S. D. V. Stephens

notes / Basic Neuroscience and Geometry of Neuroscience / chapter 2: Brain and Behavior

5 Social and Reproductive Behavior

This lecture has three aims: to understand the role of key hormones and neurotransmitters in social behaviors; to explain differences in these findings between species; and to describe how changes in hormonal systems lead to phenotypic changes (changes in an animal’s observable traits, including its behavior). It opened with a memory test on the Hebbian principles from the previous lecture (see Check yourself).

Estrogen and testosterone

Cognition

  • Estrogen is essential for memory formation and maintenance. A lack of activation of estrogen receptors leads to cell death in the hippocampus.
  • Testosterone deprivation leads to loss of synapses in the hippocampus, and to an increase in amyloid plaques (clumps of protein that build up between neurons, as in Alzheimer’s disease).

The slide’s figure shows stained sections of mouse hippocampus from wild-type (“wt”) and mutant (“reln”) mice at 3 and 9 months of age, with arrows marking small clusters of dark deposits.

Sex steroids and sexual behavior

  • Estrogen is required for sexual behavior.
  • Testosterone amplifies sexual behavior.

The slide shows a pair of sandhill cranes in a courtship dance.

Aggressive behavior

Both testosterone and estrogen are associated with aggression:

  • estrogen is necessary for aggression;
  • testosterone can amplify estrogen-dependent aggression.

“Aggression” can show up as any form of competition. (The slide shows a male lion roaring, with cubs at his side.)

Seasonal changes

  • Testosterone increases during mating seasons.
  • Testosterone increases last longer in polygynous species (where one male mates with several females) than in monogamous species (where a male and a female form a mating pair).

The lecture asked why in both cases (see Check yourself). The slide shows a male satin bowerbird decorating his bower with blue objects while a female looks on.

The winner effect

Definition 5.1 (The winner effect). In the winner effect, winning an aggressive encounter increases the number of testosterone and estrogen receptors, and winning fights increases the likelihood of winning future aggressive encounters. The loser effect is the opposite: losing decreases sex steroid receptors and decreases the likelihood of winning in the future.

  • The increase in receptors is likely increasing the capacity to respond to testosterone and estrogen during aggression.
  • Winning increases resilience against traumatic stressors.

The slide features Dr. Temitayo O. Oyegbile, with a photo of mice. The next slide is a photo of a football coach shouting on the sideline: competition in human form.

Parental care

  • Estrogen regulates the formation and maintenance of parental behavior.
  • Testosterone decreases parental behavior.
  • Monogamous species have an increase in testosterone around the time of birth. (Why? See Check yourself.)

The slide shows a cat nursing her kittens. A discussion slide followed: generally, monogamous species have an increase in testosterone after their offspring is born, but in humans we see the opposite, where males have a decrease in testosterone after a child is born. Why might this be? (See Check yourself.)

The hypothalamus–pituitary–gonadal (HPG) axis

The hypothalamus–pituitary–gonadal axis: GnRH from the hypothalamus, LH and FSH from the pituitary, testosterone from the gonads, aromatase converting testosterone to estrogen, and negative feedback from both hormones to the pituitary and hypothalamus.

The production of the sex steroids is controlled by a chain of three organs, the hypothalamus–pituitary–gonadal (HPG) axis:

  1. The hypothalamus releases gonadotropin-releasing hormone (GnRH).
  2. GnRH acts on the pituitary, which releases luteinizing hormone (LH) and follicle-stimulating hormone (FSH).
  3. LH and FSH act on the gonads (testis or ovary), which produce testosterone.
  4. Aromatase converts testosterone into estrogen.

Testosterone and estrogen also act back on the hypothalamus and the pituitary (the curved arrows). What happens if the hypothalamus releases too much GnRH? More GnRH means more LH and FSH, and so more testosterone and estrogen. The pathway is self-regulating, though: if too much testosterone or estrogen binds to receptors in the hypothalamus, GnRH release decreases.

