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52nd APS President (1979-1980)
Ernst Knobil
(b. 1926)
Knobil was born in Berlin, Germany, as a boy attended the Lycee Claude
Bernard in Paris, and then continued his education in science in the United
States at Cornell University (B.S., 1948; Ph.D., 1951) after two years of
service with the U.S. Army (1944-46). From 1951 to 1953 he was a Milton
Research Fellow in the laboratory of Roy O. Greep of the Harvard School of
Dental Medicine. This led to an appointment in the Department of Physiology
of the medical school (1953-61), which he left to become Richard Beatty
Mellon professor and chairman of the Department of Physiology at the School
of Medicine at the University of Pittsburgh (1961-81). From 1974 to 1981 he
was also director of the Center for Research in Primate Reproduction at
Pittsburgh. In 1981 he moved again to become dean of the School of Medicine
at the University of Texas Health Science Center (1981-84), H. Wayne
Hightower Professor of Physiology, and director of the Laboratory for
Neuroendocrinology (1981-). He now continues as a full-time investigator in
his laboratory.
Knobil's membership in APS dates from 1955. He was a member of the
Editorial Board of the Society's journals from 1960 to 1966 and of the
Editorial Board for the American Journal of Physiology: Regulatory,
Integrative and Comparative Physiology from 1976 to 1978. From 1979 to
1982 he was editor of the American Journal of Physiology: Endocrinology
and Metabolism. With Wilbur Sawyer he edited the volumes on the
pituitary gland and the hypothalamus of the Handbook of Physiology.
His first committee service was on the Committee for the Use and Care of
Animals (1962-67). He served on Council from 1969 to 1972 and later served
on the Ray G. Daggs Award Committee (1976-79). He became president elect in
1978. As president he was instrumental in increasing APS activities in
public affairs with a focus on the issue of animal experimentation.
Early in his career Knobil was honored by designation as a Markle Scholar
at Harvard (1956-61). He received the Ciba Award in 1961 and the Fred Conrad
Koch Award in 1982, both from the Endocrine Society. In 1983 he was chosen
for the Carl G. Hartman Award of the Society for Study of Reproduction, and
in 1985 he was given the Axel Munthe Prize in Reproduction. He has the
degree Dr. Hon. Causa from the University of Bordeaux (1980) and an honorary
Sc.D. from the Medical College of Wisconsin (1983). He is a fellow of AAAS
(Boston) and an honorary fellow of the American Association of Obstetricians
and Gynecologists, as well as of the American Gynecological Society.
Outside APS, Knobil has served on editorial boards for Endocrinology
(1959-75), Annual Review of Physiology (1968-77; editor-in-chief,
1974-77), Psychoneuroendocrinology (1974-77), Neuroendocrinology
(1976-80), and Endocrine Reviews (1980-84). A recent addition to the
list is Chronobiology International (1984-). For AAMC, Knobil served
as a member of the Executive Council for three years (1971-74), of the
Administrative Board of its Council of Academic Societies for four years
(1970-74), and more recently as a member of the Steering Committee on
Information Sciences and Medical Education (1984-). He was a member of the
Council of ACDP (1968-71; president, 1969-70); in 1983 he received its
Distinguished Service Award. Of the Deutsche Gesellschaft fur Endokrinologie
he is an honorary member, as he is of the Japan Endocrine Society; he holds
membership also in Great Britain's Society for Endocrinology. Other
memberships include the American Society of Zoologists, the International
Society for Research in Biology of Reproduction, the Society for Study of
Reproduction, the International Society for Neuroendocrinology, the Society
for Experimental Biology and Medicine, and the National Society for Medical
Research, which he served as a member of the Board of Directors (1977-79).
For three years (1968-71) he was a member of the Council of the Endocrine
Society (president, 1976-77). He has held a variety of responsibilities for
the International Society of Endocrinology, especially in reference to its
international congresses; from 1976 to 1984 he was chairman of its Executive
Committee and served as president from 1984 to 1988. From 1980 to 1983 he
was a member of the U.S. National Committee for IUPS and is currently
chairman of its Commission on Endocrinology (1982-).
