Howard Edwin Morgan

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.