LIBERATING THE FEMALE RAT
Beach was a minority voice in a sexually conservative era, but increasingly, scientists could not avoid hearing the political and social arguments voiced by people such as Betty Friedan, whose 1963 best-selling book The Feminine Mystique exploded the suburban family idyll. As Friedan founded the National Organization for Women (in 1966), other movements for social change— civil rights, the antiwar movement, and, with the Stonewall Riot of 1969, the gay liberation movement—gained national visibility.122 By the time of the publication of Money and Ehrhardt’s Man and Woman, Boy and Girl, in 1972, a groundbreaking work about the biology of sexual development, the women’s liberation movement was clearly a force to be dealt with. Money and Ehrhardt figured that they would satisfy no one. ‘‘The advocates of male supremacy like to quote the findings of Chapter 6’’ that claimed that fetal hormones affect brain development, ‘‘while neglecting the findings of Chapters 7 and 8,’’ which discuss the importance of the environment in the formation of gender identity. ‘‘The advocates of women’s liberation, by contrast, attend chiefly to Chapters 7 and 8 and neglect Chapter 6.’’123
And in 1974 the psychologist Richard Doty published an article entitled ‘‘A Cry for the Liberation of the Female Rodent: Courtship and Copulation in Rodentia’’ (see figure 8.3). Doty, who had done his postdoctoral studies under Frank Beach’s supervision, noted that females per se were understudied. During the 1960s, only 20 percent of the articles on rat copulation published in the Journal of Comparative and Physiological Psychology focused on females. Another 68 percent studied males alone, while 12 percent examined both.124 Doty also critiqued the standard system for measuring sexual behavior in the laboratory, a critique made consequential because such measurements lay at the heart of experiments supporting the O/A theory.125
In developing the best ways to observe male sexual behavior, most scientists tried to keep the behavior of the test female constant. They usually put male rats in a small observation cage and permitted them time to sniff it out and become accustomed to their surroundings. Once the male rat was comfy, the scientist introduced the female. The male rat might mount her a few times while she arched her back in the lordosis posture, permitting intromission and ejaculation. Experienced males come to know the procedure very well, becoming so excited, one rat runner wrote, ‘‘that when eventually the female
FIGURE 8.3 : Liberating the female rat. (Source: Alyce Santoro, for the author)
is introduced, the male will no longer inspect whether or not she is in es – trous,’’ but just attempts to mount.126 Even today, most researchers try to minimize female variability, often using circular test chambers so that the females have no corners to back into to prevent mounting. Hormone studies usually employ sexually experienced males, because in inexperienced ones the precopulatory behaviors, including mounting, depend on the female’s soliciting behavior.127 (I can’t stop myself from thinking of this observation as the rodent variation on the tradition of having older women initiate young men into sexual adulthood.)
In fact, when experimenters introduced female choice, funny things began to happen. Doty noted experiments in which females had to want to mate—a desire expressed by pressing a bar to gain access to a stud male. In such situations females paced their sexual contacts (and thus those of the males) in a manner that perhaps more accurately reflected behavior in the wild. Varying the test situation also affected the results of experiments with pre – or perinatal hormone exposure. The psychologist Roger Gorski described experiments in which he first allowed a female rat that had been treated perinatally with androgens to become accustomed to her test area. According to the O/A theory, prenatal androgen treatment ought to have suppressed female lordosis—the measurement of her femininity. Indeed, this is what happened when the androgen-treated female was simply dropped into a test arena containing a waiting male. But when Gorski introduced the test male only after letting the female check out her new cage for a couple of hours, he found that “most females exhibited a very high LQ’’ (Lordosis Quotient, a standard measure arrived at by taking the number of male mounts that induce lordosis and dividing it by the total number of mounts). The permanent organizing effects of androgen on the female brain seemed to have disappeared.128 His result, Gor – ski noted, showed that the masculinization of androgen-treated females is context-dependent.129
That the effects of prenatal hormone treatment were contingent on the experimental test situation was not the only discovery that challenged the O/A theory during the 1970s. Some researchers, led by Frank Beach, challenged the prevailing model of rodent masculinity and femininity. Beach suggested understanding female heterosexual mating behaviors as consisting of three components: attractivity (how much she attracted males), proceptivity (how attracted the female was to a particular male and whether she solicited mating responses from him), and receptivity (a female’s passive willingness to mate).130 In the standard laboratory setup, experimenters usually measured only the passive, receptive component of female behavior. But some experiments suggested that prenatal hormones could affect receptive behaviors without altering proceptive or attractive ones.131 Thus, Beach argued, any good theory relating hormones to behavior ought to take account ofsuch com – plexity.132
A second, equally important theoretical challenge researchers posed to the O/A theory revolved around an even broader question: What relationship existed between masculinity and femininity? If an animal (or person) were extremely masculine (by whatever measure), did that mean he or she would, by definition, be unfeminine? Or were masculinity and femininity separate entities, able to vary independently of each other? (Remember Beach’s early experiments showing that lordotic males also sired offspring.) How could some individuals be masculine and feminine at the same time?