Many organizations and institutions have asked Knobil to serve as
consultant. For NIH he served on the Human Growth and Development Study
Section (1964-66), the Reproductive Biology Study Section (chairman,
1966-68), the Advisory Committee on Primate Research Centers (1969-73), the
Contraceptive Development Branch (1969-71), the Medical Advisory Board of
the National Hormone and Pituitary Program (1980-83), and the Planning
Committee for Developmental Endocrinology and Physical Growth of the
National Institute of Child Health and Human Development (1984-), an
institute he had earlier served as a member of the Population Research
Committee of the Center for Population Research (1974-77). For three years
(1966-69) he was a member of the National Advisory Council of the Institute
of Laboratory Animal Resources of NAS. He served on the Physiology Test
Committee of the National Board of Medical Examiners (1970-74) and on the
Liaison Committee on Medical Education of the American Medical Association
(AMA) and AAMC (1971-74). In addition, he has been asked to serve as
consultant to the University of Michigan School of Medicine (1973), Harvard
Medical School (1973-74), the Ford Foundation (1974-75), the Uniformed
Services University of Health Sciences (1975), the Human Reproduction Unit
of the World Health Organization (1976-), the University of Texas Health
Science Center at Houston (1979-80), and the Advisory Committee of the
Searle Scholars Program (1981-82).
Knobil's research has generated widespread interest both in this country
and abroad. He has been invited to deliver the following lectures: Bowditch
Lecture of APS in 1965; Gregory Pincus Memorial Lecture of the Laurentian
Hormone Conference in 1973; and in 1974 the Upjohn Lecture of the American
Fertility Society, the Kathleen M. Osborn Memorial Lecture at the University
of Kansas School of Medicine, and the First Annual Hopkins-Maryland Lecture
in Reproductive Endocrinology. He gave the Karl Paschkis Lecture of the
Philadelphia Endocrine Society in 1975 and in 1978 was Sigma Xi Lecturer at
the University of Florida and Second Alza Lecturer in Palo Alto. In 1979 he
gave two lectures: the R. D. O'Brien Lecture at Cornell University and the
First Transatlantic Lecture of the Society for Endocrinology of Great
Britain. The following year (1980) he presented the Lawson Wilkins Lecture
of the Pediatric Endocrine Society and the Scientific Lecture for the
American Gynecological Society. In 1981 he was at Johns Hopkins for the
Second Bard Lecture, in San Francisco for the Sixth Herbert McLean Evans
Memorial Lecture, and at the University of Uppsala for the Fourth Carl
Gemzell Lecture. Since then he has given the Fourth Annual James H. Leathem
Lecture at Rutgers Medical School and the Geoffrey Harris Memorial Lecture
of the International Society of Neuroendocrinology, both in 1982; the Ayerst
Lecture of the American Fertility Society in 1983; and the Distinguished
Guest Lecture for the Society for Gynecological Investigation and the Potter
Lecture of Thomas Jefferson University, both in 1984.
Knobil's training in endocrinology involved only two laboratories.
"I began my career as a graduate student in zoology in the laboratory of
Samuel L. Leonard, an endocrinologist who was himself a student of F. L.
Hisaw and P. E. Smith, two of the founders of American endocrinology. My
mentor was a participant in the early development of this discipline in the
United States and imparted to me a very personal sense of belonging to this
branch of physiology. His Ph.D. thesis recounted the discovery of the
ovulatory hormone, now known at LH (Am. J. Physiol. 98: 406-416,
1931). From Leonard, I acquired an abiding love for endocrinology, which has
given me immense pleasure and happiness throughout my career. My formal
training was completed under the preceptorship of Roy O. Greep, who had been
a fellow student of Leonard in Hisaw's laboratory at the University of
Wisconsin."
"At Harvard, Greep introduced me to the world of academic medicine, which
I entered as an ingenuous postdoctoral research fellow. His unfailing and
often undeserved support is directly responsible for the path my
professional life has followed. He introduced me to E. M. Landis in the
medical school, who offered me an instructorship in his department (1953)
the year before he became president of the APS. My years as a member of his
faculty had a profound influence, in both style and substance, on the
development of the Department of Physiology at the University of Pittsburgh
School of Medicine, which I served as founding chairman for twenty years."
"My scientific contributions worthy of any note have all focused on the
physiology of the primate adenohypophysis (e.g., refs. 2 and 3). My
laboratory established the zoological specificity of growth hormone with the
finding that only growth hormone of primate origin is physiologically active
in primates (4). This discovery led to the development of a radioimmunoassay
for human growth hormone (6) and permitted a host of investigations dealing
with control of growth hormone secretion (5, 7)."
"A decade or so ago we began a systematic investigation of control of the
menstrual cycle, with the rhesus monkey as an experimental animal (10). This
was occasioned by recognition of major species differences in control of
reproductive processes and of the fact that findings obtained on one species
could not necessarily be extrapolated to another. In the course of this
effort, we were the first to describe the time course of progesterone during
the ovarian cycle of any species (8, 9) and to elucidate the endocrinologic
events of early pregnancy. This provided new insight into the control of the
corpus luteum and the role of progesterone in regulation of gonadotropin
secretion (11)."
