[MUSIC PLAYING]

Stanford University.

Is this on?

Yes.

Well, congratulations.

Everybody has survived

the midterm including

the TAs so far, who

recently have been let out

of their entombment
with thousands

of pages of exam pages.

So those are rolling along,

and everybody is still awake

so that's good.

OK.

So we have now
officially entered

the second half of the course.

And organizational
things-- readings

will now be read hopefully.

OK.

So that's not useful.

Books.

Books.

Now is a good time to

actually go and start

reading those books.

And again, if you are not

up to all of both of them,

some recommend the

chapters have come along.
Pay attention to those.

And again, a subset of you-- may

not know it yet, but about 25%

of you will have your life

transformed by that chaos book,

whereas 25% will resent

the purchase price

and my forcing you to do this.

Also, another major,

major transition here,

which is, as far as I can

tell so far, I have run out

of steam turning
the extended notes

into actual expository

writing, so they're just

going to take the form of really

poorly organized outlines.

So that happens.

OK.

What else?

What we now
transition to is going

to be our strategy for the

rest of our course, which

is to look at various subjects.

And coming down the pike after

the lectures today and Friday

on sexual behavior will be

aggression, competition,

cooperation, empathy,
potentially language

use, schizophrenia.

Somewhere in there,
there is going

to be a week or so
on that chaos stuff.
But for all of
these subjects, we

are now going to follow

the general strategy.

We will start off looking

at what the behavior is.

And what we're going

to try to do there,

in addition to looking at in
lots of different species,

is to be as objective as
any good old ethologist

in considering the
fixed action patterns

of the particular behavior.

That established, with the

promise that, in lots of cases,

what we think comprise

some of these behaviors

turn out not to be the case.

That established,
what we will start

is our inexorable march to

the left, the timeline that

constitutes everything
we learn now,

stepping back and saying, OK.

One millisecond before

that behavior occurred,

what was going on in the brain?

What parts of the brain?

What neurotransmitters?

All of that.

Stepping back before that, a

second before, a minute before,

whatever, what was

it in the environment
that triggered the brain
to produce that behavior?

What was the acute

releasing stimulus?

Stepping back, what do

hormone levels that hour,

that day, some such

time span of that,

what acute hormonal exposure

had to do with sensitizing you

to the environmental
stimulus which

released the nervous system

into generating the behavior.

Marching all the way back, throw

in culture some place or other.

Perinatal effects, early

developmental hormonal stuff,

eventually considering what do

the genetics of the individual,

of the population, of
the species-- evolution

kicking in there
some place or other--

what do all of these things

and something having to do

with ecology thrown in

there just for good measure.

Working our way back

in each of these cases,

understanding what was

the biology of one second

before, one minute, one hour,

one million years before that

gave rise to it.

Back to our two major

themes from day one,

we are now about to be

unbound, unfettered,
by our buckets, our
categorical buckets,

and instead explore

their interactions.

The other being that notion

from the very first class, which

is that any given

point, if we're

talking about chronic

hormonal effects

on this behavior, what

we're talking about

is the way those hormone

patterns were shaped

during this period,

the way genes

contributed to the
enzymes that make

the hormones, the receptors.

The second you're talking

about genetics, all of this

is becoming apparent.

At every one of these

points, whatever

point we are talking

about is going

to be the end product of

everything to the left

and just as temporary sort

of footing before going

to the things on the right.

So this will be our

strategy forever after now.

So we start off doing

this with looking

at sexual behavior,
the neurobiology,

the endocrinology, the early

experiences, et cetera, et
cetera.

So to inaugurate the second

half of the course and the fact

that it's starting off with

lectures on sexual behavior,

it has to start off with the

stupid obligatory sex joke.

OK.

So the Martians come to

Earth, and they turn out

to be great guys.

They are really terrific.

They get along wonderfully

with Earthlings.

All of them like

each other a lot.

They're all becoming

great friends.

They pass their hours learning

about each other's planets.

What's the weather like there?

What are sports like here?

What are recipes?

Everybody's getting
along terrifically.

And eventually, the

Earthlings and the Martians

are getting along well

enough that they get around

to asking the question that

everybody is really interested

in, which is, well, how do you

folks go about reproducing?

So the decision was made.

The Martians are

going to go first.
So they clear out a big space,
and a whole bunch of Martians

come in there and they stand

on top of each other heads,

and their noses flash

different colors,

and steam comes out

of various orifices,

and there's clanking

noises and whatever.

And out pops a new

little Martian.

And the Earthlings say,

wow, that was great.

Love the concept.

And I got great video of that.

And that's terrific

and all of that.

OK

So that's worked out.

And now it's the humans turn.

So a willing couple is found,

and some space is cleared out.

And the Martians sit down

there with their video cameras

as well.

And clicking away.

And this couple goes at it.

And they finish the whole thing

in a sweaty mess at the end.

And the Martian say,

that's wonderful.

That's so interesting.

You Earthlings are just endless.

And the fascinating

things you do,
but we have one question
though, which is,

so where's the new human?

And they said, oh,

that takes nine months.

They say, well, why were they

in such a rush at the end?

So our first question

here is, why were they

in such a rush at the end?

OK.

Three possible answers.

Choice number
one-- vote for it--

why were they in such

a rush at the end?

Number one, because they

were acting with this fervent

desire to do something for

the good of the species.

Just seeing if any hands go up.

That's a good thing.

Option number two,

doing that because you

want to maximize the number

of copies of your genes

passed on to the
next generation.

Option number three,

because it feels good.

OK.

One hand goes up.

And I'm not sure what that

indicates about everyone,

but I will remind you from

the survey in the first class

there that a far greater

percentage of you

want to learn about depression

than about sexual behavior.

So there you have it with

the Stanford experience.

OK.

Because it feels good.

And what we deal with

here right off the start

is this important dichotomy

between proximal and distal

explanations for behavior.

Explanation, a
distal explanation,

for sexual behavior,

parentheses,

why were they in such

a rush at the end?

Passing on copies of
your genes, the effects

of hormones, and
certain reward pathways

in the brain, all of that.

Proximal mechanism being

that it feels good.

And starting off right

off the bat the thing

to make sense of
with sexual behavior

is it is driven by this
amazing little loop here

of sensory stimuli,
and feedback,

and immediate sensations that

drive the behavior coming out.

And all this stuff down here

is for the doctoral thesis

somewhere down the line.

That's not what
the motivation is.

Probably more than any other of

the behaviors we will look at,

the driving forces are

very proximal ones.

Nobody sits there and figures

out how many copies of genes

they are passing on and thus are

willing to speed up to produce

a new human nine months later.

It is instead, in
species after species,

it is proximal
motivating mechanisms

for generating the behaviors.

OK.

So beginning to look at
the actual behaviors,

there is a funny duality to

making sense of sexual behavior

across different species,

a funny sort of contrast.

The first one being that, well,

all species go about sex--

or all vertebrate
species go about sex

in a roughly similar way.

Yet, you don't want to be

too similar to the species

next door.

There's an interesting
sort of dichotomy there.

All sorts of vertebrate

species are doing things

with pelvic thrusting

and orgasms and-- hey,
stay tuned that's
coming-- and ejaculation,

and lordotic reflexes,

and things of that sort.

Highly conserved fixed action

patterns across lots and lots

of different species.

None the less, amid that,

you've got this other problem,

which is you want to have these

fixed action patterns being

specific to your species.

You do not want to mess up.

So there is this
strange simultaneity

of very, very conserved

basic building

blocks of the fixed action

patterns of sexual behavior.

But along with that, a lot of

selectivity within species.

Now how does that

selectivity begin to work?

What you get is this very

interesting interplay,

this intercalation, between the

releasing stimuli and the fixed

action pattern.

What you get is this

chaining of behavior.

In other words, the fixed

action pattern of one

of the individuals constitutes

the releasing stimulus

for the other individual's

fixed action pattern, which

constitutes the releasing

stimulus for this individual's
fixed action pattern.

This chaining of
transitions there,

of interplay between
these two, which

is where you get the

species specificity from.

OK.

So in terms of making
sense of that, of course,

any of this in terms of looking

at the general features of how

to think about sexual behavior

across species, of course, what

you have to have out the wazoo

is your basic ethology credo

of interviewing an animal
in its own language

about its sexual

behavior, wonderfully

summarized in this quote by

Martha McClintock, researcher.

I think I've used

this quote already,

which is, in her

particular case,

studying female rat sexual

behavior, which turns out

to be this very ornate

process involving

a lot of running around

on the part of the female.

Studying female rat

sexual behavior in a cage

is like trying to study swimming

behavior of a dolphin in a bath

tub.

You need to get it in

the natural setting,
or else you are going to
lose all sorts of insight.

In the particular realm of

female rat sexual behavior,

the standard picture

for decades and decades,

the centuries where our

finest minds have looked

at rats having sex,

the standard dogma

has been that the female role

is a very passive receptive one.

And it turns out it's a

very passive receptive

one if she doesn't

have enough room

to run around and do all sorts

of courting stuff on her own,

all sorts of proceptive sexual

behaviors, which she can't see,

if you're studying an animal

in a setting where they can't

speak in their own language.

So a big, big vote for

ethological logical principles

when it comes to this.

All right.

So just to get some jargon

under our belt right

from the start here, in terms

of how the professionals talk

about sex when they're

talking about sex

and trying to sound

like professionals,

here are some of the terms.

Old outdated term,

Freudian term,
that nonetheless has
entered the general world

of referring to sexual arousal

and motivation-- libido.

Libido, as we will see that

commonplace everyday usage

term, is perhaps more

technically described

as horniness.

But it can also be

described as one term

within a trio of the terms

that people in the business

really use most frequently--

attractivity, proceptivity,

and receptivity.