Definition 5.2 (Negative feedback). In negative feedback, the output of a pathway acts back on the pathway to reduce its own production. In the HPG axis, testosterone or estrogen binding to receptors in the hypothalamus decreases GnRH release, which brings their levels back down.

The lecture then asked what happens when parts of this loop fail (see Check yourself).

Vasopressin and oxytocin

  • Vasopressin and oxytocin are made by neurons in the hypothalamus (in its paraventricular and supraoptic nuclei), whose axons carry them to the posterior pituitary, from which they are released.
  • They play key roles both in the brain and in the periphery (the rest of the body).
  • They are highly implicated in sociality.

The slide’s figure locates the hypothalamus deep in the brain and shows neurons of its paraventricular nucleus and supraoptic nucleus sending axons down to the posterior pituitary. It also gives the chemical structures of the two hormones: each is a chain of nine amino acids, and they differ at only two positions (isoleucine and leucine in oxytocin, phenylalanine and arginine in vasopressin).

Oxytocin

  • Oxytocin depolarizes the membrane potential in muscle fibers, which allows uterine contractions.
  • There is a large oxytocin release after birth, and after sex. Oxytocin is commonly referred to as the “love” hormone.
  • Oxytocin increases affiliative behavior (behavior that seeks out and keeps closeness with others).
Action potential: from rest at −70 mV past threshold at −55 mV, depolarization as sodium flows in to a peak near +40 mV, repolarization as potassium flows out, hyperpolarization, and return to rest, with small failed initiations below threshold.

To show what “depolarize” means, the slide recalls the action potential: (1) a stimulus arrives at a membrane in its resting state (−70 mV); (2) once the membrane passes threshold (−55 mV) it depolarizes as Na ions flow in, up to a peak near +40 mV; (3) it repolarizes as K ions flow out; (4) it overshoots into hyperpolarization below rest; and (5) it returns to the resting state. Stimuli that do not reach threshold produce only small “failed initiations.”

Vasopressin

  • In the periphery, vasopressin causes water retention.
  • It is associated with aggression and dominance, in particular territorial aggression and mate-guarding. The lecture called vasopressin “applied” sex steroids.
  • It matters for the difference between monogamous and polygynous mating systems (next section).

The slide shows two betta fish (Siamese fighting fish) facing off with their fins flared.

The importance of receptors

  • Monogamous voles have more oxytocin and vasopressin receptors than polygynous voles.
  • Increasing vasopressin receptors induces pair bonding (a lasting preference for one partner) and parental care; blocking vasopressin receptors blocks these behaviors.
  • Monogamous voles have an extended microsatellite (a stretch of DNA made of a short sequence repeated many times) in the vasopressin receptor gene.

As the slide’s title says, receptors are what matter here: the number of receptors for a hormone shapes the social behavior it produces. (The slide shows a pair of voles with their pups, framed by hearts.)

The lecture closed with a reading on vasopressin, oxytocin and PTSD (post-traumatic stress disorder): ScienceDirect article.

Check yourself

Question 5.3 (Memory test). Reflect on the three Hebbian principles of synaptic plasticity. Give a real-world example for each.

Show solution

Use it or lose it: a skill you haven’t practiced in years has partly faded. Use it to improve it: practice (repetition) or an intense stimulus such as a drug strengthens synapses. Regional and functional specificity: learning Latin will not make you good at basketball.

Question 5.4 (Seasonal changes). Why does testosterone increase during mating seasons? And why might testosterone increases last longer in polygynous species than in monogamous ones?

Show solution

Testosterone amplifies sexual behavior and aggression, which shows up as competition for mates, so it rises when mating is at stake. A polygynous male keeps competing for further mates over a longer period, while a monogamous male settles with one partner and moves on to parental care, which testosterone decreases.

Question 5.5 (Discussion). Generally, monogamous species have an increase in testosterone after their offspring is born. In humans, males instead have a decrease in testosterone after a child is born. Why do you think this is?

