males have an X and a Y chromosome.
Sex dierentiation

The testes and ovaries, respectively the male and female gonads, arise
from the same tissue embryonically.
Testes develop under the influence of the Y chromosome. If no Y
chromosome is present, ovaries develop by default.
Illustration from J.P.J. Pinel, Biopsychology. Allyn & Bacon, 2011
Sexual neurodevelopment
While sex is ultimately a matter of genetics, the

sex chromosomes merely get the ball rolling
on sexual dierentiation.
The vast majority of sexual development takes

place under the influence of hormones, and there
are many non-genetic reasons that these can vary.
The brain, like many other parts of the body,

contains sexual dimorphisms that develop
under the influence of sex hormones. This
process begins in-utero, and continues through
puberty.
Sexual dimorphisms are structural or functional
features that differ substantially between sexes.
Testosterone, produced mainly by the testes, is

the primary androgen. Estradiol, produced
mainly by the ovaries, is the primary estrogen.
However, males produce small amounts of

estrogens, and females produce small amounts of
androgens.
Note for hormone enthusiasts: estrogen actually
refers to a family of three chemically similar
hormones: estrone, estriol, and estradiol (the most
potent estrogen). Androgen refers to two
hormones: testosterone, and dihydrotestosterone.
Testosterone and estradiol are both steroid

hormones, and they are chemically quite
similar to each other. T can be converted
into E by the enzyme aromatase, but this is a
one-way change.
Sexual neurodevelopment
The sex hormones can act in an organizational and an

activational capacity.
When hormones permanently influence development, they are

said to be acting in an organizational capacity.
When hormones affect short-term behavior, they are acting in
the activational capacity.
Feminized behavior: the lordosis

response.
Sexual development of the brain consists of three

separate, dissociable processes:
Masculinization: allows the brain to produce male sexual

behavior.
Feminization: allows the brain to produce female sexual
behavior.
Defeminization: the process through which the brain loses the
ability to produce female sexual behavior.
Testosterone is produced only by males. It is converted to

estradiol in the brain, where it has the organizational eect
of masculinizing and defeminizing the brain.
Feminization is considered the default state in the rodent

brain; if not exposed to the masculinizing and
defeminizing influence of testosterone (specifically its
metabolite estradiol), the brain takes on the feminine form.
Defemenized behavior: absence of a

lordosis response.
Masculinized behavior: mounting.
Sexual dimorphisms in the brain

There are many sexually dimorphic areas in
the brain. These areas show structural,
neurochemical, and molecular dierences
between males and females.
The frontal cortex, hypothalamus, and
amygdala have a high degree of sexual
dimorphism.
The medial preoptic area (mPOA) is a

nucleus in the hypothalamus. A subregion
called the sexually dimorphic nucleus
(SDN) It is 5x larger in male rats than in
female rats. This is often abbreviated as
SDN-POA.
The difference in size is regulated by the
organizational effects of testosterone (after it is
converted to estradiol).
The sexually dimorphic nucleus of the

media preoptic area in male and
female rats. Note the size difference.
Sexual dimorphisms in the brain
Sexually dimorphic areas of the brain develop under the organizational

influence of the sex hormones.
Injection of testosterone at 4 days of age masculinizes the female SDN-POA.
Castration at birth feminizes the male SDN-POA.
The stages of sexual behavior

Reproductive behavior can be divided into
several stages:
1. Sexual attraction
Brings males and females together.

Synchronized with reproductive readiness.
2. Appetitive behavior
Behaviors aimed at establishing or

maintaining sexual interaction. In rats this
consists of hopping, darting, ear wiggling,
singing, sniffing, etc.,
3. Consummatory behavior (copulation)
Sexual intercourse itself.
4. Postcopulatory behavior
Includes everything that follows sex.

