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
Arnold A. Berthold
Courtesy of the National Library of Medicine.
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).
Berthold, A.A., 1849. Transplantation der Hoden. Arch.
Anat. Physiol. Wissenschr. Med. pp. 42-46. In German.
A.A., 1849. Transplantation of testes. English
translation by D. P. Quiring,
1944, Bull. Hist. Med. 16,
Frank A. Beach (1911-1988)
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.
- 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.
- 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.
F.A., 1976. Sexual
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 C. Young
William Caldwell Young (1899-1965)
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.
- 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.
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
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.
- 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
- Dantchakoff V. 1937. Realisation du sexe a volonte. Bull.
Biol. 3, 1-53.
Daniel S. Lehrman (1919-1972)
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.
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.),
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.