Show solution

The lecture left this open. One answer consistent with it: testosterone amplifies aggression such as territorial defense and mate-guarding, which may be how fathers of other monogamous species protect their young, while human fathers care for children directly, and testosterone decreases parental behavior.

Question 5.6 (HPG axis). What happens if the testosterone and estrogen receptors in the hypothalamus are impaired?

Show solution

The hypothalamus can no longer sense testosterone and estrogen, so negative feedback fails: GnRH release is not turned down, LH and FSH keep stimulating the gonads, and testosterone and estrogen levels climb too high.

Question 5.7 (HPG axis). What happens if aromatase isn’t converting enough testosterone into estrogen?

Show solution

Estrogen falls while testosterone builds up. This lecture showed estrogen is needed for memory, sexual behavior, aggression and parental behavior, so these suffer, and testosterone cannot make up for it, since it only amplifies estrogen-dependent behavior.

Question 5.8 (Memory check). What best describes testosterone?

  • Increases during mating season
  • Is more associated with aggression than estrogen
  • Increases parental behavior
  • Is made from converted estrogen
Show solution

Increases during mating season. Estrogen is the hormone necessary for aggression (testosterone only amplifies it), testosterone decreases parental behavior, and estrogen is made from testosterone, the other way round.

Question 5.9 (Memory check). What best describes estrogen?

  • Decreases during mating season
  • Is more associated with aggression than testosterone
  • Increases parental behavior
  • Decreases aggression
Show solution

Increases parental behavior: estrogen regulates the formation and maintenance of parental behavior. Both hormones are associated with aggression, and estrogen is necessary for it, so it does not decrease aggression.

Question 5.10 (Memory check). True or false: both testosterone and estrogen are essential for memory.

Show solution

True. Without estrogen-receptor activation, hippocampal cells die; without testosterone, the hippocampus loses synapses and amyloid plaques increase.

Question 5.11 (Memory check). The hypothalamus releases:

  • a. Testosterone and estrogen
  • b. GnRH
  • c. LH
  • d. FSH
Show solution

b. GnRH. LH and FSH come from the pituitary, and testosterone and estrogen from the gonads (estrogen via aromatase).

Question 5.12 (Memory check). Which hormones trigger the creation of testosterone?

  • a. GnRH
  • b. LH
  • c. FSH
  • d. B and C
Show solution

d. B and C. In the lecture’s HPG diagram, LH and FSH from the pituitary act on the gonads, which make testosterone; GnRH acts only on the pituitary. (Strictly, LH is the main trigger of testosterone synthesis; FSH acts mostly on other cells of the gonads.)

Question 5.13 (Memory check). Why don’t LH and FSH create estrogen?

  • Estrogen is created by progesterone
  • Estrogen is testosterone converted by aromatase
  • Estrogen is created by GnRH directly
  • They do, you’re lying, this is a trick question you scoundrel
Show solution

Estrogen is testosterone converted by aromatase. LH and FSH trigger the gonads to make testosterone, and aromatase then turns testosterone into estrogen.

Question 5.14 (Memory check). Which is most important for hormonal regulation of social behavior in mammals?

  • The amount of hormone released
  • The amount of hormone in the periphery
  • The number of receptors
  • The amount of enzymes
Show solution

The number of receptors. Monogamous and polygynous voles differ in their oxytocin and vasopressin receptors, and adding or blocking vasopressin receptors switches pair bonding and parental care on or off; the winner effect also works through receptor number.

Question 5.15 (Memory check). Monogamous voles have ________ compared to polygynous voles.

  • Less vasopressin receptors
  • More vasopressin receptors
  • Less vasopressin hormone levels
  • More vasopressin hormone levels
Show solution

More vasopressin receptors (and more oxytocin receptors).

Question 5.16 (Memory check). Oxytocin is implicated in

  • Parental care
  • Pair bonding
  • Affiliative behavior
  • All of the above
Show solution

All of the above. Oxytocin increases affiliative behavior, is released in large amounts after birth and after sex, and monogamous, pair-bonding voles have more oxytocin receptors.