Refractory period, copulatory lock,
parenting, pair-bonding, etc.,
Control of male sexual behavior

Destruction of the mPOA destroys male
consummatory sexual behavior in every
species examined.
Electrical stimulation of this nucleus elicits
copulatory behavior in male rats.
Interestingly, destruction of the mPOA does not
appear to aect appetitive sexual behavior.
Males with lesions in the mPOA will still work to

gain access to a female, but once they do, they fail
to show the appropriate consummatory behavior.
Neurons in the mPOA project to a region in

midbrain called the lateral tegmental field.
This tract is essential for the function of this circuit

disrupting it is equivalent to lesioning the mPOA.
The amygdala appears to be responsible for

appetitive sexual behavior.
In this study, male rats with lesions in the

mPOA can press a bar to gain access to a
female. They will, but fail to successfully
mate with them after.
Control of female sexual behavior

The ventromedial nucleus of the
hypothalamus (VMH) contains circuits
that are critical for female sexual behavior.
Destruction of the VMH in female rats
destroys female sexual behaviors
(lordosis, etc.,).
Females with this lesion also tend to attack
male suitors.
The influence of the VMH on sexual

behavior is dependent on projections to
the periaqueductal gray (PAG) in the
midbrain.
Diagram showing both male and female

sexual circuits in the rat brain.
Destruction of this tract mimics the effects

of VMH lesion.
Activational eects of sex hormones in

males
In both males and females, sexual behavior
is stimulated by the activational eects of
the sex hormones.
Testosterone is needed for male sexual
behavior in both rats and humans.
Castrated males lose interest in sex, unless
they are given testosterone injections.
Another word for castration is orchidectomy
(see fun fact to right).
Testosterone plays a permissive role in this

equation.
Giving massive doses of testosterone does
not increase sexual behavior proportionally.
Blood levels of T are not correlated with sex
drive in healthy males.

Orchids are a type of flower. The name orchid comes

from a Greek word meaning testicle. The plant was
given this name because its roots look like testicles.
Interestingly, orchids are also used as aphrodisiacs an
example of sympathetic magic.
Activational eects of sex hormones in

females
Sexual behavior in female rats is stimulated by a
combination of estrogens and progesterone.
The levels of estrogens and progesterone peak

immediately prior to ovulation in female rats. This ensures
that they are primed for sex at the optimal time for
fertilization.
Ovariectomized female rats will lose interest in sex, unless
given a combination of estradiol and progesterone.
Estrogens and progesterone are not as tightly linked

to sexual behavior in human females.
Human sexual receptivity also peaks prior to ovulation, but

unlike rats, human females can and will have sex at any
point in their cycle.
Ovariectomy does not have a big effect on sexual behavior
in humans. Vaginal lubrication decreases, but this only
affects the mechanics of the process, not the desire itself.
Testosterone is important in regulating sexual behavior

in human females.
Injections of testosterone are effective in inducing sexual
motivation in human females.
The human menstrual cycle. The follicular

phase is characterized by high estrogen
and low progesterone. The luteal phase is
characterized by high progesterone and
low estrogen.
Contrasting humans and rats

Sexual attraction is a largely hormonal and
pheromonal process in rats.
Rats will show sexual responses to odors from a
sexually receptive member of the opposite sex. They
will even cross an electrified grid to get closer.
Human sexual attraction is a much more complex

aair, with influences at the pheromonal, hormonal,
and physical level, as well as higher level cognitive
influences.
People devote substantial amounts of cognitive time
and energy to sex. They think of it, dream about it,
and make plans about it.
All of this suggests that the cortex is important for sex in
humans.
Dissecting desire
Conventional wisdom holds that male sexual arousal
is a simplistic thing, capable of being evoked by
pictures and tactile sensations.
Female arousal is taken to be a more complex aair,
with more emphasis on emotional and contextual
cues.
According to a recent study, this may not be entirely
true. Women are perfectly capable of arousal through
visual stimuli, and may in fact be less discriminating
than men in that regard.
Chivers made male and female participants watch
videos consisting of things like: nude exercise, solitary
masturbation, sexual intercourse, and bonobos having
sex.
She found that heterosexual women were aroused by
the human sex videos irrespective of who was in the
video.
Males, on the other hand, were only aroused by their
preferred category of sexual partner.
Thankfully, nobody was particularly aroused by the
bonobo sex.
Neural correlates of male orgasm

