Chapter 6: Hormones and Behavior
The discussion below focuses on how sex-specific hormonal differences may lead to differences in brain development and behavior. Although we focus mainly on sex-related differences, it’s important to recognize that gender-related differences can also impact brain development and behavior. (For a more comprehensive review of sex versus gender, see Lips, 2020.)
Hens and roosters are different. Cows and bulls are different. Human males and females are different. Humans, like many animals, are sexually dimorphic (di, “two”; morph, “type”) in the size and shape of their bodies, their physiology, and for our purposes, their behavior (cf. Fausto-Sterling, 2000). The behavior of males and females differs in many ways. Young females generally excel in verbal tasks relative to young males, who are nearly twice as likely as females to suffer from dyslexia (reading difficulties) and stuttering. Young males generally perform better at visuospatial tasks. More than 90% of all anorexia nervosa cases involve young females. Young males are twice as likely to suffer from schizophrenia. Young males are much more physically aggressive and generally engage in more rough-and-tumble play (Berenbaum et al., 2008). Many sex differences, such as the difference in aggressiveness, persist throughout adulthood. For example, many more males than females are serving prison sentences for violent behavior. The hormonal differences between males and females may account for adult sex differences that develop during puberty, but what accounts for behavioral sex differences among children prior to puberty and activation of their gonads? While societal and cultural factors play a role (discussed below), hormones are a major determinant of sex differences. Hormonal secretions from the developing gonads determine whether the individual develops in a male or female manner. The mammalian embryonic testes produce androgens, as well as peptide hormones, that steer the development of the body, central nervous system, and subsequent behavior in a male direction. The embryonic ovaries of mammals are virtually quiescent and do not secrete high concentrations of hormones. In the presence of ovaries or in the complete absence of any gonads, morphological, neural, and later, behavioral development follows a female pathway.
Gonadal steroid hormones have organizational (or programming) effects on the brain and behavior (Phoenix et al., 1959). The organizing effects of steroid hormones are relatively constrained to the early stages of development. An asymmetry exists in the effects of testes and ovaries on the organization of behavior in mammals. Hormone exposure early in life has organizational effects on subsequent rodent behavior; early steroid hormone treatment causes relatively irreversible and permanent masculinization of rodent behavior (mating and aggressiveness). These early hormone effects can be contrasted with the reversible behavioral influences of steroid hormones provided in adulthood, which are called activational effects. The activational effects of hormones on adult behavior are temporary and may wane soon after the hormone is metabolized. Thus, typical male behavior requires exposure to androgens during gestation (in humans) or immediately after birth (in rodents) to somewhat masculinize the brain and also requires androgens during or after puberty to activate these neural circuits. Typical female behavior requires a lack of exposure to androgens early in life, which leads to feminization of the brain and also requires estrogens to activate these neural circuits in adulthood. But this simple dichotomy, which works well with animals with very distinct sexual dimorphism in behavior, has many caveats when applied to people.
If you walk through any major toy store, you will likely observe some aisles filled with pink-packaged toys and adjacent aisles without any pink packages. Remarkably, you will also see a strong self-segregation of girls and boys in these aisles. Toy manufacturers are often accused of making toys that are gender biased, but it seems more likely that boys and girls enjoy playing with specific types and colors of toys. Indeed, toy manufacturers would immediately increase their sales if they could sell toys to both sexes. Boys generally prefer toys such as trucks and balls, and girls generally prefer toys such as dolls. Although it is doubtful that there are genes that encode preferences for toy cars and trucks on the Y chromosome, it is possible that hormones might shape the development of a child’s brain to prefer certain types of toys or styles of play behavior. It is reasonable to believe that children learn which types of toys and which styles of play are appropriate to their gender. How can we separate the contribution of physiological mechanisms from learning in order to understand sex differences in human behaviors?
To untangle the contributions of physiological mechanisms from learning in sex-specific behaviors, animal models are often used. Unlike in humans, where sex differences are usually only a matter of degree (often slight), in some animals, members of only one sex may display a particular behavior. As noted, often only male songbirds sing. Studies of such strongly sex-biased behaviors are particularly valuable for understanding the interaction among behavior, hormones, and the nervous system.
A study of vervet monkeys calls into question the primacy of learning in the establishment of toy preferences (Alexander & Hines, 2002). Female vervet monkeys preferred girl-typical toys, such as dolls or cooking pots, whereas male vervet monkeys preferred boy-typical toys, such as cars or balls. There were no sex differences in preference for gender-neutral toys, such as picture books or stuffed animals. Presumably, monkeys have no prior concept of “boy” or “girl” toys. Young rhesus monkeys also show similar toy preferences.
What, then, underlies the sex difference in toy preference? It is possible that certain attributes of toys (or objects) appeal to either boys or girls. Toys that appeal to boys or male monkeys, in this case, a ball or toy car, are objects that can be moved actively through space and can be incorporated into active, rough-and-tumble play. The appeal of toys that girls or female vervet monkeys prefer appears to be based on color. Pink and red (the colors of the doll and pot) may provoke attention in infants.
Society may reinforce such stereotypical responses to gender-typical toys. The sex differences in toy preferences emerge by 12 or 24 months of age and seem fixed by 36 months of age, but are sex differences in toy preference present during the first year of life? It is difficult to ask preverbal infants what they prefer, but in studies where the investigators examined the amount of time that babies looked at different toys, eye-tracking data indicate that infants as young as 3 months showed sex differences in toy preferences: girls preferred dolls, whereas boys preferred trucks. Another result that suggests, but does not prove, that hormones are involved in toy preferences is the observation that girls diagnosed with congenital adrenal hyperplasia (CAH), whose adrenal glands produce varying amounts of androgens early in life, played with masculine toys more often than girls without CAH. Further, a dose-response relationship between the extent of the disorder (i.e., degree of fetal androgen exposure) and the degree of masculinization of play behavior was observed. Are the sex differences in toy preferences or play activity, for example, the inevitable consequences of the differential endocrine environments of boys and girls, or are these differences imposed by cultural practices and beliefs? Are these differences the result of receiving gender-specific toys from an early age, or are these differences some combination of endocrine and cultural factors? Again, these are difficult questions to unravel in people.