Quick, get to work on poems

about those three terms.

But what you've got

here is attractivity,

how attractive an individual

is to someone else.

Receptivity, how
receptive that individual

is to the interest of
the other individual.

Proceptivity, the
active behaviors

that are carried out in

response to being attracted to.

And thus, you could say

because of the attractiveness

of this organism,
this other organism

began proceptive behaviors

which did or did not

prove to meet with receptive

fixed action patterns
and responses.

The very words any

of us would use

to describe what
goes on at a party.

OK.

So we've got that triad

there, the terms that

are much more in common than

terms like libido or arousal

or motivation.

These are the more common ones.

What is another realm, in

terms that are much more used,

these are much

more zoology terms.

What's used far more

often in clinical medicine

is a description, a
dichotomy, between motivation

and performance.

And that is never used more

frequently than in the realm

of making sense of
sexual motivation

in men as dissociable
from erectile function

versus dysfunction,
motivation being very, very

different from performance.

So that's another
realm of distinctions.

Other realms as well--

desire, orgasms, arousal,

all sorts of commonplace terms.

The performance versus

motivation dichotomy
and the proceptivity,
receptivity, [INAUDIBLE]

are the major terms

that are used.

Next issue, in terms

of getting to this,

how do people find

out information

about sexual behavior?

One option is to sit there

with night viewing goggles.

And that's very useful

for nocturnal species.

But how do people find out

about human sexual behavior?

All sorts of ways

over the years,

starting with anonymous

questionnaires.

But a really clever technique

was worked out in the 1980s.

A biological mathematician
named Joel Cohen

getting at how to get

people to tell you

about very embarrassing things

about their sexual lives.

And this was prompted in the

80s when AIDS first swept in,

and it was wildly,

wildly taboo at the time

in virtually every
corner of this country

to admit to having a less

than standard, white bread

sexual orientation.

Take a look at the extended

notes to see the trick

that Joel Cohen came up

with, a very clever device

in order to figure out

what percentage of people

are doing what sexually

without asking anybody

to give an answer that

they would find, perhaps,

to be embarrassing or grounds
for all sorts of persecution.

OK.

So beginning to look at
aspects of behavior and other

features of the rightmost

end of all of this.

We start off with the

most central puzzle

in making sense of any of

this stuff, which is, so

what's up with female orgasms?

And we've got right off

the bat the simple problem

of making sense of this

biological phenomenon and one

where fertility is
not dependent on it.

One does not need to have

orgasms to become pregnant,

to give birth, to pass

on copies of one's genes.

So what's the deal with orgasms?

First off, a question

we will wind up

asking with a whole lot of the

behavior coming down the line

is, are we the only species?

And all sorts of

careful studies have

shown that we are

not the only species.

Other apes, other

primate species, as well

monkeys and apes, show

orgasm among the females.

And that is detectable by

all sorts of physiology

we'll hear about in a while.

We are not the only species.

One of the really

bizarre, pathetic things

about trying to do
research in this area

was one of the first papers

that ever demonstrated something

which physiologically was

identical to female orgasm

in rhesus monkey
females, which wound up

being a paper in
this journal Science.

Down there in the

footnotes, the authors

had to indicate this did not

make use of any federal grant

money to carry out the study.

Just to give you a sense of

where some of the stuff sits.

OK.

So what's up with female orgasm?

It is not necessary
for conception.

It is not necessary to
increase the number of copies

of your genes in
future generations.

Despite that, there

is some evidence
that it facilitates
fertilization.

And the technical term that's

always been given for that

is, bizarrely
enough, facilitation.

The notion is something about

the vaginal fluid, something

about the biochemistry of,

increases sperm motility.

Sperm swim faster and harder

and jump upstream back to spawn

or whatever it is
the sperm are doing

with more avidity, with

more energetic displays,

in an environment of
more vaginal secretion.

And orgasm greatly

increases that.

So the argument there

being that orgasm

facilitates fertilization
through the sperm facilitation

process.

Evidence for that has always

been a little bit indirect.

It is not airtight
that that happens.

Another argument for why

this increases fertility.

And this is sort of

an interesting one.

And the notion here is that

what an orgasm does is,

among other things,

exhausts you enormously,

causing you to be
far more likely to be
horizontal than vertical
shortly thereafter, and thus,

facilitating fertilization
because the sperm don't

have to swim straight

up against gravity.

I kid you not.

This is one of the

leading models out there.

Then there's the orgasm

facilitates female conception

out of reinforcement
theory, which is back

to the it feels good and thus

you are more likely to do it

again and increasing the

likelihood of passing on copies

of your genes.

All of this is wonderful.

All of these possible

mechanisms where,

even though female orgasm is

not necessary for conception,

it nonetheless increases
the likelihood of.

That's great.

However, what most of

the studies have shown,

though, is there
is no relationship

between the fertility of

a woman and her propensity

towards orgasm.

It does not seem to play

out in so far as any

of this facilitation
or horizontal
swim enhancement techniques
or whatever actually occurs--

these are not big enough

of effects to actually make

a difference in terms
of reproductive success.

So what else?

What else is known about it?

There is a certain
degree of heritability,

of propensity towards
orgasm in females.

And this is shown with

all our standard behavior

genetics techniques in
terms of comparing dizygotic

versus monozygotic twins.

We know what to do in terms

of not overvaluing findings

like those.

Nonetheless, they are there.

So if a basic puzzle is,

why do females have orgasms

if it's not necessary for

conception and, in fact,

the evidence is not great

that it even facilitates it,

why on top of all of

that, why such things

as clitoral orgasms, which

the studies generally show

are more easily

brought about than

are vaginal ones,

what's up with that?

Even more the same question.

Now somewhere in
there is a lurking
the heart breaking
possibility in making

sense of why female orgasm

exists the dreadful possibility

that what we're dealing

with here is a spandrel.

And that it is a spandrel.

It is simply baggage
brought along

that those guys have to go

through this orgasm physiology

to do any stuff with sperm

and pass on copies of genes

and all of that.

And it's simply baggage

that the same physiology

occurs in females.

That orgasm is simply

a spandrel in women.

And the counter scenario

that's always given

is, this is exactly

equivalent to the notion

that nipples are

spandrels in men,

in that women, female

mammals, need to go about all

this lactation business.

And that's part of the

whole package deal.

And just as it would be way

too much work to evolve females

without orgasms,
it would be way too

much to get rid of those

useless nipples on men.

OK.
Let's have a quick survey.

Yes?

So could a lot of
those questions

also be applied to why

do male have orgasms?

Because ejaculation can

occur without orgasm.

OK.

So why do males have orgasms?

Ejaculation can occur--

it is far more voluminous

in the face of an orgasm.

So that's easily framed in

terms of an adaptive mechanism.

OK.

Quick survey here.

How many people--

OK, how many guys

who have those useless

nipples, how many of you

go for the nipple as

spandrel in guys theory?

Whoa.

Is that slow in the hands?

OK.

Should I raise my hand?

Should I not raise my hand?

OK.

Women in the room, how many

go for the female orgasm

is merely spandrel theory?

If I recall, there was one other
question somewhere a few weeks

back that only got one

person fessing up to it,

and it came somewhere

around there also.

So I don't know if
that's the lighting

or if people tend to
sit in the same places.

OK.

So not a whole lot of enthusiasm

for the spandrel concepts here.

Nonetheless, that needs to

be seriously entertained.

OK.

So now looking at other features

of the fixed action patterns,

and amid all these

different species

doing lordotic stuff and

orgasms and ejaculation and all

of that, what are some of

the realms of sexual behavior

that are relatively

unique to humans?

First off, one that used to be

thought to be absolutely unique

was non-reproductive sex.

This is a world of difference

than those species where

the female ovulates for like

2 and 1/2 hours every year,

and everybody mates

at that point.

Or species where
somebody comes into heat,

a female is in estrus.
Humans have
non-reproductive sex.

And that was viewed

as absolutely unique.

What it is now clear is it

is not completely unique.

There are lots of

other species that do,

probably most famously bonobo

chimps and various cetaceans

like dolphins.

Nonetheless, it is certainly
a specialty among humans.

What else?

Foreplay, that used to be in the

category of human specialties.

And it is clear by now that

bonobo chimps, for example,

have vastly more

patience with foreplay

than average humans do.

We are not the only

species with that either.

Huge, huge controversy.

How unique is homosexuality

to human behavior?

Human sexual behavior?

And what's clear increasingly

is we're not the only species

with that either.

The original view, when people

would view homosexual behavior,

male-male, female-female,
in other species,

it would be animals
in captivity.

And it would be
the, why is there
so much homosexuality
in prisons argument--

lack of alternatives.

This was not normal,

natural behavior.

What is clear from

ethological field studies

is we are by no means
the only species

to have homosexual behavior.

What else?

One of the things that we do

seem to be fairly unique about

is having egalitarian
sex, which is

to say that there are

no human cultures where

as part of the central

tenets of that culture people

are restricted, only a

subset of individuals

are allowed to reproduce.

And this is a
world of difference

from various species.

For example, New World

monkeys, marmosets,

where it is only one

male and one female

in a group that does

the reproducing.

Instead, humans in every culture

ever seen have egalitarian sex.

What else?

What else is highly human?

People used to
think exclusively so
this endless quest for variety.

And again, just take a look

at these bonobo chimps,

and you'll see how small minded

we are when it comes to this.

But something else

that is indeed

unique to human sexual

behavior is the notion

that this is something

you do in private.

There is no other species where

the majority of sexual behavior

is conducted intentionally
outside the sight of everybody

else.

That is rather unique to humans.

What else about

human sexual behavior

seems to be specializations?

