In Memory of Robert W. Goy.

Robert W. Goy, pioneering investigator of the origins of behavioral sex differences, educator, and Primate Center director, died January 14, 1999, from cardiovascular and metabolic complications. He would have been 75 on January 25th.

Goy was a professor of psychology and director of the Wisconsin Regional Primate Research Center at UW-Madison from 1971 to 1989. His seminal research advanced the notion that exposure to the male sex hormone testosterone during fetal development "organized" the developing nervous system to express masculine characteristics. This basic principle of hormone action has been found to operate in animals from lizards to nonhuman primates, and is an important aspect of human development. In addition, Goy made significant contributions to our understanding of the role that early social experience plays in developing the expression of masculine and feminine behavior. For more than 35 years, Goy mentored Ph.D. students and postdoctoral fellows who have become leaders in the fields of primate behavior and neuroendocrinology. A long time member of the NIH Psychobiology Research Panel, Goy was a strong and consistent supporter of innovative research in this field. Many of today's established researchers benefited from Goy's ability to recognize new and exciting research approaches before they became widely accepted.

Goy was born in Detroit, and received his undergraduate and doctoral degrees in psychology from the University of Michigan in 1947 and the University of Chicago in 1953, respectively. He then joined the laboratory of W. C. Young at the University of Kansas, where some of the most important advances in the emerging field of behavioral endocrinology were made over the next 10 years. In the early 1950s, W. C. Young's laboratory team, through extensive studies of guinea pigs, demonstrated that the presence of specific gonadal hormones turned on, or activated, adult patterns of reproductive behavior. Using inbred guinea pig strains, Goy, along with Jaqueline Jakway, demonstrated that the sensitivity to these activating effects was genetically regulated. Scientists are only now beginning to understand the mechanisms producing this sensitivity.

In 1959 Goy and Young, with colleagues Charles Phoenix and Arnold Gerall, published the first unambiguous evidence that prenatal exposure to elevated levels of the male sex hormone testosterone masculinized both the reproductive anatomy and behavior of genetically female offspring. This landmark study advanced the argument that the fetal hormonal environment permanently organizes the developing nervous system to produce either masculine or feminine patterns of behavior. This organizational effect of hormones became one of the key concepts in behavioral neuroendocrinology and revolutionized the way in which hormonal influences on behavior were subsequently studied. The concept radically altered views of human sexual development when scientists recognized that human genetic anomalies could alter the natural prenatal hormonal environment and permanently alter an individual's anatomy and behavior.

In 1963, Young's laboratory group moved to the newly established Oregon Regional Primate Research Center outside Portland to expand its sexual differentiation studies to nonhuman primates. To prepare for this next research phase, Goy had been a visiting scientist at the Wisconsin Regional Primate Research Center (WRPRC) from 1961 to 1963. At the WRPRC, he studied the measurement of sex differences in juvenile behavior of rhesus monkeys with noted primate psychologist Harry Harlow.

In 1964, Goy, Young, and Phoenix began investigating the effects of prenatal hormone alterations in rhesus monkeys. They produced the first masculinized genetic female rhesus monkey and demonstrated that the principles developed in guinea pigs applied to nonhuman primates and, by extension, to humans. These landmark studies also showed that differences in male and female juvenile rhesus monkeys' social behavior, which occur when the young monkeys are not secreting gonadal hormones, were organized by the prenatal hormone environment. This was the first clear evidence that prenatal hormones actually altered the structure of the nervous system, instead of changing its sensitivity to the activating effects of gonadal hormones. Subsequent work in other laboratories throughout the world have unequivocally provided evidence of specific structural changes within the developing nervous system--changes organized by hormones during the period of sexual differentiation.

Following W. C. Young's death in 1966, Goy headed the Division of Reproductive Physiology and Behavior at the Oregon Regional Primate Research Center. Goy, Phoenix, and colleague John Resko continued their research into elucidating the role of gonadal hormones in the activation and organization of behavioral sex differences.

