Founders of Behavioral Neuroendocrinology

Behavioral neuroendocrinology has a rich history as a discipline. Given this, it is not surprising that many names come to mind as important contributors. However, four names are likely to come to every SBN member’s mind as being true Founders: Arnold A. Berthold, who conducted the first experiments in behavioral endocrinology during the 1800’s, and three individuals who forged the modern science of behavioral neuroendocrinology – Frank A. Beach, William C. Young, and Daniel S. Lehrman.

Arnold A. Berthold (1803-1861)

Founder Pic Courtesy of the National Library of Medicine.

The first recognized experiments in behavioral endocrinology were conducted by Professor Arnold Adolph Berthold at the University of Göttingen in 1848-1849. Berthold hypothesized that intact testes are necessary for the development of male-typical characters, and he therefore conducted a series of experiments using castration and testes replacement in roosters. He found that males that were castrated as juveniles later showed deficits as adults in behaviors such as aggression, mating and crowing. They also failed to develop the large size and plumage characteristics of normal males. However, Berthold discovered that all of these effects could be reversed if the subject’s testes, or the testes of another male, were implanted into the body cavity. These re-implanted testes became vascularized and produced sperm. Given that their nerve connections had been severed, Berthold correctly concluded that the testes influence behavior not by the actions of nerves, but by secreting a substance into the bloodstream. This conclusion was not widely accepted by his contemporaries, and unfortunately, Berthold never followed up with additional research. He died in 1861, and his findings remained uncited for 50 years. Indeed, the behavioral relevance of gonadal secretions was largely unrecognized until Frank Beach began his classic studies in the 1930’s. Material drawn from “An Introduction to Behavioral Endocrinology,” Second Edition, by Randy J. Nelson (Sinauer: 2000).

Recommended Readings

  • Berthold, A.A., 1849. Transplantation der Hoden. Arch. Anat. Physiol. Wissenschr. Med. pp. 42-46. In German.
  • Berthold, A.A., 1849. Transplantation of testes. English translation by D. P. Quiring, 1944, Bull. Hist. Med. 16, 399-401.

Frank A. Beach (1911-1988)

Founder Pic Contributed by Benjamin D. Sachs, University of Connecticut. Courtesy of the National Academy of Sciences of the United States of America.

Frank Beach was the principal founder of behavioral endocrinology: He named and defined the discipline (Beach, 1975), wrote the first survey (Beach, 1948) and history (Beach, 1981) of the field, organized the annual West Coast sex conference, which eventually morphed into the SBN, and founded the journal Hormones and Behavior. SBN’s Early Career Award is named in Beach’s honor, a tribute both to his scientific accomplishments and to his teaching and mentorship of graduate students and postdoctoral fellows.

Beach, a psychologist, developed an interest in the neural control of instinctive behavior in Karl Lashley’s lab. While an assistant curator of Experimental Biology at the American Museum of Natural History (ANHL), Beach audited an endocrinology course at New York University, for which he synthesized what was then known about the hormonal regulation of behavior. This term paper grew into Hormones and Behavior (Beach, 1948), the discipline’s first book.

Beach became the founding curator of the Department of Animal Behavior at the ANHL, which later included T.C. Schneirla, Lester Aronson, Daniel Lehrman, and Jay Rosenblatt. Beach’s contacts with biologists fostered his appreciation of the natural behavior of animals, and he established contacts with European ethologists, whose naturalistic approach to behavior he valued.

In 1946, Beach accepted a professorship at Yale, and in 1958 he moved to the University of California, Berkeley, where he supervised over 30 PhD’s and postdoctoral fellows, established a field station for behavioral research (Beach, 1969), and spent the rest of his life.

Beach contributed significantly to several areas of behavioral neuroendocrinology, e.g., sexual differentiation and development, interactions between hormones and experience, and differences among species in the relative influence of hormones on behavior. His most influential writings were conceptual and emphasized the value of a comparative approach (Beach, 1967). He stressed the importance of close observation and accurate description of behavior (Beach, 1976). He believed that the concept of instinct needed de-scenting (Beach, 1955), partly because it tended to inhibit, rather than promote, investigation of the ontogeny of behavior. He polemically questioned whether perinatal hormones affect sexual behavior by organizational effects on the nervous system, arguing that it was more parsimonious to conclude that hormonal effects on genital differentiation had secondary effects on the brain and behavior (Beach, 1971).

By force of his personality and example, Beach influenced hundreds of scientists in several disciplines. But he had no illusions of infallibility. Faced with persuasive data, he changed his position on issues, and he often expressed the wish that all science were published in ink that disappeared after 10 years, so that the weight of previous science, and especially theory, would not hinder progress.