One that is fairly unique,

if not entirely so,

is the subset of humans who

psychopathological confuse

sexual behavior with violence.

And that seems not to have

a whole lot of precedence.

So immediately one
asks other domains.

Masturbation, that is not

remotely a human specialty.

That has all sorts of

other species as well.

And that used to get the,

well, what else is there

to do when you're sitting in

the zoo-- for the animals-- what
else is there to do?

It is not natural.

It is a default whatever.

But looking out

in the real world,

and there is plenty of that.

And one of the most like

implausible suggestions

for an adaptive
just so story thing

is, why do male

primates masturbate

to the point of ejaculation?

They tend to eat

the semen afterward.

Whoa!

Great source of protein,

go the adaptationists.

Everything has an
adaptive basis.

This one does not ring

terribly true to me.

What else?

Fantasy.

Fantasy in humans, is
that unique to humans?

Obviously, we haven't a clue.

But here's one suggestion to

me that this is not actually

the case.

And this was years ago where

I was watching my baboons,

and there was this

one low ranking kid,

this snivelly adolescent kid,

where the nearest thing he has
ever gotten to a
female in his life

with some high ranking female

in a bad mood beating on him.

And he's sitting there

minding his own business.

And along comes, I think by

any baboon male standards,

the hottest female in the troop,

who has a peek estrus swelling,

is no doubt ovulating that day.

Comes walking along,

followed two feet

behind by the huge

menacing male,

who was in the

consortship with her.

And our guy just sits there

and doesn't even quite

look at what's happening.

Every now and then, his eyes

go up, watching them go past.

And as she walks past,

he gets an erection

and then goes off

and masturbates.

OK.

The charming-- I don't

know if this is even

the word you can use

in this setting--

but the charming notion is

that we've just seen evidence

for some sort of internal

fantasy life going on this guy.

OK.

You could be a killjoy

instead and say, no, no.
She was giving off
wafts of pheromones,

and that was what was

responsible for it.

Nonetheless this is
about as far as we

can get at asking this

critical question,

are we the only species that

does this fantasizing stuff?

Marriage.

Clearly, we've heard about

monogamous pair bonding

species.

In terms of the formal structure

of marriage, it is universal.

All human cultures have

some version of it.

Across all human cultures,

more than 90% of people

wind up in that
culture's equivalent

of a permanent,
stable relationship.

And this is the case

in polygamous cultures.

We've already heard

that business.

Even though historically, the

majority of human cultures

have been polygamous,

nonetheless, amid them,

the vast majority of

individuals have been

in monogamous relationships.

Amid that, nonetheless,

what is also

clear is amid that highly,

highly prevalent pattern
of monogamous relationships,
there's a lot less monogamy

going around than

you would think.

And this was first sorted

out-- people like Alfred Kinsey

when first working out

that questionnaire approach

to people's sexual
behavior, what became clear

was there is a lot less

faith within pair bonding,

within humans in this

country and has since

shown in all sorts

of other societies

than one would

originally assume.

There is social monogamy but

not necessarily anywhere near

as high of rates
of sexual monogamy.

And what the paternity

studies have shown

is in most Western
European countries,

the rate at which children

have been fathered

by an individual other
than the person claiming

marriageable credit for doing

so ranges between 10% and 40%

of children.

How's that for a number?

OK.

What else?

What else tends to be a feature

of human sexual behavior?
What you have is, of course,
not only intrinsic in the fact

that there's a difference

between social and sexual

monogamy.

You have cheating.

That is a human specialty

in every culture.

What else is absolutely

wildly human?

This notion of romance.

And romance is,

by most estimates,

a relatively new invention

in most cultures,

maybe a couple of centuries old.

And what is an even

newer invention

is the notion that romance,

passion, et cetera,

should persist, should last

throughout the entire duration

of the lifetime's marriage.

That is an utterly
novel concept.

That is perhaps 30,

40, 50 years old

in most Westernized countries.

That's a new one as well.

What that, of
course, ushers in is

looking at issues of divorce.

Across all cultures, the

average duration of marriages

are two to four years.

And people have

made the argument

that that is the typical

duration of children being

dependent on a high
degree of parenting,

of both parents being around.

That is the average interbirth

interval, two to four years,

in most traditional
human cultures.

What's the term

being described then

if that is the "natural"

point at which most

marriages dissolve and turn into

other monogamous relationships?

The term that is

given is that humans

tend toward being serial

monogamists, moving from one

monogamous relationship to
another with, on the average,

a lag time roughly corresponding

to the interbirth interval.

So that's charming.

What else?

What else do you have?

All sorts of other aspects

of human sexual variety,

but ultimately, when you

look at human sexual behavior

versus other species,

we are so boring.

We are so limited when you look

at the range of unlikely things

going on out there.

Species that are

regularly hermaphroditic--
and people, in fact,
have done studies

on how is it that a
hermaphroditic animal does not

try to have sex with itself?

And these are usually

worm type things.

And that's some version

of an incest avoidance.

At the same time, there are

other species where individuals

change sex opportunistically.

All sorts of fish species

where that happens.

Lots of species that

are parthenogenetic,

where an individual reproduces

without the benefit of anybody

else's genetic input.

Even stranger, there's a bunch

of snake species that are

parthenogenic, but the

females cannot reproduce

parthenogenically unless
they mate with males.

They do not actually get

any sperm from the males,

and they get no

genetic contribution,

but something about

that is necessary

for the parthenogenetic

event to occur.

OK.

So all sorts of bizarrities that

make our fixed action patterns

look really pretty dull.

But nonetheless, these are the
backbones of the human fixed

action patterns,
and some of them

wildly unique, some

of them far less

than people used to

think, some of them

very, very unprecedented.

OK.

So what this allows us to do

now is make our first big step.

What's going on in the brain?

What is the neurobiology

of sexual behavior

producing those fixed

action patterns?

And what you better bet

right off the bat is we

are talking about

the limbic system.

This is all limbic

system until we

see ways in which it's not

just all the limbic system.

But it is heavily
centered-- no surprise--

in the limbic system.

And this was being noted

first around the 1930s, 1940s

with the first experiments where

there were lesion studies done

damaging different parts of

the limbic system in animals.

And what would be noted

was animal sexual behavior

would change.

And this was eventually

termed a profile,

termed after the two scientists

who pioneered this stuff called

the Kluvre-Bucy Syndrome,

which is when you damage some

of these strange mysterious

rhinencephalonic structures

in there, you change

the sexual behavior.

You change, for

example, in monkeys,

whether they are attempting

to mate with another monkey as

opposed to an inanimate object.

They change aspects of

the fixed action patterns

of the behavior and such.

And out of it, this was

one of the main driving

forces on people saying,

nose-brain, well, that's great.

But actually, what

we're looking at

is a part of the brain that

has lots to do with emotion

and emotionally
related behaviors.

That was one of

the driving forces

on the limbic system being

pulled together as a concept.

OK.

So what areas within the

limbic system are relevant?

First pass, there are

different hot spots

in there depending on gender.

Among females, probably

the most important area

is a subsection of the
hypothalamus called

the ventral medial

hypothalamus involved

in female sexual behavior.

What's the evidence for that?

Just go back to last

Friday's lecture--

lesion studies, stimulation

studies, recording studies.

Destroy the VMH, you do not

get sexual behavior anymore

from a female.

Stimulate it, and you will

get the same behaviors

that you would normally

only see in an ovulating

female rat for example.

All the sorts of

tools we heard about.

Reinforcing this
even more is this

is the hot spot in the

hypothalamus for receptors

for estrogen and progesterone.

So that makes lots of sense.

Meanwhile, another region of

the brain that is typically

involved in sexual
behavior in females,

a region in the midbrain.

The midbrain, which seems to

have something to do with some

of the hormonal aspects

of sexual behavior

that are specific to females.

Finally, back to
that lordosis reflex,

you got to have a

spinal cord to pull off

the full array of typical

mammalian female sexual

behavior.

So spinal pathways,
which do not exist

in males, lordotic reflexes,

the back arching reflex,

is exclusively a female one.

Meanwhile, over on the

other side of things,

there are regions

in the brain that

tend to be more specialized

for sexual behavior in males

than in females-- a different

part of the hypothalamus

called the medial preoptic area.

And the exact same

sort of evidence

is for the ventral

medial hypothalamus

in females-- lesion studies,

stimulation, recording studies,

all that sort of

thing, and-- you

guessed it-- whopping

great amounts

of testosterone receptors,
androgen receptors,

within the medial preoptic area.

Very interestingly, something we

will hear more about next week

or so, is another
region of the brain
is involved in male
sexual behavior, which

is the amygdala.

Mhm.

That's kind of interesting.

The amygdala.

We've already heard about

amygdala fear, anxiety,

and all of that.

But the amygdala also plays a

very major role in aggression.

And there's a little

bit, a small domain,

of amygdaloid function
in males that's

involved in sexual behavior,

involved in sexual motivation.

Medial preoptic area is much

more about sexual performance

in males.

Amygdala is much more

about sexual motivation.

And all sorts of people have

speculated fairly reasonably,

I think, that this

may have something

to do with the fact explaining

why, among humans, it

is far more likely to

be males than females

who go about confusing

sexuality with aggression.

That it's got something

to do with this weird role

of the amygdala in
male sexual motivation,

male sexual arousal.

What else?

OK.

Males have penises, thus

they're the only ones

who can have penile

erections, and to do

that, autonomic nervous system.

And what we will hear

about, what you already

heard about in the introduction

to the autonomic nervous

system, but also in

the zebra's book is

that whole business in order to

manage that, to pull that off

initially, it is
parasympathetic nervous system

that establishes the erection.

The process of arousal involves

the transition to sympathetic.