In 1971, Goy succeeded Harry Harlow as Director of the WRPRC, and continued in this role for 18 years. At Wisconsin, Goy initiated studies on how early experience affects the development of adult reproductive behavior in rhesus monkeys. He was the first to recognize that the standard laboratory rearing paradigm in common use for rhesus monkeys produced appropriate juvenile social behavior but deficient adult sexual behavior, particularly for males. So Goy developed a unique laboratory rearing environment using carefully selected groups of mothers and infants. The environment preserved important aspects of the social environment a rhesus monkey would normally encounter in its natural habitat. With colleagues David Goldfoot and Kim Wallen, Goy demonstrated the important role that early experience plays in the expression of juvenile and adult sex differences in behavior. This research, in addition to continuing studies of the prenatal hormone role in behavioral development, advanced the notion that the prenatal hormonal environment produces behavioral predispositions which are then shaped and molded by early social context. In Goy's view, both biological and social influences were crucial to the development of masculine and feminine patterns of behavior.

During his tenure as director, Goy mentored numerous graduate students, postdoctoral fellows, and visiting scientists in behavioral endocrinology and primate development. Through his teachings Goy combined a caring and thoughtful personal style with a sparkling and masterful intellect. Many of his students have become leaders in the fields of neurobiology, neuroendocrinology, behavioral endocrinology, and primatology.

In 1988 Goy, with colleagues Mary McBrair and Fred Bercovitch, published a study demonstrating that very short prenatal exposure to androgen could masculinize juvenile patterns of behavior. Most importantly, by altering the time during gestation when the female fetus was exposed to androgen, Goy masculinized the female offspring's behavior without masculinizing her reproductive anatomy or neuroendocrine function. This remarkable finding was the first to separate the psychological effects of prenatal hormonal manipulations from their effects on reproductive anatomy. This separation between physical and psychological effects suggests a possible cause of human transsexuality.

In addition to his pioneering contributions to our understanding of sexual differentiation, Goy made equally important contributions to the study of the neural control of sexual behavior. With colleagues Jeff Slimp and Ben Hart he demonstrated that medial preoptic lesions eliminate male rhesus monkey sexual behavior without eliminating sexual motivation. Similarly, with colleagues Ei Terasawa, Stan Wiegand, Thom Nass, Bill Byne, and Ruth Bleir, Goy contributed to our understanding of the organization of the endocrine hypothalamus and its role in regulating the ovarian cycle.

Throughout his career, Goy championed the role that hormones play in activating sex-typical patterns of behavior and how the hormonal environment of an individual is a critical component of one's psychological makeup. In the 1970s, the male response to changes in female physical attractiveness was thought to control rhesus monkey sexuality. Yet during this time, Goy presented the first evidence that the female's ovarian hormones modulated her own sexual motivation, not simply how attractive she was to a male. This view of hormonally modulated female sexual motivation, which took 15 years to demonstrate definitively, solidified our understanding of both human and nonhuman primate sexuality.

Following his retirement in 1989, Goy remained active in behavioral neuroendocrinology. He supported the fledgling Society for Behavioral Neuroendocrinology and collaborated on studies at the Yerkes Regional Primate Research Center that addressed the effect of prenatal suppression of naturally occurring androgens in fetal males. He also collaborated with the WRPRC and the Mayo Clinic on the role of prenatal androgen excess in the development of infertility and diabetes in women. Goy served as a frequent consultant to the NIH and various professional societies. He became the second editor of Hormones and Behavior, following Frank Beach's retirement. He remained editor until the publication became the official journal of the Society for Behavioral Neuroendocrinology in 1996. Goy was an editorial consultant for several scientific journals and books, and he authored or co-authored nearly 200 scientific articles.

Awards Goy received include the Kenneth Craik Award in Physiological Psychology from Cambridge University and the Distinguished Scientific Contribution Award by the American Psychological Association. The latter is awarded to individuals who demonstrate "outstanding theoretical or empirical contributions to basic or applied research in psychology." Last year, Goy was honored for his lifelong contributions at a special symposium of the Inaugural meeting of the Society for Behavioral Neuroendocrinology.

Kim Wallen Emory University and Yerkes Regional Primate Research Center

Address correspondence to Kim Wallen, Ph.D., Emory University, Department of Psychology, Atlanta, GA 30322.