"More recently we have delineated the neuroendocrine control system that
governs secretion of gonadotropic hormones throughout the menstrual cycle of
the rhesus monkey. In sum, it comprises the arcuate nucleus of the
hypothalamus, which mediates an hourly discharge of the hypothalamic
hormone, GnRH, with relatively simple negative and positive feedback loops
between estrogens and the pituitary gland. We now understand control of the
menstrual cycle for the first time. Consequently the ovarian cycle of
monkeys is understood better than that of any other species (12, 13)."
For his past president's address (14), Knobil abandoned the societal
introspection of his immediate predecessors to consider the classic volume
by William Harvey, On the Generation of Animals. Because spermatazoa
were unknown, being at that time invisible, Harvey remained puzzled as to
just what the male fluid may contribute to development and differentiation
of the ovum. Knobil noted that Harvey's treatise is "tedious, repetitive,
impossible to summarize, encapsulate, or even to sample adequately." It was
begun when he was a student in Padua (ca. 1600), under the teaching of
Fabricius, and apparently concluded only shortly before he was persuaded,
somewhat against his better judgment, to permit its publication (1651).
Knobil remarked that it is five times longer than Harvey's De Motu Cordis---and,
one judges, five times less conclusive. Nevertheless, Knobil found it to be
"vaguely reminiscent" of Claude Bernard's writings.
Commenting on his contributions to neuroendocrinology, one of his friends
recently stated that Knobil has qualities that remind one of Claude Bernard,
especially in his ability to analyze the integrative and relational aspects
of endocrine systems and their controls. Knobil has done this for
endocrinology of primates more successfully than anyone of his generation.
Selected Publications
1. Greep, R. O., E. Knobil, F. G. Hofmann, and T. L. Jones. Adrenal
cortical insufficiency in the rhesus monkey. Endocrinology 50:
664-676, 1952.
2. Knobil, E., A. Morse, F. G. Hofmann, and R. O. Greep. A histologic and
histochemical study of hypophyseal-adrenal cortical relationships in the
rhesus monkey. Acta Endocrinol. 17: 229-238, 1954.
3. Knobil, E., R. C. Wolf, R. O. Greep, and A. E. Wilhelmi. Effect of a
primate pituitary growth hormone preparation on nitrogen metabolism in the
hypophysectomized rhesus monkey. Endocrinology 60: 166-168, 1957.
4. Knobil, E., A. Morse, R. C. Wolf, and R. O. Greep. The action of
bovine, porcine and simian growth hormone preparations on the costochondral
junction in the hypophysectomized rhesus monkey. Endocrinology 62:
348-354, 1958.
5. Knobil, E., and R. O. Greep. The physiology of growth hormone with
particular reference to its action in the rhesus monkey and the species
specificity problem. Recent Prog. Horm. Res. 15: 1-69, 1959.
6. Knobil, E., and R. O. Greep. The detection of growth hormone in
plasma. In: Hormones in Human Plasma, edited by H. N. Antonaides.
Boston, MA: Little, Brown, 1960.
7. Knobil, E. Tenth Bowditch Lecture. The pituitary growth hormone: an
adventure in physiology. Physiologist 9: 25-44, 1966.
8. Neill, J. D., E. D. B. Johansson, and E. Knobil. Levels of
progesterone in peripheral plasma during the menstrual cycle of the rhesus
monkey. Endocrinology 81: 1161-1164, 1967.
9. Neill, J. D., E. D. B. Johansson, and E. Knobil. Patterns of
circulating progesterone concentrations during the fertile menstrual cycle
and the remainder of gestation in the rhesus monkey. Endocrinology
84: 45-48, 1969.
10. Monroe, S. E., L. E. Atkinson, and E. Knobil. Patterns of circulating
luteinizing hormone and their relation to plasma progesterone levels during
the menstrual cycle of the rhesus monkey. Endocrinology 87: 453-455,
1970.
11. Knobil, E. On the regulation of the primate corpus luteum. Biol.
Reprod. 8: 246-258, 1973.
12. Knobil, E. On the control of gonadotropin secretion in the rhesus
monkey. Recent Prog. Horm. Res. 30: 1-46, 1974.
13. Knobil, E. The neuroendocrine control of the menstrual cycle.
Recent Prog. Horm. Res. 36: 53-88, 1980.
14. Knobil, E. William Harvey and the physiology of reproduction.
Physiologist 24(1): 3-7, 1981.
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