Full blast sympathetic

nervous system

needed for ejaculation, that's

what all of that is about.

Exclusive to males.

But then it turns out it's

not exclusive to males

because it's virtually the

exact same physiology underlying

clitoral erections in females,

the same exact sort of thing,

which, of course, brings up

the dangerous possibility

of another spandrel in
our laps here in terms

of making sense of that.

I didn't say that just now.

Did I say spandrels in your lap?

OK.

Bringing up that
possibility that is not

specific to male physiology.

What is specific,
of course, is stuff

that's going on with penises

in terms of blood flow.

There's generally a
dichotomy between species,

between whether or not

males get vascular erections

or muscular erections.

Vascular erections, you increase

blood flow into the penis,

and you stop it from

going out the other end.

And thus, you get a

vascular-driven engorgement

as DH Lawrence would no
doubt have described it.

Alternatively, in
lots of species

there are muscular erections.

There is a muscle, for example,

found in rodent penises called

the erector [? levae ?]

muscle-- well,

that's not too surprising

that it's called that--

and a whole bunch of

cell bodied neurons

in the spinal cord responsible

for pulling up the sail

or whatever it is you do there.

And what you get are

differences in general.

The muscular-driven
erections occur a lot faster.

The vascular ones,

the hemodynamic ones,

last a lot longer.

Take your pick, but

it's essentially

the exact same autonomic

physiology in both cases.

Finally, one other

thing, a factoid,

a useful one we heard last

week, which is insofar

as there is very
similar physiology

to orgasms in both sexes,

there is that difference

in recovery time,
how long it takes

for the sympathetic

nervous system

to go back to
baseline post orgasm.

And on the average,

a substantial sex

difference in that.

Yes, everybody managed

to guess it last week,

which direction it went.

A far slower recovery time

in females than in males.

In terms of underlying
neurobiology,

something that was a

major finding in the field

were brain regions that

differed in size depending
on your gender,
including in humans.

And this ushered in a whole

world of sexual dimorphism

in the brain.

And there have now been shown

to be all sorts of brain regions

where, on the average,

you get differences

in the size of nuclei.

You get differences

in the number

of axons going through a

bundle of fiber, all of that.

And we will hear about some

more of those down the line.

But the one that has gotten

the most attention in terms

of sexual behavior is a
cluster of tiny nuclei

in the hypothalamus
called the INAH cluster,

the Interstitial Nucleus of

the Anterior Hypothalamus.

Do not write down

what that stands for.

But it's a little

nucleus in there

there, a little subset

of neuronal cell bodies,

where you get a very

substantial sex difference

in the size of this area,

where on the average,

it is about twice the

size in men as in women.

And back to the

other week's rant

about statistical
significance versus magnitude,

this is a big effect.

It is almost,
almost in the range

where you can identify

the sex of somebody

by looking at the size of

this nucleus in their brain

post-mortem.

In rodents, you pretty much can.

A very reliable two-fold

difference, males

larger than females.

As we'll hear in a while,

one really interesting

exception to that.

OK.

So either some areas of the

brain that are preferentially

involved and activated

by sexual behavior,

depending on your gender,

or regions that differ in

size substantially
by your gender.

But then, at the end

of the day, there's

all sorts of things that

are absolutely in common.

Again, the physiology of

orgasm, exactly the same.

What clinically the

picture is is males having

problems with the whole system.

The problem tends to be

too rapid of a transition

from parasympathetic
to sympathetic.

In other words, the world

of premature ejaculation.

The more typical

medical problem in women

is failure of the
transition from

parasympathetic to sympathetic,
inability to reach orgasm.

And it is, of course, a huge

social, cultural, political,

philosophical
argument whether that

counts as a pathology or
normal human variability.

I'm not going anywhere

near that one.

But nonetheless, that is

the more common pattern.

Neurobiology that's
absolutely in common

between the sexes, which

is all of that stuff

at the very beginning of why

are they in such a rush at end,

the neurobiology of pleasure,

and the neurobiology of reward,

and of anticipation.

And this is this

whole world of-- as we

know already-- dopamine.

The role of dopamine

in sexual behavior

is virtually identical
in both sexes, which

is to say it plays a huge role.

You find circumstances

where you deplete dopamine
from the relevant
brain regions--

back to last Friday--

limbic system.

You remember that

ventral tegmental area,

which sends that big

dopaminergic projection

to the nucleus accumbens,

which then passes it

on to all sorts of
places in the brain.

Deplete that
pathway of dopamine,

and you're not going to

get a whole lot of interest

in sexual behavior.

You're not going to get a whole

lot of libido proceptivity.

What's the classic

circumstance where

you see depletion

of dopamine there

and loss of proceptive libido?

Clinical depression.

That's one of the defining

symptoms of depression

amid the various numerous

forms of pleasure

that go down the tubes.

Loss of sexual interest, one

of the defining symptoms.

So the dopamine system.

The general term

given for that is

the mesolimbic dopamine

system to distinguish it

from some of the other ways that

dopamine is used in the brain.

The mesolimbic
dopamine system is

absolutely central to
the reinforcing aspects

of sexual behavior.

So what's the evidence for that?

First off, back to

that distinction

that I think I brought up

last week-- I wasn't paying

attention-- but I
think I talked about,

which is the dopamine

system there is not

so much about reward, it's about

the anticipation of reward.

Did I talk about that in

monkeys pressing levers?

Yes.

OK.

I should probably read the

extended notes at some point

or look at the film of this.

OK.

What you see there

is the dopamine

is about the anticipation of.

And the dopamine,

as we heard, is

about also fueling the behaviors

needed to achieve the reward.

Dopamine in this
mesolimbic pathway

as driving
goal-directed behavior.

And that is certainly the

case with sexual behavior.

By now, there is a whole

literature involving humans

where you stick them

in brain scanners

and you do something

or other sexually

arousing or interesting or
something or other to them.

And then you see what parts

of the brain activate.

And it's these dopamine pathways

consistently way up there.

Showing just how subtle

this can be, how's this?

You take men-- there's been a

whole literature by now showing

that you present people in

brain scanners with pornography.

And that must have been a really

interesting in human subjects

release form you worked out.

But showing in both sexes,

what you tend to see

is activation of
dopaminergic regions.

We will hear in a little

while a sex difference

in that domain that will

probably not surprise anyone.

But how's this for subtle?

You take a guy, and you

show him the picture

of someone of the opposite

sex if he is heterosexual.

And you show him the

picture of this individual.

And if it is someone who he

assesses as being attractive,
you don't necessarily get
this dopaminergic pathway

to activate.

It depends.

What this study showed was

if the person is making what

would pass for eye contact, if

they were looking straight out,

the dopamine system activates.

And if they're looking

elsewhere, it doesn't activate.

How's that for a classic

male sort of responsiveness?

If it looks as if this
attractive person is

looking at you, it activates.

Even more distressingly

from this study,

when you show men--

on the average,

blah, blah--
pictures of women who

they would rate as

being unattractive,

it's when they're looking

away that the dopamine system

activates.

Oh my god!

What is going on here?

This is pitiful.

What also has been shown is

the exact same eye contact

phenomenon of gay men looking

at pictures of attractive men.

Another theme we're going

to see over and over, which
is sexual orientation
being pretty much

trivial in terms of
how it influences

some of this neurobiology.

Just switch the gender

of the other individual,

and it works exactly the same.

Now when you look at this

business about dopamine rising

in anticipation of a reward
rather than a response

to the reward itself, it

brings up one of the-- it

doesn't bring that

up-- it brings up

one of the all-time

interesting studies

that was published

about a decade ago.

OK.

So the paradigm I
described last week,

you put on the light, which

tells the monkey that,

OK, we're starting

one of those sessions

where if you press

the levers adequately,

you will get a reward.

And they now carry

out this behavior.

And as a result, they

get the reward here.

And as we saw, dopamine

doesn't go up after the reward.

It goes up at this point.

This is the I how to do this.

This is going to be great.

This is terrific.

Here's where you get

the rise in dopamine.

This is not only

the anticipation,

but if you don't

have this rise, you

don't get the behavior,

the goal-directed behavior.

Now in this brilliant

study, what they did

was transition from

a paradigm where,

OK, the monkey presses the lever

10 times and gets the reward.

Now what you do is

the monkey works,

and it gets the reward

only half the time.

It gets only a 50% reward

rate unpredictably.

And what happens to dopamine?

OK.

Got your choice.

What's your vote?

It doesn't rise as much.

It rises the exact same amount.

It rises even higher.

OK.

You guys all understand

anticipation and goal--

it does this.

It's one of the

biggest rises you
will find in dopamine in
the brain short of cocaine.

What have you just

introduced into there?

This is, I'm all over it.

I know how this works.

This is going to be great.

I have mastery and control.

I am the captain of
my own lever pressing.

This is all about that.

What's this about?

This is what dopamine

does when you've

introduced the word

maybe into the equation.

And that is incredibly

reinforcing.

And people will work like

mad in contexts of maybe

far more so than when they

work in contexts of certainty.

Psychologists have
known this forever.

This is intermittent
reinforcement.

You never get more

behavior out of an organism

than when you have

introduced a maybe into it.

And part of the

brilliance of this study

was what they then did.

Now animals either got

reward 25% of the time

or 75% of the time.

On a certain level,
these are diametrically

opposite manipulations.

In one, you're
getting more rewards.

In the other,
you're getting less.

What's the thing

they have in common?

They both had smaller

maybes than the 50% version.

And what you see is it

would look like this.

100%, 25% or 75%, 50%

maximizing the maybe.

And one of the most

brilliant things

that various social

engineers do with humans

is convince people that there's

a 50% maybe when it is not

50% in the slightest.