Even when behavioral sex differences appear early in development, there seems to be some question regarding the influence of societal expectations. One example is the pattern of human play behavior during which males are more physical; this pattern is seen in a number of other species, including nonhuman primates, rats, and dogs. Is the difference in the frequency of rough-and-tumble play between boys and girls due to biological factors associated with being male or female, or is it due to cultural expectations and learning? If there is a combination of biological and cultural influences mediating the frequency of rough-and-tumble play, then what proportion of the variation between the sexes is due to biological factors, and what proportion is due to social influences? Importantly, is it appropriate to talk about “normal” sex differences when these traits virtually always arrange themselves along a continuum rather than in discrete categories?
Sex differences are common in humans and in nonhuman animals. Because males and females differ in the ratio of androgenic and estrogenic steroid hormone concentrations, behavioral endocrinologists have been particularly interested in the extent to which behavioral sex differences are mediated by hormones. The process of becoming female or male is called sexual differentiation. The primary step in sexual differentiation occurs at fertilization. In mammals, the ovum (which always contains an X chromosome) can be fertilized by a sperm bearing either a Y or an X chromosome; this process is called sex determination. The chromosomal sex of homogametic mammals (XX) is female; the chromosomal sex of heterogametic mammals (XY) is male. Chromosomal sex determines gonadal sex. Virtually all subsequent sexual differentiation is typically the result of differential exposure to gonadal steroid hormones. Thus, gonadal sex determines hormonal sex, which regulates morphological sex. Morphological differences in the central nervous system, as well as in some effector organs, such as muscles, lead to behavioral sex differences. The process of sexual differentiation is complicated, and the potential for errors is present. Perinatal exposure to androgens is the most common cause of anomalous sexual differentiation among females. The source of androgen may be internal (e.g., secreted by the adrenal glands) or external (e.g., exposure to environmental estrogens). Turner syndrome results when the second X chromosome is missing or damaged; these individuals possess dysgenic ovaries and are not exposed to steroid hormones until puberty. Interestingly, women with Turner syndrome often have impaired spatial memory.
Female mammals are considered the “neutral” or “default” sex, as additional physiological steps are required for male differentiation; more steps bring more possibilities for errors in differentiation. Some examples of male anomalous sexual differentiation include 5α-reductase deficiency (in which XY individuals are born with ambiguous genitalia because of a lack of dihydrotestosterone and are reared as females, but masculinization occurs during puberty) and androgen insensitivity syndrome or TFM (in which XY individuals lack receptors for androgens and develop as females). By studying individuals who do not neatly fall into the dichotic boxes of female or male and for whom the process of sexual differentiation is atypical, behavioral endocrinologists glean hints about the process of typical sexual differentiation.
We may ultimately want to know how hormones mediate sex differences in the human brain and behavior (to the extent to which these differences occur). To understand the mechanisms underlying sex differences in the brain and behavior, we return to the birdsong example. Birds provide the best evidence that behavioral sex differences are the result of hormonally induced structural changes in the brain (Goodson et al., 2005). In contrast to mammals, in which structural differences in neural tissues have not been directly linked to behavior, structural differences in avian brains have been directly linked to a sexual behavior—birdsong.
Sex differences in human brain size have been reported for years. More recently, sex differences in specific brain structures have been discovered (Figure 5). Sex differences in a number of cognitive functions have also been reported. Females are generally more sensitive to auditory information, whereas males are more sensitive to visual information. Females are also typically more sensitive than males to taste and olfactory input. Women display less lateralization of cognitive functions than men. On average, females generally excel in verbal, perceptual, and fine motor skills, whereas males outperform females on quantitative and visuospatial tasks, including map reading and direction finding. Although reliable sex differences can be documented, these differences in ability are slight. It is important to note that there is more variation within each sex than between the sexes for most cognitive abilities (Figure 6).
- Rooster and hen © Photo by Egor Myznik on Unsplash is licensed under a CC0 (Creative Commons Zero) license
- Girl on chair © Noba is licensed under a Public Domain license
- dimorphic nuclei © Noba is licensed under a CC BY-NC-SA (Attribution NonCommercial ShareAlike) license
- average sex difference © Noba is licensed under a CC BY-NC-SA (Attribution NonCommercial ShareAlike) license
- According to the World Health Organization, sex refers to “the different biological and physiological characteristics of females, males and intersex persons, such as chromosomes, hormones and reproductive organs.” Gender refers to “the characteristics of women, men, girls and boys that are socially constructed. This includes norms, behaviors and roles associated with being a woman, man, girl or boy. Gender varies from society to society and can change over time.” ↵
The induction of male traits.
The induction of female traits.
The process by which individuals develop the characteristics associated with being male or female. Differential exposure to gonadal steroids during early development causes sexual differentiation of several structures including the brain.
The point at which an individual begins to develop as either a male or a female. In animals that have sex chromosomes, this occurs at fertilization. Females are XX and males are XY. All eggs bear X chromosomes, whereas sperm can either bear X or Y chromosomes. Thus, it is the males that determine the sex of the offspring.
The sex of an individual as determined by the sex chromosomes (typically XX or XY) received at the time of fertilization.
The sex of an individual as determined by the possession of either ovaries or testes. Females have ovaries, whereas males have testes.
An enzyme required to convert testosterone to 5α-dihydrotestosterone.