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58th APS President (1985-1986)
Howard Edwin Morgan
(1927 - 2009)
In the life of APS, little of significance happens solely within one
year, confined to the twelve months of a single presidency. Rather, in the
normal sequence from councillor through past president, each president takes
part in important deliberations and decisions over a period of several
years. For example, during the three years he was in presidential offices,
Morgan became closely involved in planning for the Centennial Celebration
because the long process of making these plans began to come to a focus in
1984-87. He was instrumental in making the final agreement for a project
many years in the making---the joint publication with IUPS of News in
Physiological Sciences. He also took an active part in the lengthy
consideration of how to ensure a broader representation of the several
sections by modifying governance of the Society. Finally it was in the year
when Morgan was president elect that Orr E. Reynolds retired from the
position of executive secretary-treasurer of APS and Martin Frank was
appointed to that office. Morgan became therefore the first president to
hold office in collaboration with Frank, as Berne had been the first to
serve with Reynolds in 1973. Morgan brought to the office extensive
experience not only with the Society's journals (see later) but also in the
deliberations of the Porter Physiology Development Committee (1968-1980).
Morgan was born in Bloomington, Illinois, and began his college education
there with one year at the Illinois Wesleyan University (1944-45). He then
moved directly into medical school at Johns Hopkins University, where he
received his M.D. degree in 1949. His original intention was to become an
obstetrician-gynecologist, a career he began on the house staff of the
hospital of Vanderbilt University (1949-53). The following year (1953-54) he
was instructor in these disciplines. He then became for a year a fellow in
medical research in the unit of the Howard Hughes Medical Institute
established in the Department of Physiology at Vanderbilt (1954-55). But the
following year he was back in obstetrics and gynecology as assistant chief
of that service on active duty in the U.S. Army Station Hospital at Fort
Campbell, Kentucky. He then returned to Vanderbilt, and for the next ten
years (1957-67) he was an investigator in the Hughes Institute, with faculty
rank that progressed from assistant professor (1959-62), to associate
professor (1962-66), and professor (1966-67). Morgan then became the first
professor and chairman of the Department of Physiology in the Milton S.
Hershey Medical Center of the Pennsylvania State University in Hershey,
Pennsylvania. From 1973 he has been also Associate Dean for Research, and in
1974 was honored by designation as the Evan Pugh Professor of Physiology. In
1982 he was further honored by appointment as a scholar of the Howard Hughes
Medical Institute. Morgan wrote briefly of his training:
"Because I entered physiological research after eight years of clinical
training, research, and practice in obstetrics and gynecology, my training
was entirely as a postdoctoral fellow. Charles R. Park served as my
preceptor and guided me into studies of the effects of insulin on glucose
uptake and sugar transport. With a solid background obtained in Park's
laboratory, I later was able to undertake the new areas of investigation
that have characterized the remainder of my career."
Before he became a member of APS (1965), Morgan had been elected to the
Biochemical Society (Great Britain, 1960) and the American Society of
Biological Chemists (1962). He holds membership also in the Biophysical
Society (1965), the Cardiac Muscle Society (1969; president, 1976-77), ACDP
(president, 1975-76), and AHA (Board of Directors, 1984-). In the Basic
Science Council of AHA, Morgan served on the Executive Committee from 1973
to 1979 and again from 1981 to the present (chairman, 1983-). He has been a
member of the Executive Committee of the American Section of the
International Society for Heart Research (1976-79; president, 1979-82). From
this office he became president elect of the International Society (1980-83)
and has served as president (1983-86). Morgan therefore has served as
chairman or president of a major scientific organization in all but two of
the past eleven years, and in the year 1985-86 he held three such offices
simultaneously.
In addition to the honors noted above, including those from the Howard
Hughes Medical Institute and from his own university, Morgan has received an
Award of Merit from AHA (1979), the Carl J. Wiggers Award from the
Cardiovascular Section of APS (1984), and an honorary fellowship in the
American College of Cardiology (1985). He was elected to APS Council in 1983
and became president elect the following year.