That's what Las Vegas is about.

That's an entire world of very

smart psychologists making

people think in
circumstances where there's

like one tenth of 1% of a maybe

going on there that is actually

a 50%.

And when you do that, you

get dopamine like crazy,

and you get

goal-directed behavior

as a result. Really,
really powerful.

And this is so strongly the case

that this explains an extremely

cynical thing that a

guy I knew in my dorm

back when used to

say all the time.

How's this for like a

dispirited view of what

life is like, but possibly

absolutely accurate, which is,

a relationship is the price you

pay for the anticipation of it.

How's that for a grim worldview?

Go figure.

This guy had-- what a string

of disastrous relationships.

But what you see here

is introduce a maybe,

and it is very, very powerful.

One final piece of the dopamine

system here that is pertinent,

which is, as you

might expect from all

of our molecular
biology stuff, there's

all sorts of different

dopamine receptor subtypes.

And two of them are pertinent

to this world of sexual behavior

and reward, what is called

the shockingly the D1 dopamine

receptor and the D2

dopamine receptor.

And what studies show is

in monogamous species, what

happens is right after mating,

when a pair bond is first

formed, the second that's over

with, levels of the D2 receptor

go way down.
You down regulate the
levels of the receptor,

and you up regulate the

levels of the D1 receptor.

What's that about?

If you drive down the D2

levels before they even mate,

they don't form a pair bond.

If you prevent the decline in

the D2's after they've mated

and pair bonded or if you

prevent the rise in the D1's,

they'll pair bond.

And then, 8 and

1/2 minutes later,

they will go and pair

bond with somebody else.

The D2's seem to mediate the

rewarding anticipatory aspects

of pair bonding.

The D1's, on a certain

rodential level,

seem to mediate the pleasure

of the monogamous, the truly

monogamous features
of the pair bond.

So a very interesting
interaction between the two.

OK.

One last thing about

dopamine, and this one

is like even more depressing

than relationships

are the price you pay.

This was a study which

was really like someday

may come to haunt you majorly.

And in this study,
what they did--

it was another one of those

brain imaging study ones.

And what they did was they

took people in two categories.

In both cases,
these are people who

had found their beloved,

their beloved, the person who

was their soul mate,

the person in whose arms

they were going to die

someday, the person.

And they divided it

between these two groups.

One was a group where they had

known the person in whose arms

they were going to die

for like 2 and 1/2 weeks.

And the other is when

they had been together

for more than five years.

So you put somebody

in the brain scanner,

and you start flashing up

at speed, subliminal speeds,

of pictures of
individuals they know.

Important control in the study.

And embedded in there is a

picture of their beloved.

And suddenly, somewhere

along the way,

up flashes the picture

of their beloved.

Be in a short-term relationship
and the dopamine system
goes crazy and
activates like mad.

Now, you come back

five years later

into that same relationship

with the beloved,

and you do the exact same thing.

And you flash up their picture,

and the dopamine system

doesn't activate.

What activates instead was

that anterior cingulate thing

we heard about on Friday

having to do with empathy,

and comfort, and all of that.

In other words, what we see here

is the neurochemical transition

from one's beloved

from causing your blood

to run scalding hot

to your beloved being

like a comfortable old armchair.

This is one depressing study.

So let's take a five-minute

break to contemplate that one.

OK.

And then we will resume.

Lots of good questions

just now during the break.

Disappointingly few
along the lines of,

I've got a friend who.

So not a bunch of those.

But let's see.

A number of questions.
First one, can I repeat what
I said about the D1 and the D2

receptors?

OK.

These are different types

of receptors for dopamine.

In other words, they all respond

to the same neurotransmitter

dopamine, but in different ways.

And these receptors are found

on different neuron types.

So you're getting into all

sorts of different pathways.

What you see is, in rodents,

in pair bonding rodents,

they better have

elevated levels,

they better have D2

receptors on neurons

being fed by this mesolimbic

reward dopamine pathway.

They have to have D2 receptors

to form the pair bond

for the attachment to occur.

The second that happens, you

need to have low levels of D2

and high levels of D1 to remain

faithful in your pair bonded

relationship.

So what you see in these voles

is right after the pair bond

occurs, there's down regulation

of the D2's and up regulation

of the D1's.

And if you prevent

that from happening,

the pair bond that's been

formed does not prove lasting.

So that's what I
was saying there.

Somebody brought up
the great question,

which I was going to say

something about and forgot,

which is, well, how

about like the D2 D1

ratios in humans and their

sexual behavior stuff?

And there's been

one study showing

that a higher ratio

of D2 to D1 predicts

more stable relationships.

Small effect.

Not replicated yet.

But nonetheless, that's

kind of interesting.

So on a certain
level then, D2 seems

to be required, at
least shown in rodents.

Who knows about us?

D2 is about the formation

of the attachment.

D1 is about the maintenance

of it, the faithfulness of it,

if you will.

Next, somebody
bringing up the issue

in terms of female orgasm.

Maybe what female orgasm is

about is a mate selection

mechanism, as in
individuals who increase
your likelihood of
having orgasms are ones

you are more likely to

lower your D2 receptors for.

But the one problem

with that one

that makes wonderful sense--

what the studies tend

to show though is the

likelihood of orgasm

is much more a function

of who the female is

than who the male is that they

are with, arguing against that.

Let's see.

Finally-- no.

Not that.

OK.

So we already covered that.

And there was one

additional question.

OK.

So what's the driving

force in terms

of the proximal reinforcing

pleasurable aspects

of sexual behavior?

What's up with why

only some species--

us predominantly-- have
non-reproductive sex,

can have sex all the

time, versus other species

that only do for reproduction?

What that means is

in other species,

the endocrinology of
ovulation is the thing

that makes sex pleasurable.

And we will see shortly

what that's about.

In females, it's the

hormones associated

with ovulation that

sensitize various tactile

receptors to respond in ways

that mediate proximal pleasure.

And in males, it's

the female giving off,

for example, the right

pheromones, the right releasing

stimuli, driven by the

right hormone levels that

constitute the proximal signal

of pleasurable anticipation.

And what you find in humans

is it doesn't work that way.

You do not need, for

example, in women

the elevated levels of estrogen

typical of ovulation in order

to have tactile responsiveness

to sexually arousing stimuli.

Stay tuned though.

It's easier though when

estrogen levels are higher.

OK.

Final brain region

relevant to all of this

is the frontal cortex.

We already got a first pass at

the frontal cortex last week.

And frontal cortex regulating

your behavior, impulse control,

all that sort of thing,

gratification postponement--

this plays a large role

in sexual behavior.

What's the easy

immediate explanation

that one can come up with,

what the frontal cortex does

is it makes you be appropriate

in your sexual behavior.

It teaches you the

appropriate context.

It teaches you what aspects

of proceptive sexual behavior

is not a good idea.

It keeps you from doing

things you would regret vastly

afterward.

That's a very easy

version of it.

And commensurate
with that, you see

lots of circumstances
of individuals

with frontal cortical damage

doing highly inappropriate

sexual behavior.

One example of it, and one

of those horrifying things

that can happen--

this was a case

that actually happened in

a nursing home in Martinez

in the East Bay a

number of years ago.

This was a man in his 80s

who had had stroke damage

to his frontal
cortex, who was found
to have raped a woman
there, another 80-year-old

with Alzheimer's disease.

Damage the frontal

cortex, and all sorts

of the, "this is not sexual

behavior that you do"

constraints go down the tubes.

Just as importantly though,

what the frontal cortex,

with all of it's giving

you the discipline

to do the right thing,

some of the time,

what that takes the

form of is getting

you to do proceptive
sexual behavior.

For example, you

are trying to do

some courtship of some

other antlered ungulate

that you are courting.

And this is terrifying because

there is another individual

challenging you.

And there's the

frontal cortex that

is getting you to carry

out those sexual behaviors

to that point, even if it is

a terrifying circumstance.

Nonetheless, what the frontal

cortex mostly is about

is reigning in sexual behavior.

It's not changing the fixed

action patterns of sex.

It's changing the context

in which the fixed action

patterns occur.

So now, we are ready

to look at one more

feature of the
neurobiology, which

is when somebody is having sex,

what hormonal responses are

triggered?

Notice this is not here.

This is not what hormones

have to do with bringing

about sexual behavior.

This is, what are the hormonal

responses to sexual behavior?

Starting off in females,

including human females,

having sex increases

secretion of

progesterone-derived hormones.

And that has something to do

with reinforcing the pleasure.

Interestingly, in females
the world over, having sex

increases the level of

testosterone-related hormones

in the bloodstream, androgens.

Women, females, generate

androgens maybe 5%

the levels you see in males.

And they come out of

the adrenal glands.

And they seem to play

a very central role

in mediating sexual motivation,

sexual arousal, in females.

How is that's shown?

Obvious experimental
studies with lab rats.

How is that's shown in humans?

When women have any of a number

of types of diseases where you

have to take out

the adrenal glands,

sexual motivation, sexual

arousal, goes down.

Give them replacement

androgens, and

sexual arousal, sexual

proceptivity, returns, so

androgens there playing a role.

But probably most

importantly, in terms

of hormones triggered by
sexual behavior in females,

is the release of oxytocin.

Oxytocin is really interesting.

We've heard about

oxytocin twice already.

One, is when it's coming out

of the posterior pituitary.

And the second is that minor

business the other day,

last Wednesday,
of oxytocin being

another one of those

hypothalamic hormones that

helps to release ACTH

from the pituitary.

Remember, it doesn't
directly release.

It's a modulator, a CRH action,

if-then clause, et cetera.

But those are the two ways

we've heard about oxytocin.
Oxytocin also works in the
brain as a neurotransmitter

and neuromodulator.