In this study, males were placed in PET
brain scanners and stimulated to
ejaculation by their female partners.
While this was happening, researchers
noted activation of the VTA, as well as
activation of other, somewhat
unexpected areas like the occipital lobe
and cerebellum.
Researchers did not detect any changes
in the hypothalamus, and actually saw a
decrease in activity in the amygdala and
surrounding entorhinal cortex.
Neural correlates of female orgasm

To study orgasms in females, researchers went to
greater lengths to ensure that they were real (male
orgasms provide their own physical corroboration).
In this study, the researchers used measures of rectal

pressure, which is known to increase during orgasm.
Women were placed in a PET scanner and clitorally

stimulated by their partners.
They were asked to imitate an orgasm while being
stimulated. This was used as the baseline for
subsequent brain scanning.
Researchers found increased activity in clitoral

sensory areas, as well as in the cerebellum. More
notable, however, was the massive decrease in
activity seen throughout the brain (left temporal
lobe, amygdala, prefrontal cortex, orbitofrontal
cortex, etc.,)
This suggests that fear, moral reasoning, and
conscious control are all temporarily suspended
during orgasm.
However its hard to tell if this is a cause or
consequence of orgasm.
Sexual orientation
Humans can be classified according to three basic dimensions:
Biological sex: determined by the action of genetics and prenatal hormones.
Sexual identity: an individuals feelings of being a man or woman (or both, or
neither).
Sexual orientation: physical, emotional, and sexual partner preference.
None of these three dimensions has clear-cut boundaries. Each are

continua along which people can fall nearly anywhere.
Homosexuality
Homosexuality is simply one extreme on the continuum of human
sexual orientation. Over the years, there have been many theories
for why it exists.
Note that a societys tendency to theorize about the origin of
homosexuality is to an extent a reflection of an implicit perception that it
is abnormal. We do not spend much time asking where
heterosexuals come from, but thats an equally reasonable question.
Psychoanalytic theories suggested that homosexuality was

caused by weak, detached fathers and overprotective mothers.
Behavioral theories suggested that homosexuality resulted from
a conditioned preference to same-sex sexual stimuli.
Neither psychoanalytic nor behavioral theories of
homosexuality have much empirical support.
Homosexual individuals routinely assert that they had felt attraction to
the same sex early in childhood, and felt different the entire time.
Biological theories of homosexuality

Homosexuality appears to have a large genetic
component. Twin studies suggest a concordance rate
of around 50% in monozygotic twins.
In spite of this, no specific genes have been conclusively
linked to homosexuality. Studies that claim to have found
genetic links are rarely replicated.
It is often assumed that homosexual males have

dierent levels of sex hormones. This is not true.
Orchidectomy reduces sexual behavior in both gay and
straight males it does not redirect it.
Homosexual behavior is seen in most animal species.

It is especially common with animals in captivity.
It has been suggested that homosexuality develops due
to overcrowding, or perhaps simply due to an absence of
alternatives (the so-called prison theory).

It has been suggested that diering exposure
to prenatal testosterone may explain the
development of homosexuality.
This does seem to be the case in rats and
other animals tested.
Castration of newborn male rats leads to a
higher rate of feminized sexual behavior in
adulthood.
Injecting newborn female rats with testosterone
leads to masculinized and defeminized sexual
behavior in adulthood.
It is much more dicult to confirm this in

humans. Lesbian women, but not gay men,
show evidence of masculinized digit ratios.
This suggests that they were exposed to
slightly higher levels of testosterone during
development.
You can measure your own digit ratio.

Measure the index and ring fingers from
the basal crease to the tip, then divide
the index finger by the ring finger (2D/4D).