In areas related to cardiology, Morgan has provided scientific counsel to
the Research Committee of the Pennsylvania Heart Association (1967-71;
chairman, Research Peer Review Committee, 1983-); the Research Council of
the New York City Heart Association (1974); the US:USSR Exchange Program for
Problem Area 3, Myocardial Metabolism (coordinator, 1974-83); and AHA. He
served as a member of the Physiological Chemistry A Research Study Group of
AHA (1973-75; chairman 1976-79) and of the AHA Research Committee (1974-79
and 1980-81). In 1977-78 he was vice-president for research, chairman of the
Research Committee, and a member of the Board of Directors of AHA. NIH has
called on him for membership in the Metabolism Study Section (1967-71), on
an ad hoc committee for the National Heart Center Program (1973), on a
Cardiology Advisory Committee (1975-78), and on the Advisory Council of the
National Heart, Lung and Blood Institute (1979-83). In 1982 Morgan was asked
to be chairman of a special panel appointed by this latter institute "to
review allege misconduct at Brigham and Women's Hospital/Harvard Medical
School." Finally, he now holds membership on the U.S. National Committee for
IUPS (1984-87).
Another important feature of Morgan's career is his association with
scientific journals. Beginning with the Editorial Board of the American
Journal of Physiology (1967-73), he became editor of Physiological
Reviews (1973-78), associate editor of the American Journal of
Physiology: Endocrinology and Metabolism (1979-81), and editor of the
American Journal of Physiology: Cell Physiology (1981-84). For much of
this time he served on the Publications Committee (1979-85; chairman,
1981-85). Other journals for which he has provided editorial assistance
include Circulation Research (1971-76 and 1982-), the Journal of
Biological Chemistry (1973-78 and 1980-85), the Journal of
Cardiovascular Pharmacology (1977-82), and the Journal of Molecular
and Cellular Cardiology (1974-; associate editor, 1979-83). Of this
listing, his influence was perhaps the greatest on Physiological Reviews.
During his tenure as editor it grew significantly in international
reputation and influence.
Morgan's research interest is the physiological regulation of
intermediary metabolism. For many of his studies he has used the isolated
and perfused rat heart. He has described his work as follows:
"Initial studies dealt with the mechanism of action of insulin on glucose
uptake and the nature of glucose transport. Insulin was found to accelerate
glucose transport, a stereospecific, saturable process in the cell membrane
(1). A kinetic model of sugar transport was proposed, based on studies in
rabbit erythrocytes (2). This model and its mathematical description have
been used by many other investigators in characterizing transport phenomena.
Experiments measuring the rate of glycogen utilization led to investigation
of the allosteric control of phosphorylase a and b and to the
discovery that phosphorylase b activity was increased by 5'-AMP
[adenosine 5'-monophosphate] and inhibited by ATP [adenosine triphosphate]
and G-6-P [glucose 6-phosphate] (3). This mechanism of allosteric control
accounted for the differential effects of anoxia and glucagon and for
acceleration of glycogen utilization in working hearts."
"My interest in the effects of heart work on cardiac metabolism led to
development of the isolated perfused working rat heart (4) that has been
used extensively both in our laboratory and elsewhere for study of the
effects of mechanical performance on carbohydrate, fat, and protein
metabolism. In this model, perfusion medium is introduced into the left
atrium over a range of atrial filling pressures and is pumped against a
variable outflow resistance. With this model, myocardial oxygen consumption
was found to depend on the aortic pressure to which the heart was exposed;
greater oxygen consumption was accompanied by faster utilization of
oxidative substrates."
During the next phase of my research career, my interest shifted to
identification of factors that control growth of the heart and that can lead
to cardiac hypertrophy. Initiation of peptide chains on myocardial ribosomes
was found to become a rate-controlling step during in vitro perfusion and to
be accelerated by insulin, fatty acids, and other noncarbohydrate
substrates, leucine, increased cardiac work, and exposure to higher aortic
pressure (5, 6). A rigorous method for estimation of rates of protein
synthesis was developed that depended on measurements of the specific
activities of phenylalanyl-tRNA (7). Protein degradation also was identified
as a site of control of protein turnover that is affected by insulin,
diabetes, energy availability, noncarbohydrate substrates, leucine, cardiac
work, and increased aortic pressure (6, 8). The factor that links cardiac
work to faster rates of protein synthesis and slower proteolysis (9) appears
to be stretch of the ventricular wall, because these effects could be
observed in hearts arrested with tetrodotoxin and containing a ventricular
drain. In these preparations, an increase in aortic pressure stretched the
ventricular wall, accelerated protein synthesis, and inhibited proteolysis.
These events appear to represent early changes in the hypertrophy process."