And what does it do there?

It appears to play
a very central role

in forming attachments,
a very central role

in forming of pair bonds.

And it, along with dopamine

and the D2 receptors,

are critical for female

voles of monogamous species

to form pair bonds.

Female humans, when having

sex, secrete lots of oxytocin

and activate oxytocin

pathways in the brain.

And it appears to play a role

in the formation of attachment.

Interestingly-- how's this--

a whole body of research

now showing that if

you introduce oxytocin

into the brains of

humans experimentally,

they become more trusting.

Amazing body of research where

you take aerosolized oxytocin

and you spritz it

up people's noses.

And what they showed

in the studies

were, number one,

one type of study.

You then play a clip of

somebody making an argument
for some stance in
some debate, and people

believe the person more.

They find their argument

more convincing.

They trust the person more.

Or the other version

that's been shown

is you spritz oxytocin

up the noses of people,

and you make them more

cooperative in their game

theory play, the

ways in which they

go about playing prisoner's

dilemma and other games

we will hear about

later on as well.

This has given rise to a whole

new field-- I kid you not--

called neuromarketing.

The notion that if only you

could spritz oxytocin up

the noses of people right before

your television ad comes on,

they're going to believe

you when you say it

will make you happy

to buy our thing.

And they will fall for it.

There are actually

neuroendocrinologists

making a living now selling

their wares to neuromarketing

people-- self-proclaimed ones.

No doubt they are

spritzing oxytocin up

the noses of those

capitalists to get

them to hire them to do this.

But oxytocin playing

a role in this.

So that's kind of interesting.

Because what's oxytocin

mostly doing in the body?

The vast majority

of oxytocin is not

this stuff up in the

brain in these pathways,

some of them impinging on

dopamine-releasing neurons.

The vast majority is not

the oxytocin sitting there

in the hypothalamus doing

something or other to ACTH

in the pituitary.

The vast majority is this stuff

coming out of the posterior

pituitary.

And what does oxytocin

have to do there?

It has to do with milk letdown.

It has to do with nursing.

And suddenly, instead,

it's playing a role

in forming sexual pair bonding.

And the argument is

made that attachment,

monogamous attachment, sexual

attachment, is in some way,

evolutionarily a descendent
of the neurobiology

of mother-offspring attachment.

That that's where it is

originally being driven by.
So oxytocin playing
a role there as well.

Meanwhile, over at the

male end of things,

up go testosterone
levels during sex.

In a surprisingly linear
way, the more sexual behavior

in a male, the higher

testosterone levels

are found afterward.

Critically, critically--
stay tuned for a little

while-- these are elevations

of testosterone in response

to sexual behavior.

As we'll see, the evidence

that high testosterone levels

make males more sexually active

is basically nonexistent.

So critical, critical
proviso here.

This is sex increasing

testosterone secretion, not

the other way around.

What else?

Meanwhile, back to the

posterior pituitary.

Was that a question?

No.

OK.

That was a head scratch.

OK.

Back to the posterior pituitary.

The other hormone coming

out from there, vasopressin.
And oxytocin is to females
as vasopressin is to males.

And we've already

heard something

about this back in

the molecular genetics

stuff in those if-then clauses

and unlikely ways in which you

get mutations.

vasopressin, vasopressin
also is found

in the nervous system, where it

serves as neuromodulatory role.

And vasopressin is critical

for forming a pair bond.

Back to that business

about when you

look at monogamous versus

polygamist species,

what's going on?

What you have uniquely

in the monogamous

species is expression of
the vasopressin receptor

gene on neurons that

released dopamine.

In other words, male

secrete vasopressin.

And if you are of a species

where vasopressin now goes

and stimulates dopamine

neurons, you decide you really,

really, really liked having

sex with this other vole.

And you come back for more.

And that's the driving force on

the formation of the pair bond.

Incredible studies
showing that if you

take male voles from

the polygamist species

and, due to gene

transfer techniques--

I've mentioned the

study already--

but you now stick

vasopressin receptors

into those dopamine neurons,

those polygamist males

now become pair bonding males.

They now become monogamous.

So really interesting.

What you then see

in these studies is

you look at these monogamous

species where the males have

vasopressin receptors on
these dopamine neurons,

and you look at

individual males,

and the ones who

have more receptors

there are forming

pair bonds faster.

It takes fewer rounds

of mating with a female

to form a pair bond.

So what about primates?

So you start off looking at two

different primate species, one

pair bonding, marmoset monkeys,

New World marmoset monkeys,

and then one classic

tournament species,

polygamous primate
species, rhesus monkeys.
And what you see is
you've got the variant,

the monogamous vole,

vasopressin receptor gene

variant in the pair

bonding monkey species,

in the marmosets.

And you get the

polygamous version

of the gene in the

rhesus monkeys.

So it maps on there as well.

So the more of this receptor

in these dopamine pathways

within monogamous
species, the more rapidly

they form a pair bond.

And what you see is differences

in the mere presence of them

in comparing pair bonding

versus non-pair bonding rodents

and monkeys.

So how about humans?

First thing that comes

up is, among the apes,

you also find the two

variants, the monogamous vole

variant of the
vasopressin receptor gene

and the polygamous male variant.

So what species
do you see it in?

In chimpanzees, you see

the polygamous vole species

version.

That makes lots of sense.

They are a major polygamist
species in their behavior.

But then, this

beautiful dichotomy

comes crashing down

when you see that you've

got the monogamous gene

version in bonobos.

And as we will hear

about in a while,

bonobos are the most

hyperpolygamously, hyper

varietyish sexually behaving

organisms on the entire planet.

They are as far as you could

get from a monogamous species

as you can ever ask for,

if you ask for such things.

And you've got the wrong type

of vasopressin receptor gene.

Whatever is going on,

it's more complicated

than the have the

version that winds up

on the dopamine
neurons and you are

going to have 50th wedding

anniversaries if you are a vole

or you are a marmoset.

And it's got to be more

complicated than that.

So how about humans?

And what you see

is not explicitly

as much genetic
variability as some people

having the monogamous vole

version and some people

have the polygamist.

But nonetheless,
you get variation.

The gene basically is

about halfway in between.

Whoa.

We keep having that

theme over and over,

all these different ways

of looking at body size,

and sexual dimorphism,

and imprinted genes,

and all that stuff.

And humans keep winding

up being about a halfway

between a classic
monogamous pair

bonder and a classic

polygamist tournament species.

So the basic human version of

it is somewhere in between.

But you get variations.

You've got genetic variations

that either look a little bit

more like the monogamous vole

version or the polygamous vole

version.

And what studies have now

shown-- two different studies

independently showing this--

have the monogamous vole

version.

And with the small effect, you

are more likely to get married,

you are more likely

to remain married,

and both you and

your partner are
more likely to rate the
marriage as stable and happy.

That's kind of interesting.

Finally, in terms of
the role of vasopressin,

in terms of attachment
in males, all of that,

and in terms of
social connectiveness,

a large body of
studies now have shown

mutations in the
vasopressin receptor gene.

OK.

Anybody want to guess

what disorder you find it?

I put that in the

extended notes, haven't I?

People have read it already.

I know, I fell for it last time.

I am not falling for

that stupid trick

again of asking you to prove.

He sniffed some oxytocin.

So does anybody want to

guess which the-- OK.

What do you now all

know is there's now

been a lot of
demonstrations of mutations

in the vasopressin
receptor gene in family

pedigrees with autism, a disease

of very, very little attachment

to other humans.

So we've got written

all over the place here
some sort of role of oxytocin
in female attachment formation

and vasopressin.

And we've already

gotten interesting hints

that this applies to humans.

And we've already

gotten interesting hints

that individual differences

in the molecular biology

of these genes
predicts something

about individual differences in

the stability of relationships

in humans.

OK.

So everything we've been hearing

about with the neurobiology--

and we're still living in

this part, in this bucket here

temporarily-- has
been built around

heterosexual relationships.

What's known about

the neurobiology

of sexual orientation?

What has been found is most

strikingly one landmark study,

one that got on the

cover of Time magazine,

one that had a gigantic,

gigantic impact,

which was looking back at

that hypothalamic nucleus,

that INAH, I-N-A-H. Yes.

Where what we saw

before was very reliably

on the average, men, their

size of it is about twice

the size as in women.

And in other species,

males about twice

the size as in females.

And a study was

done in the late 80s

by a neuroanatomist
names Simon LeVay.

LeVay is one of the all-time

great neuroanatomists.

He was trained by
Hubel and Wiesel,

was a professor at
Harvard Med School

for a while before moving

to the Salk Institute.

And what he did was

look post-mortem

at the brains of a bunch of

individuals where he knew

the sexual orientation, men.

And what he showed was gay

men, on the average, had

this nucleus on the

average was half the size

that you saw in

heterosexual men.

On the average, it was

about the same size

as you saw in
heterosexual women.

Amazing landmark study.

Everybody learned about

the homosexual brain

from this study.

Hugely widely reported.

And this is kind of interesting.

OK.

What's interesting about it?

First off, the question you need

to ask is how much variability?

Fair amount.

It wasn't all that

reliable of a difference.

On the average, it was

about a half the size.

Next thing you

would want to know

is, has anybody

replicated it since then?

Yes.

Next thing you

would want to know

is, where did LeVay

get the brains from?

And these were

predominantly from gay men

who had died of AIDS.

Is that a confound?

Is that going to perhaps

atrophy this part of the brain?

Nobody knows.

So that remains as a
caveat in that study.

What was striking

though was everybody

learned about this finding.

This became famous.

LeVay became extremely

famous for this.
And what was also
very interesting

about it was the political

context of this finding.

A few years before

that, another group

had reported another

difference in the hypothalamus

based on sexual orientation.