Recommended Readings

  • Beach, F.A., 1948. Hormones and Behavior. Paul B. Hoeber, New York.
  • Beach, F.A., 1955. The descent of instinct. Psychol. Rev. 62, 401-410.
  • Beach, F.A., 1967. Cerebral and hormonal control of reflexive mechanisms involved in copulatory behavior. Physiol. Rev. 47, 289-316.
  • Beach, F.A., 1969. Locks and beagles. Am. Psychol. 24, 971-989.
  • Beach, F.A., 1971. Hormonal factors controlling the differentiation, development, and display of copulatory behavior in the ramstergig and related species. In: Tobach, E., Aronson, L.R., and Shaw, E. The Biopsychology of Development. Academic Press, NY, 249-296.
  • Beach, F.A., 1975. Behavioral endocrinology: An emerging discipline. Am. Scientist. 63, 178-187.
  • Beach, F.A., 1976. Sexual attractivity, proceptivity, and receptivity. Horm. Behav. 7, 105-138.
  • Beach, F.A., 1981. Historical origins of modern research on hormones and behavior. Horm. Behav. 15, 325-376.
  • More of the history and flavor of Frank Beach is available from his autobiographies (e.g., 1978, Pioneers in Neuroendocrinology, 2, 19-35. Plenum Press), biographies (e.g., Dewsbury, D.A., 1997, Biog. Memoirs Nat. Acad. Sci. 73, 3-22), and obituaries (e.g., 1988, Horm. Behav. 22, 419-443).

William Caldwell Young (1899-1965)

Founder Pic Contributed by Kim Wallen, Emory University. Courtesy of the National Library of Medicine.

William Caldwell (“WC”) Young, a founder of behavioral neuroendocrinology, had a career spanning almost forty years. Trained with Carl Moore at The University of Chicago, Young was interested in hormonal influences on behavior, but Moore dissuaded him saying: “… the behavior of animals was utterly capricious, unordered by hormonal events, and unrelated to variables of significance to reproductive biology” (Goy, 1967). Fortunately, Young ignored Moore and while at Brown University in 1928 studied hormonal modulation of female sexual receptivity. His studies demonstrated that female guinea pig sexual receptivity varied reliably with cyclic changes in ovarian morphology (Young, Dempsey & Myers, 1935; Myers, Young & Dempsey, 1936). Subsequent classic experiments varied the timing and dosages of estrogen and progesterone given to ovariectomized females, demonstrating that female sexual receptivity required sequential estrogen and progesterone treatment (Dempsey, Hertz, & Young, 1936; Collins, et al. 1938). Between 1939 and 1946, Young worked at Yale Laboratories in Florida and then at Cedar Crest College. He then moved to the Department of Anatomy at the University of Kansas, where he attracted numerous postdoctoral trainees, including Arnie Gerall, Robert W. Goy, Charles H. Phoenix, and Elliot Valenstein. The Kansas laboratory produced ground-breaking studies of guinea pig behavior including genetic influences on estrogen sensitivity (Goy and Young, 1957). Young also demonstrated that the response of males to androgen was determined by individual responsiveness and not androgen amount, a core principle of behavioral endocrinology (Grunt and Young, 1953).

By 1956 Young had been diagnosed with cancer. He set three personal goals before cancer took his life: to finish editing “Sex and Internal Secretions”; to publish 100 research articles; and to make a lasting contribution to the emerging field of Behavioral Endocrinology (Gerall, personal communication). He achieved all three goals, with the last his most important contribution-- the principle that prenatal androgens could organize the nervous system during critical periods of development (Phoenix, Goy Gerall & Young, 1959). The Young group demonstrated that prenatally exposing genetic female guinea pigs to elevated androgens permanently suppressed their capacity to display feminine sexual behavior (defeminization) and significantly enhanced their display of masculine sexual behavior (masculinization). From these striking behavioral results they suggested that the prenatal androgen had permanently altered the tissues underlying sexual behavior and proposed that, similar to the reproductive duct systems, androgens ‘organized’ the developing nervous system during critical period of development. The 1959 study, while controversial, ultimately fixed the ability of hormones to alter neural development as a cornerstone of behavioral neuroendocrinology.

Young moved to the Oregon Regional Primate Research Center in 1963, to continue studies of the organizing actions of androgens using rhesus monkeys. It was here that Young, along with Goy and Phoenix, first demonstrated that prenatal exposure of genetic females to androgens masculinized juvenile behavior and required no hormonal activation, likely reflecting alteration of the nervous system. Young, along with Frank Beach, formed the West Coast Sex meetings, where after Young’s death in1966, the WC Young Award was given each year to the most promising graduate student. Young was self effacing and many would likely be surprised to discover that principles of behavioral endocrinology that they take for granted arose in his laboratory.