Young’s 1959 paper had implied that masculinity and femininity were graded, mutually exclusive responses. The more feminine a guinea pig, the less masculine she was. The psychobiologist Richard Whalen, another of Beach’s students, found that for rats, the situation was not so straightforward. Under the right circumstances, he could produce males or females that both mounted and exhibited high frequencies of lordosis. In other words, masculine and feminine were not mutually exclusive responses, but rather had what scientists called an ‘‘orthogonal’’ relationship133 (see figure 8.4). Later, Whalen and Frank Johnson complicated matters by manipulating the hormone doses and times of stimulation in order to show that masculinization itself had at least three independent physiological components.134 Whalen proposed an orthogonal model of rat sexuality in which, he argued, masculinity and femininity varied independently ofeach other. The same animal could be both highly masculine and highly feminine, highly feminine but not at all masculine (or vice versa), or might score low on both scales.
A. Linear Model
B. Orthogonal Model
figure 8 .4: A: A linear model of masculinity and femininity. As an animal becomes more feminine, it must also become less masculine. B: An orthogonal model of masculinity and femininity. The animal in the upper-right-hand corner shows many feminine and many masculine traits. The animal in the lower-left-hand corner shows only a few feminine or a few masculine traits.
(Source: Alyce Santoro, for the author)
While Beach’s and his students’ work echoed feminist insistence that passivity did not define femininity and that masculine and feminine behaviors showed significant overlap, other researchers seemed to be drawing from the same pool of ideas. For example, Whalen published his orthogonal model the same year that the psychologist Sandra Bem popularized the idea of androgyny by designing a scale to measure their independent variations in humans. The fact that neither was aware of the other’s work at the time suggests that the concept of independent masculinity and femininity was ‘‘in the air,’’ although the precise route bringing the idea to Whalen and Bem remains difficult to pin down.135
Following Whalen’s lead, scientists modified their terminology. Defeminization came to mean the suppression of female-typical behavior (such as lordosis) in genetic females, while masculinization applied to the enhancement of male-typical behaviors in genetic females. Parallel terminology applied to genetic males, for which demasculinizing treatments decreased the frequency of male typical behaviors, while feminizing ones increased female-typical behaviors. Using these words had the unexpected effect of ‘‘encouraging] questions about spontaneous bisexuality that might be overlooked with a different theoretical framework.’’136
The climate of the 1970s, with its focus on human androgyny, the clamor of the women’s movement, and the nascent gay rights movement, helped make visible certain problems with the way scientists envisioned the biology of rodent sex. Even at the biochemical level, it turned out, sexual distinctions were far from clear-cut. In fact, during the 1970s biochemists realized that testosterone, that most masculine of molecules, usually exerted its influence on brain development only after it had been transformed (through a biochemical process called aromatization) into estrogen! The scientists who discovered the phenomenon, which again made it difficult to conceptualize these steroid hormones as specific sex hormones, echoed the 1930s reactions to the finding of estrogenic activity in stallion urine: they called aromatization paradoxical or surprising. Nevertheless, attention had returned to the role of estrogen in sexual development.137