And this was-- there it is.

And what you found was in

this part of the brain,

on the average, it would tend

be bigger in women than in men.

And what these guys

reported was in gay men,

it tended to be bigger
than in straight men.

What was puzzling

about the study

was this was a part of

a hypothalamus having

to do with regulation
of your kidneys.

And it was totally ignored

and completely bizarre,

except there was

one thing that was

done with it, which

was this was viewed

as a totally offensive
study in the gay community.

This was viewed as an

attempt by scientists

to pathologize sexual
orientation, to say, you see,

we found something wrong in

the brains of homosexual men.

This part of the brain is

bigger than it should be.

It's bigger than

it's supposed to be.

There's probably two

reasons why that occurred.

The first one was that

the scientists who did it

were straight.

And the second reason being

that they were a Dutch group.

And I am willing to
bet unconsciously there

was something central European

Nazi echoes of Germanic

sounding authors producing

this finding, which there

is no shortage of history in
the gay community for being

skittish about Nazi

notions of what normality

is in human behavior
and human brains.

This finding got widely

condemned in the gay community.

Out came LeVay with his

finding, and he became the most

beloved neuroanatomist ever in

the history of gay communities.

One probably important

feature reason for it

is that LeVay was

gay, very openly so.

Another reason was the part of

the brain he found made sense.

It had something to do with

sexual behavior as opposed

to the totally puzzling thing.

So what was this about?

Very interestingly, the
explanation almost certainly

is that this was the

part of the hypothalamus

next door to the area

that LeVay studied.

And if this part were

smaller in gay men

simply because of just

physical constraints stuff,

this part could get

bigger in gay men.

Because this one was

taking up less area.

That's probably
what was going on.

What's really fascinating

though is the political context

that this research was done in.

And this was that first group,

the senior author of it,

a man named Dick Schwab.

And he got death threats

because of that study reporting,

ooh, here is-- easily

interpreted as-- something

wrong in the brains of gay man.

And Simon LeVay became

the hero in the community.

This was utterly embraced

by the community.

Because in large part,

how it was interpreted as,

this is biology.

This isn't choice.

This is biology.

Look at this.
This is ridiculous,
us saying, oh my god.

If we have gay teachers

in the classroom,

we will turn the Boy Scouts

of America into a gay men.

Oh my god, if we have blue-eyed

teachers in the classroom--

the usual argument that this is

absolute gibberish to demonize

sexual orientation as a choice.

Look at this.

There's a neurobiology of it.

And this was sufficiently that

I've seen people in the Castro

District in San Francisco--

this has disappeared somewhat,

showing the half life of

neuroanatomical knowledge.

But during that time, people

in the Castro District up

in the city there, which

is a very gay community,

I have seen people with

t-shirts saying-- the other term

people never really wanted

to embrace this INAH,

so its nickname was the

sexually dimorphic nucleus

of the hypothalamus.

And I have seen people

with t-shirts saying,

the only small thing about me is

my sexually dimorphic nucleus.

They were actually

selling t-shirts

that would say this during

gay pride parades around 1990

or so.

Isn't it great when

people learn neuroanatomy

out in the general public there?

OK.

So an interesting
brain difference

confounded by people
who died of AIDS.

It's still not clear

what that means

in terms of possibly
negating the finding at least

independently replicated.

Fascinating piece
of not science,

but the political

context of science.

This politically incorrect

to an extreme, this very,

very widely embraced.

One interesting thing is

that paper by LeVay of

was published in
the journal Science.

Again, constantly mentioned as

probably the most influential

science journal in this country.

And it was published--

what year was it? '88.

'88, '92, '90?

OK.

I'm getting the year wrong.

But it was just before the

Clinton election against Bush
where one of the big issues
was gays in the military.

That was the first thing

that Clinton turned to

after he was elected.

I happen to know the man

who is the editor of Science

at the time.

And they timed the publication.

That paper came out in

late October of that year.

And they held it for

that time because they

knew that this was going to be

an issue in the election, which

was kind of cool that they did

that, although some people may

disagree.

OK.

But we hurtle on.

So what other biological,

neurobiological differences,

as a function of
sexual orientation?

Another one that comes through

over and over and over again,

which-- what do
you make of this--

is apparently there is a
reliable gender difference

in the length of
the second finger

versus the fourth finger,

the ratio of the two.

And just to show how bi--

whoa, did a lot of hands

just go up in this auditorium.

And just to show how
biologically compelling

the explanation is for

it, I don't actually

remember which like-- who's

got the greater four to two

ratio, which sex, or whatever.

But it is a very--
now people are

checking each other's hands.

OK.

The first wave of-- and now

all this chimp hand inspection

stuff happening in here.

And what has been shown

quite reliably since then is,

on the average, gay men tend

to have the finger length

ratio of straight women

rather than of straight men.

A small effect there.

Another even more

bizarre finding,

which is there is something

called the autoacoustic reflex.

And what that is is if you sit

there and plug your ears up

with your fingers, you will

hear a noise that is just

coming from the intrinsic

vibration of something

right there in your

ears, and that's

the autoacoustic reflex

generating some low Hertz

sound in there.

And the rate of the oscillation

differs by sex in humans,
which no doubt
explains everything

about the tragic

wars of the sexes

and why people just don't

understand each other

by gender because of their ears

vibrating at different speeds.

But what these studies

have also shown

was gay men having the

autoacoustic reflexive

vibratory speed more

typical of straight women

than straight men.

Again, a very small effect.

What are all of these about?

The assumptions
are it's got to do

with something with prenatal

hormone environment.

Stay tuned.

We will be coming back to this.

Now somewhere in
there you may ask, OK,

well what about the neurobiology

of sexual orientation in women?

Vastly smaller literature.

Far, far less studied.

What has been shown so far

are only two endpoints.

One is the same deal with the

fourth to second finger ratio.

On the average, gay women

have the ratio more typical

of straight men
than straight women.
The other thing
that's been shown

is the same autoacoustic

reflex thingy going on there.

Final realm of
neurobiology, rather than

issues of gay versus

straight, what

is the neurobiology
of transsexuality?

And that used to be

considered to be purely

a domain of psychopathology.

If being gay used to be

a certifiable psychiatric

disorder-- up until the early

1970s, the American Psychiatric

Association in their textbook,

the Diagnostic Statistical

Manual, you could be

psychiatrically certified

as ill.

A psychiatric disorder was

being homosexual or lesbian.

And then in what had to

have been one of the more

all-time blow out

committee meetings ever,

they decided that,

no, actually it's

not a psychiatric disorder.

And overnight, about

40 million Americans

were cured of a
psychiatric disease.

The notion of transsexuality

as a psychiatric disorder

has had much, much

longer shelf life.

What's the neurobiology of that?

To date, there have been

a handful of studies,

and they show essentially

the same thing-- really,

really interesting.

Another region of
the brain that shows

a sex difference in
its average size--

don't even worry about

the name of this.

It's called the bed nucleus

of the stria terminalis.

It's where the amygdala

begins to send its projection

into the hypothalamus.

Another one to those

gender differences.

There is one type

of neuron in there

with a certain type of

neurotransmitter, where very,

very reliably it is about

twice the size in males

than in females.

Sufficiently so that
even in human brains,

you could pretty confidently

determine the sex of somebody

by seeing the number

of these neurons.

You'll see, I'm not

even saying the name

of the neurotransmitter.

It's irrelevant.
It's just another one
of those differences,

a dimorphism in a region of
the brain, a really, really

reliable one.

And this was a study done by

some superb neuroanatomists

looking at transsexuals.

And what they showed

was very interesting,

which was very, very reliably

and a very powerful effect.

What you would see

in their large sample

size of transsexuals
brains post-mortem

was people would have this

part of the brain, the size not

of their sex that they were

born with, but rather of the sex

they insisted they

always actually were.

Wow.

Immediate questions
one must ask.

OK.

Well, maybe this

is due to the fact

that when people change

gender, transsexual procedures,

there's a whole lot

of hormones involved.

And maybe that's doing something

to this part of the brain.

Critical control
that they had was

this was looking

both at transsexuals
who had made gender changes and
those who went to their death

bed saying this is

not the sex that I am,

I got the wrong body, but

never made the change.

It wasn't a function
of having actually

gone through the transition

and the endocrine manipulations

with it.

Another control
they had, which was

looking at men who would get

a certain type of testicular

cancer where they

would have to be

treated with certain

feminizing hormones.

In other words, very similar

to some of the endocrine

treatments of male-to-female
transgendered individuals.

And post-mortem, you didn't

see the changes there.

It has nothing to do
with the hormones.

It had to do with the person

insisting from day one

that they got the wrong body.

And this was a landmark

study, fabulously

well done and controlled,

and replicated once

since then showing that

what transsexualism used

to be thought of
is that people who

think that they're a different

gender than they actually are.

What this study suggests is

what transsexualism is about

is people who got the

wrong gendered body.

And these are people who are

chromosomally of one sex.

In terms of their gonads,

they're of that sex.

In terms of their hormones,

they're of that sex.

In terms of their genitalia

and their secondary sexual

characteristics,
they are of that sex.

But they're insisting,

that's not who I really am.

This part of the brain

agrees with them.

Also very interestingly

that study

was done by the same Dutch

scientists who did this one.

Again, this is very

complex terrain

in terms of what these

things wind up implicating.

Interestingly, that study was

published right around the time

that the city of San Francisco

did something very cool, which

was for city employees

now, medical insurance

will cover transgender

operations.

However, there is no evidence

that the obscure endocrine

journal published out

of Latvia or something
did that like the afternoon
before the San Francisco

commissioners had their

meeting on that one.