Recommended readings:

  • Goy, R.W. Young, W.C., 1967. Anat. Rec. 157, 3-11.
  • Young, W.C., Dempsey, E.W., Myers, H.I., 1935. Cyclic reproductive behavior in the female guinea pig. J. Comp. Psychol. 19, 313-335.
  • Dempsey, E.W., Hertz, R., Young, W.C., 1936. The experimental induction of oestrus (sexual receptivity) in the normal and ovariectomized guinea pig. Am. J. Physiol. 116, 201-209.
  • Myers, H.I., Young, W.C., Dempsey, E.W. 1936. Graafian follicle development throughout the reproductive cycle in the guinea pig, with especial reference to changes during oestrus (sexual receptivity). Anat. Rec. 65, 381-401.
  • Collins, V.J., Boling, J.L., Dempsey, E.W. et al. 1938. Quantitative studies of experimentally induced sexual receptivity in the spayed guinea-pig. Endocrinology 23, 188-196.
  • Feder, H.H., Resko, J.A., Goy, R.W. 1968. Progesterone concentrations in the arterial plasma of guinea-pigs during the oestrous cycle. J. Endocr. 40, 505-513.
  • Goy, R.W., Young, W.C., 1957. Strain differences in the behavioral responses of female guinea pigs to alpha-estradiol benzoate and progesterone. Behaviour, 10, 340-354.
  • Grunt, J.A., Young, W.C., 1953. Consistency of sexual behavior patterns in individual male guinea pigs following castration and androgen therapy. J. Comp. Physiol. Psychol. 46, 138-144.
  • Phoenix, C.H., Goy, R.W., Gerall, A.A. et al. 1959. Organizing action of prenatally administered testosterone propionate on the tissues mediating mating behavior in the female guinea pig. Endocrinology 65, 369-382.
  • Dantchakoff V. 1937. Realisation du sexe a volonte. Bull. Biol. 3, 1-53.

Daniel S. Lehrman (1919-1972)

Founder Pic Contributed by Jay S. Rosenblatt, Rutgers University

As a lifelong naturalist and expert ornithologist, Dan Lehrman was particularly suited to apply Geoffrey Harris’ discovery of the hypothalamic control of pituitary hormone secretions to unraveling the hormonal consequences of male-female behavioral interaction during the breeding cycle in the ring dove. Robert Hinde, his close colleague at Cambridge University, with a similar ornithological background, did corresponding studies on the canary. Hormonal effects on reproductive behavior were already well known, chiefly through the research of Frank Beach, but the discovery that reproductive behavior can in turn affect hormone secretions completed the picture and established the field of behavioral endocrinology.

Dan Lehrman’s research on the reproductive cycle of the ring dove showed how early phases of the cycle arose through behavioral interactions stimulating hormonal secretions which, in turn, stimulated the subsequent phases of behavioral interaction. This approach applied to parental behavior was the basis for Dan’s now classical chapter covering the entire field of avian and mammalian parental behavior for Sex and Internal Secretions, edited by W.C. Young in 1961. In addition to his contributions to behavioral endocrinology, Dan played the principal role in the response of American comparative psychologists to Konrad Lorenz’s theory of instinct in his famous critique published in 1953. He brought to bear on this theory evidence from developmental psychobiology, a broad knowledge of neurophysiology, and the rigor of the scientific method to question both the evidence presented in support of the theory and the philosophical views on which it was based. As a student of T.C. Schneirla, he believed that there were irreconcilable philosophical differences between the behaviorist approach of American comparative psychologists and instinct theory of Lorenzian ethologists that Lehrman described in his last writings. Importantly, Dan had a strong influence on the ethological theory of the other leading ethologist at the time, Niko Tinbergen, who, for the first time, included developmental analysis as one of four areas necessary for the comprehensive study of behavior.

Dan established his laboratory at Rutgers University in Newark in the early 1950’s, which eventually became the Institute of Animal Behavior, the only one of its kind for many years. Many of its graduates are among the leading behavioral endocrinologists today.

Recommended readings

  • Lehrman D.S., 1953. A critique of Konrad Lorenz’s theory of instinctive behavior. Quart. Rev. Biol. 28, 337-363.
  • Lehrman, D.S., 1959. On the origin of the reproductive cycle in doves. Trans. N.Y. Acad. Sci. 21, 682-688.
  • Lehrman, D.S. 1961. Hormonal regulation of parental behavior in birds and infrahuman mammals. In: W.C. Young (Ed.), Sex and Internal Secretions. Williams and Wilkins, Baltimore, pp. 1268-1282.
  • Lehrman, D.S. 1970. Semantic and conceptual issues in the nature-nurture problem. In: L.R. Aronson, E. Tobach, J.S. Rosenblatt, and D.S. Lehrman (Eds.), Development and Evolution of Behavior. pp. 17-52.
  • Lehrman, D.S. 1971. Behavioral science, engineering, and poetry. In: E. Tobach, L.R. Aronson, and E. Shaw (Eds.), Biopschology and Development. Academic Press, New York, pp. 459-471.
  • Rosenblatt, J.S. 1995. Daniel Sanford Lehrman: June 1, 1919 – August 27, 1972. Biogr. Mem. Natl. Acad. Sci. 66, 3-21.