6.1: Introduction

This chapter describes the relationship between hormones and behavior. Many readers are likely already familiar with the general idea that hormones can affect behavior. Students are generally familiar with the idea that sex-hormone concentrations increase in the blood during puberty and decrease as we age, especially after about 50 years of age. Sexual behavior shows a similar pattern. Most people also know about the relationship between aggression and anabolic steroid hormones, and they know that administration of artificial steroid hormones sometimes results in uncontrollable, violent behavior called “roid rage.” Many different hormones can influence several types of behavior, but for the purpose of this chapter, we will restrict our discussion to just a few examples of hormones and behaviors. Behavioral endocrinologists are interested in how the general physiological effects of hormones alter the development and expression of behavior and how behavior may influence the effects of hormones.

To understand the hormone-behavior relationship, it is important to describe hormones. Hormones are organic chemical messengers produced and released by specialized glands called endocrine glands. Hormones are released from these glands into the blood, where they may travel to act on target structures at some distance from their origin. Hormones are similar in function to neurotransmitters, the chemicals used by the nervous system in coordinating animals’ activities. However, hormones can operate over a greater distance and over a much longer time than neurotransmitters (Focus Topic 1). Examples of hormones that influence behavior include steroid hormones such as testosterone (a common type of androgen), estradiol (a common type of estrogens), progesterone (a common type of progestogen), and cortisol (a common type of glucocorticoid) (see Table 1, A-B). Several types of protein or peptide (small protein) hormones also influence behavior, including oxytocin, vasopressin, prolactin, and leptin.

Hormones coordinate the physiology and behavior of individuals by regulating, integrating, and controlling bodily functions. Over evolutionary time, hormones have often been co-opted by the nervous system to influence behavior to ensure reproductive success. For example, the same hormones, testosterone and estradiol, that cause gamete (egg or sperm) maturation also promote mating behavior. This dual hormonal function ensures that mating behavior occurs when animals have mature gametes available for fertilization. Another example of endocrine regulation of physiological and behavioral function is provided by pregnancy. Estrogens and progesterone concentrations are elevated during pregnancy, and these hormones are often involved in mediating maternal behavior in the mothers.

Not all cells are influenced by each and every hormone. Rather, any given hormone can directly influence only cells that have specific hormone receptors for that particular hormone. Cells that have these specific receptors are called target cells for the hormone. The interaction of a hormone with its receptor begins a series of cellular events that eventually lead to the activation of enzymatic pathways or, alternatively, turn on or turn off gene activation that regulates protein synthesis. The newly synthesized proteins may activate or deactivate other genes, causing yet another cascade of cellular events. Importantly, sufficient numbers of appropriate hormone receptors must be available for a specific hormone to produce any effects. For example, testosterone is important for male sexual behavior. If men have too little testosterone, then sexual motivation may be low, and it can be restored by testosterone treatment. However, if men have normal or even elevated levels of testosterone yet display low sexual drive, then it might be caused by a lack of receptors, so treatment with additional hormones will not be effective.

How might hormones affect behavior? In terms of behavior, one can think of humans and other animals as composed of three interacting components: (1) input systems (sensory systems), (2) integrators (the central nervous system), and (3) output systems or effectors (e.g., muscles). Hormones do not cause behavioral changes. Rather, hormones influence these three systems so that specific stimuli are more likely to elicit certain responses in the appropriate behavioral or social context. In other words, hormones change the probability that a particular behavior will be emitted in the appropriate situation (Nelson, 2011). This is a critical distinction that can affect how we think of hormone-behavior relationships.

We can apply this three-component behavioral scheme to a simple behavior, singing in zebra finches. Only male zebra finches sing, and they sing to attract mates or ward off potential competitors from their territories. If the testes of adult male finches are removed, then the birds reduce singing, but castrated finches resume singing if the testes are reimplanted or if they are treated with either testosterone or estradiol. Although people often consider androgens to be “male” hormones and estrogens to be “female” hormones, testosterone is commonly converted to estradiol (Figure 1). Thus, many male-like behaviors are associated with the actions of estrogens! Indeed, all estrogens must first be converted from androgens because of the typical biochemical synthesis process. If the converting enzyme is low or missing, then it is possible for females to produce excessive androgens and subsequently develop associated male traits. Again, singing behavior is most frequent when blood testosterone or estrogen concentrations are high.

Estrogens are somehow involved in singing, but how might the three-component framework help us formulate hypotheses to explore the role of estrogen in this behavior? Estrogens could affect birdsong by influencing the sensory capabilities, central processing system, or effector organs of a bird. By examining input systems, we could determine whether estrogens alter the birds’ sensory capabilities, making the environmental cues that normally elicit singing (like females or competitors) more salient. Estrogens also could influence the central nervous system, for example, by altering neuronal architecture or the speed of neural processing. Finally, the effector organs, like the muscles in a songbird’s vocal apparatus, could be affected by the presence of estrogens. We do not understand completely how estrogens, derived from testosterone, influence birdsong, but in most cases, hormones can be considered to affect behavior by influencing one, two, or all three of these components, and this three-part framework can aid in the design of hypotheses and experiments to explore these issues.

How might behaviors affect hormones? The birdsong example demonstrates how hormones can affect behavior, but as noted, the reciprocal relation also occurs—behavior can affect hormone concentrations. For example, the sight of a territorial intruder may elevate testosterone concentrations in resident male birds and thereby stimulate singing or fighting behavior. Similarly, male mice or rhesus monkeys that lose a fight decrease circulating testosterone concentrations for several days or even weeks afterward. Comparable results have also been reported in humans. Testosterone concentrations are affected not only in humans involved in physical combat but also in those involved in simulated battles. For example, testosterone concentrations were elevated in winners and reduced in losers of regional chess tournaments.

People do not have to be directly involved in a contest to have their hormones affected by the outcome of the contest. Male fans of the Brazilian and Italian soccer teams were recruited to provide saliva samples to be assayed for testosterone before and after the final game of the World Cup soccer match in 1994. Brazil and Italy were tied, but Brazil won on a penalty kick at the last moment. The Brazilian fans were elated, and the Italian fans were crestfallen. When the samples were assayed, 11 of 12 Brazilian fans who were sampled had increased testosterone concentrations, and 9 of 9 Italian fans had decreased testosterone concentrations, compared with pre-game baseline values (Dabbs, 2000).

In some cases, hormones can be affected by anticipation of behavior. For example, testosterone concentrations also influence sexual motivation and behavior in women. In one study, the interaction between sexual intercourse and testosterone was compared with other activities (cuddling or exercise) in women (van Anders et al., 2007). On three separate occasions, women provided a saliva sample from pre-activity, post-activity, and the next morning, analyses showed that the women’s testosterone was elevated prior to intercourse as compared to other times. Thus, an anticipatory relationship exists between sexual behavior and testosterone. Testosterone values were higher post-intercourse compared to exercise, suggesting that engaging in sexual behavior may also influence hormone concentrations in women.