But nonetheless,
this is a subject

with all sorts of

realms of implications.

One additional study

about transsexualism.

OK.

How many of you know about

Phantom Limb Syndrome?

OK.

You are a guy with a penis,

and you get a certain type

of penile cancer.

And what's often done is

your penis is excised.

It is cut off.

And about 60% of men who have

had to have their penises

removed because of
cancer there wind up

getting phantom penile

sensations, which

I don't want to know about.

What you see though is when you

take transgendered individuals

who go from male to

female, in other words,

as part of it having
their penises removed, 0%

rate of penile
phantom sensation.

Suggestion being that there

is something much more

"normal" in that case than

when a penis is being removed

for cancer, a whole new area

of research, very novel,

very challenging.

OK.

So this has giving us a

sense now of this bucket.

And we are now ready to move

on to, what in the environment

releases some of
these fixed action

patterns of sexual behavior?

What in the environment

is doing this or that

to the medial preoptic

area or the amygdala

or vasopressin receptor
levels or any such thing.

What are the sensory

triggers for the neurobiology

of sexual behavior?

OK.

Right off, what is obvious is

we are in ethologyville here.

It's going to depend on the

species which sensory modality

is most important.

And this is, once again, the

crushing of the limbic system

equals nose-brain concept.

Limbic system equals

nose-brain if you are a rat.

It's, once again, going

to be interviewing

an animal in its own language.

You've got species where

the releasing stimuli
are all visual.

And we've heard one

example of that already,

which were the pathetic male

turkeys getting faked out

by the Styrofoam female turkeys

with the feathers pointing

the wrong way.

Visual stimuli.

Other species are

quite visual as well.

Primates, non-human primates,

monkeys, for example.

And what studies have shown

is-- how's this for remarkable?

OK.

Here we have-- nah, forget it.

OK.

You will take a rhesus

monkey, a rhesus monkey

from a social group.

And he's sitting there,

and he can lever press

for various rewards.

And he will press a lever

a certain number of times

to get some juice as a reward.

He will press a lever a

certain number of times

to see a high ranking male

from his social group,

no doubt to keep
an eye on the guy.

He will not lever press

to see a male who is lower

ranking than him, but

he will lever the most

to see pictures of female

rhesus monkeys who are in heat.

Whoa!

Is that weird or what?

And the bigger the estrus

swelling on the female,

the more levered

pressing the male

will do to be able to see this.

So that's kind of interesting.

And this close relative of ours,

in terms of visual stimuli.

What is also known

is humans are highly

visual in their sexual

responsiveness as well.

Visual stimuli as
releasing stimuli.

What is no surprise
whatsoever is on the average,

males are more responsive

to visual releasing stimuli

than are females among humans.

And this has been

shown in various ways.

For example, now studies using

brain imaging showing that

for visually sexually arousing

material that not only

are men on the average

subjectively more responsive,

but you get more

of an activation

of the dopamine pathways

in men than in women.

What's interesting also

is that in men, you
uniquely get activation of that
area of the amygdala as well.

And again, that weird world

of the structure of the brain

heavily involved in aggression

also being involved something

about male sexual motivation.

What else?

Then there, of course, is the

world of tactile stimulation.

And what you've got is a whole

domain where, not surprisingly,

stimulate the right

tactile receptors,

and it is sexually arousing.

Are they sure?

Have they done enough

research on this?

And you will activate

dopamine regions.

All of that making

perfect sense.

What also makes perfect sense is

some types of tactile receptors

in some part of the body

activate dopamine more

than other types.

We are now in the whole

world of erogenous zones

and that whole deal.

What is also clear is

that these receptors,

these tactile receptors, their

responsiveness to stimuli

will change depending

on your hormone levels.

And what you see is in women,

tactile responsiveness,

the extent to which

tactile stimulation of skin

throughout the body, but

especially of the genitals,

tactile stimulation evokes

more dopamine activation

when somebody is ovulating.

In other words, at
ovulation, women's skin

is more sensitive to
sexually arousing touch.

In men, it requires
testosterone.

Men who are castrated, tactile

responsiveness to stimuli

goes down in terms of

finding them pleasurable,

sexually arousing.

Final domain of tactile

stuff, the specialized version

we've heard of already, that

lordosis reflex business.

Again, that's a spinal

reflex, but this is not

a spinal reflex of bopping

somebody on the knee

and their leg goes flying out.

This is spinal
reflex where you only

get the lordotic arch backing

in females-- arch backing?

Back arching.

OK.

You don't get either,

but you especially

don't get the arch

backing when you don't
have elevated estrogen levels.

Only when females are ovulating

are those tactile receptors

sensitive to pressure on
the flanks of the rear end,

and out comes the reflex there.

So tactile stimulation.

At the end of the day,

though, without question,

as agreed upon by every

scientist on earth,

the coolest sensory modality

for sexual release stimuli

are olfactory cues, pheromones.

And thus, we enter the

magnificent wonderful world

of pheromonal communication
and pheromonal sexual arousal.

All sorts of interesting

findings there.

First, at the end of

generating pheromones

that are sexually arousing.

What is required in both

species-- in both species?

Whoa.

That was an interesting slip.

What's required in
both sexes-- OK.

What's required in both kinds

is the right hormones into order

to generate pheromones.

Sexually arousing pheromones

in all the different species

look-- that's where

the species part was
coming into that sentence.

What you see is males do not

generate sexually arousing

pheromones if they lack

testosterone levels.

Ovariectomized females,
women, rats, monkeys,

et cetera, who have had

their ovaries removed

do not produce pheromones

that are sexually arousing.

What that of course

brings up is,

what are some of the chemical

constituents of pheromones?

And this is very interesting

because it brings up

another one of those weird

domains of neuromarketing,

pheromones that have

sexually arousing components.

There's all sorts of fatty

acids that play a role in that.

But a lot of what is sexually

arousing about pheromones

in different species
are breakdown products

of sex hormones.

Breakdown products of
androgens in males,

of estrogens in females.

And that winds of providing

some of these sexually

arousing aspects of those odors.

What does that

immediately tell you?

Your olfactory receptors have

all sorts of receptors there
that could pick up on remnants
of testosterone and estrogen,

things of that sort.

That makes a lot of sense.

What doesn't make

any sense at all

is the following finding,

which is perfume.

Perfume, in its classic

form, is made out

of the sweat of various animals.

OK.

Except we're going to get into

even worse domain here, which

is perfumes
traditionally, before

getting the synthetic

versions, were typically

made from the sweat

of male animals.

Hm.

What's that about?

Musk.

Things of that sort.

Chanel No.

5 is made from the sweat of

whipped male Abyssinian cats.

I kid you not.

And this even produced

protests some years ago,

animal rights groups,

about how we should not

be perfuming ourselves
with the sweat

of whipped male Abyssinian cats.

Suddenly, you've
got a real puzzle.

Along comes synthetic perfumes,

and the majority of them

are made of synthetic

versions of androgens.

Wait a second.

Perfume is made up of all

sorts of breakdown products

of male sweat.

Isn't perfume supposed to

like smell good to guys?

We have a deep
abiding puzzle here.

And there is an answer for it.

Let me survey people first off.

OK.

Guys in the room,

how many of you

basically think that

most perfumes smell

kind of appealing?

OK.

How many of you don't?

OK.

Females, how many of you think

your basic off-the-rack perfume

smells appealing?

OK.

Well, that proves something.

OK.

Just complete it, how

many of you don't?
OK.

The vast majority of perfumes

are not purchased by men.

The vast majority are

purchased by women.

In other words, most of

the marketing decisions

about what to stick

in your perfumes

are being marketed

for people who

are going to decide if

it's appealing or not,

people who have lots of

estrogen in their bloodstream

rather than androgens.

That is thought to be
the explanation for how

it is that most perfumes are

derived from male pheromones.

Yeah.

The other thing is if

I were to wear perfume

it would more so that girls

think I smell good instead

of for me to think I smell god.

OK.

Yes.

The strategizing starts.

OK.

So a whole new world of

potential neuromarketing here.

But this taps into, what are

the chemical constituents

of pheromones?
What sort of
information is carried

by pheromones, olfactory
communication, between genders?

It will tell you the

species of the individual.

It will tell you their gender.

It will tell you whether

they are gonadally intact,

whether they've been

castrated or not.

It will tell you something

about their health.

It will tell you whether

they are terrified or not.

Have the sweat from someone or

something that is terrified,

and as we already know,

it will smell differently

to the amygdala.

It will have a lot more

glucocorticoid breakdown

products in it.

And it will tell you,

as we know already,

if it's the right species, if

this person is related to you.

Final point before we then

go into the specifics of what

pheromones are doing to the

neurobiology of all this,

finally, not only do you need

to have hormones intact in order

to generate the
pheromones, you need

to have the right

reproductive hormones on board

in order to perceive them.

Men who are castrated
no longer find

the smells of
female ovarian-- OK,

I'm getting ahead

of myself here.

OK.

If you don't have the

hormones on board,

if you have no estrogen

and you're female,

or if you have no
testosterone and you are male,

you will not be

able to distinguish

the sweat of women and men.

Gonadally intact people can

at above the chance level.

Finally, women become

far better at detecting

the smell of-- distinguishing

the smell of men versus women

when they are ovulating.

Finally, finally,
that's not what you see

when you have gay individuals.

Gay men are better at

detecting the smell of gay men

than either straight

men or straight women.

So we've gotten the

first pieces of here

with the pheromonal system.

You have to be hormonally

intact to generate

pheromones that are sexually

informative and to detect them.

What we will then transition

to is, what sort of information

is being carried
in the pheromones

and what effect does it

have on the neurobiology

of depending on who's
pheromones you are sniffing?

OK.

So we will--

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