"After longer periods of exposure to pressure overload or to
thyrotoxicosis in vivo, we found that content of cardiac RNA increased and
accounted for much of the increment in protein synthesis. since ribosomal
RNA constitutes about eighty-five percent of cardiac RNA, these changes
indicated that net ribosome production was increased, either by acceleration
of rRNA transcription or processing or by inhibition of rRNA degradation
(10). These events are the focus of my current research."
During the past twenty-five years the presidents of APS have often
expressed concern about the apparent fragmentation of the science and the
development of diverse and presumably independent interests by members of
the Society. A countertendency is beautifully illustrated in the lecture
Morgan gave when he received the Carl J. Wiggers award in 1984. He described
experiments that began with a problem in classic physiology, the response of
the heart to increased load. However, he pursued the response, not only by
use of traditional physiological measurements such as oxygen consumption,
but on through analysis of pathways of protein, carbohydrate, and lipid
metabolism, until he reached the measurement of rates of synthesis and
degradation of the several forms of RNA. The experiments moved clearly and
easily from the whole organ to the level of molecular biology. His lecture
illustrates how what seem to be old-fashioned problems can be studied by
using the most sophisticated of modern techniques to provide a clearer
understanding of what really takes place in living organisms.
Selected Publications
1. Morgan, H. E., M. J. Henderson, D. M. Regen, and C. R. Park.
Regulation of glucose uptake in muscle. I. The effects of insulin and anoxia
on glucose transport and phosphorylation in the isolated, perfused heart of
normal rats. J. Biol. Chem. 236: 253-261, 1961. (Citation classic.)
[Dr. Morgan's first paper in a series on the control of glucose uptake.]
2. Regen, D. M., and H. E. Morgan. Studies of the glucose-transport
system in the rabbit erythrocyte. Biochim. Biophys. Acta 79: 151-166,
1964. [Mathematical model of glucose transport.]
3. Morgan, H. E., and A. Parmeggiani. Regulation of glycogenolysis in
muscle. III. Control of muscle phosphorylase activity. J. Biol. Chem.
239: 2440-2445, 1964.[Discovery of the allosteric control of phosphorylase.]
4. Neely, J. R., H. Liebermeister, E. Blattersby, and H. E. Morgan.
Effects of pressure development on oxygen consumption by the isolated rat
heart. Am. J. Physiol. 212: 810-814, 1967. [Development of in vitro
working rat heart.]
5. Morgan, H. E., D. C. Earl, A. Broadus, E. B. Wolpert, K. E. Giger and
L. S. Jefferson. Regulation of protein synthesis in heart muscle. I. Effect
of amino acid levels on protein synthesis. J. Biol. Chem. 246:
2152-2162, 1971. [Initial paper on control of protein synthesis.]
6. Rannels, D. E., R. L. Kao, and H. E. Morgan. Effect of insulin on
protein turnover in heart muscle. J. Biol. Chem. 250: 1694-1701,
1975. [Initial description of the effect of insulin on protein degradation
in heart.]
7. McKee, E. E., J. Y. Cheung, D. E. Rannels, and H. E. Morgan.
Measurement of the rate of protein synthesis and compartmentation of heart
phenylalanine. J. Biol. Chem. 253: 1030-1040, 1978. [Discovery of a
rigorous approach to measurements of protein synthesis.]
8. Morgan, H. E., B. H. L. Chua, N. E. O. Fuller, and D. Siehl.
Regulation of protein synthesis and degradation during in vitro cardiac
work. Am. J. Physiol. 238 (Endocrinol. Metab. 1): E431-E442,
1980. [Discovery of effects of cardiac work on protein turnover.]
9. Kira, Y., P. J. Kochel, E. E. Gordon, and H. E. Morgan. Aortic
perfusion pressure as a determinant of cardiac protein synthesis. Am. J.
Physiol. 246 (Cell Physiol. 15): C247-C258, 1984. [Discovery of
stretch as the mechanical factor affecting protein turnover.]
10. Siehl, D., B. H. L. Chua, N. Lautensack-Belser, and H. E. Morgan.
Faster protein and ribosome synthesis in thyroxine-induced hypertrophy of
rat heart.Am. J. Physiol. 248 (Cell Physiol. 17): C309-C319,
1985. [Discovery of role of increased ribosome content in cardiac
hypertrophy.]
11. Morgan, H. E., E. E. Gordon, Y. Kira, D. L. Siehl, P. A. Watson, and
B. H.-L. Chua. Biochemical correlates of myocardial hypertrophy. Wiggers
Award Lecture. Physiologist 28: 18-27, 1985.
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