A recent line of research that shows promise focuses on the fraternal
birth order eect.
The odds of being homosexual increase with the number of older
brothers one has. Some estimates suggest that the odds of being gay
increase by 33.3% for each older brother one has.
Studies of blended families show that this eect is due to the number
of boys born to one mother, not the number of boys one is reared
with.
Blended families occur when biological siblings are raised together with
adopted or step-siblings.
The maternal immune hypothesis states that some mothers

become progressively more immune to masculinizing hormones.
The more male fetuses a mother carries, the more her immune system is
exposed to androgens. Her immune system may attack these
androgens and prevent them from having the usual effects on
development.
Brain dierences in homosexual men

Given the role of the SDN-POA in promoting male-specific sexual behavior, and the
known malleability of this nucleus, perhaps its also responsible for human
homosexuality?
In 1991, Simon LeVay discovered that the 3rd Interstitial Nucleus of the Anterior
Hypothalamus (INAH3), a region that is usually larger in males than females, was
smaller in brains from homosexual men.
This area may be the human equivalent of the SDN-POA.
This observation has been semi-replicated a number of times (but it is not universal).
INAH3 in heterosexual man.!
INAH3 in homosexual man.!
So what?
Sex is a complex biological phenomenon depending on the
coordinated action of genes, hormones, and brain regions.
Human sexuality is enormously complex, and simplistic
assumptions like men are men, women are women simply
wont do.
And we havent even gotten to talk about interesting cases

like congential adrenal hyperplasia, androgen insensitivity
syndrome, ablatio penis, trans-sexuality and more!

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Lecture 23; December 3, 2013

All vertebrate, terrestrial animals go about sex roughly the

Illustration from J.P.J. Pinel, Biopsychology. Allyn & Bacon, 2011

While sex is ultimately a matter of genetics, the

The vast majority of sexual development takes

The brain, like many other parts of the body,

Testosterone, produced mainly by the testes, is

However, males produce small amounts of

Testosterone and estradiol are both steroid

The sex hormones can act in an organizational and an

When hormones permanently influence development, they are

Feminized behavior: the lordosis

Sexual development of the brain consists of three

Masculinization: allows the brain to produce male sexual

Testosterone is produced only by males. It is converted to

Feminization is considered the default state in the rodent

Defemenized behavior: absence of a

Masculinized behavior: mounting.

Sexual dimorphisms in the brain

The medial preoptic area (mPOA) is a

The sexually dimorphic nucleus of the

Sexual dimorphisms in the brain

Sexually dimorphic areas of the brain develop under the organizational

Illustration from J.P.J. Pinel, Biopsychology. Allyn & Bacon, 2011

The stages of sexual behavior

Brings males and females together.

Behaviors aimed at establishing or

3. Consummatory behavior (copulation)

Sexual intercourse itself.

Includes everything that follows sex.

Control of male sexual behavior

Males with lesions in the mPOA will still work to

Neurons in the mPOA project to a region in

This tract is essential for the function of this circuit

The amygdala appears to be responsible for

In this study, male rats with lesions in the

Control of female sexual behavior

The influence of the VMH on sexual

Diagram showing both male and female

Destruction of this tract mimics the effects

Illustration from J.P.J. Pinel, Biopsychology. Allyn & Bacon, 2011

Activational eects of sex hormones in

Testosterone plays a permissive role in this

Orchids are a type of flower. The name orchid comes

Activational eects of sex hormones in

The levels of estrogens and progesterone peak

Estrogens and progesterone are not as tightly linked

Human sexual receptivity also peaks prior to ovulation, but

Testosterone is important in regulating sexual behavior

The human menstrual cycle. The follicular

Contrasting humans and rats

Human sexual attraction is a much more complex

Neural correlates of male orgasm

Neural correlates of female orgasm

In this study, the researchers used measures of rectal

Women were placed in a PET scanner and clitorally

Researchers found increased activity in clitoral

None of these three dimensions has clear-cut boundaries. Each are

Psychoanalytic theories suggested that homosexuality was

Biological theories of homosexuality

It is often assumed that homosexual males have

Homosexual behavior is seen in most animal species.

Biological theories of homosexuality