Robert Elder Forster II

39th APS President (1966-1967)
Robert Elder Forster II
(b. 1919)

In planning for the XXIV IUPS Congress in Washington, D.C. (1968), Wallace Fenn chose Robert Forster to be chairman of the Finance Committee. The success of this committee has become legendary. With the help of what Forster called "the tremendous efforts of Wallace Fenn and of K. K. Chen," the committee raised enough money to leave the congress with a $65,000 surplus. (Friends have wondered whether Forster might have been a consultant to the U.S. Olympic Committee for the games in Los Angeles in 1984.) This surplus was turned over to APS as a trust fund, with the income of the fund devoted to support of travel grants to future congresses.

Forster was born in St. David's Pennsylvania, and has lived in that neighborhood for most of his life. He graduated from Radnor High School in Wayne in 1937 and with his family took up residence again in that area (Haverford) some twenty years ago. Much of his education and training, however, was completed in New England. From high school he enrolled in the Sheffield Scientific School of Yale University in biological sciences. At this time one could leave Yale in three years and get credit for the first year of medical school as the last year of college, so he entered the University of Pennsylvania School of Medicine in 1940. World War II accelerated medical school training, and he was able to graduate in December 1943 to take an internship in medicine at the Peter Bent Brigham Hospital in Boston. He had only an abbreviated period as a house officer, and in October 1944, having been discharged from the U.S. Navy as physically unfit, he entered the Army. After basic training at Carlisle, Pennsylvania, he was assigned to the Quartermaster Corps Climatic Research Laboratory in Lawrence, Massachusetts. This appointment was available because of the ill fortune of Clifford Barger, who had been at the laboratory but had contracted tuberculosis. At the Quartermaster Corps Laboratory, Forster did research on temperature regulation and heat exchange in the course of testing and helping design new field clothing and equipment for military personnel.

Forster had been interested in doing research as a medical student. Stimulated by his teachers, Julius Comroe and Carl Schmidt, he had measured changes in blood plasma viscosity with thiocyanate treatment for hypertension and had published his first article in the "yellow journal" of Lea and Febiger, the American Journal of the Medical Sciences (1). During several summer terms he worked at the Alfred I. DuPont Institute in Wilmington, which was then affiliated with the University of Pennsylvania School of Medicine, in biochemical research. There he first learned to operate a Van Slyke manometric gas analyzer under the tutelage of Douglas MacFadyen and Murray Angevine, who had been colleagues of D. D. Van Slyke at the Rockefeller Institute. Forster has described his subsequent training and tour of duty in the Quartermaster Corps Laboratory as follows:
"The laboratory was led by Colonel John H. Talbot, who had been on the high-altitude Andes expedition of 1935. Many distinguished individuals interested in the response of humans to extreme environmental conditions passed through the laboratory. Sir Hubert Wilkins, who had sailed in a submarine under the North Pole, acted as a consultant. He had the delightful trait of simply turning off his hearing aid when he thought the argument was going against him. Paul Siple, the Boy Scout who had been at Antarctica with Admiral Richard E. Byrd, also gave frequent advice. Cuthbert Bazett, chairman of the Department of Physiology at the University of Pennsylvania, came often for consultations."

"My time in the laboratory included a somewhat amusing experience. I had put together an electrical impedance plethysmograph, which measured the impedance of a segment of a finger at radio frequencies (and compared simultaneous changes in volume from a volume plethysmograph). The impedance plethysmograph was championed by Jan Nyboer, but unfortunately the instrument really does not measure changes in finger volume. I wrote to Alan Burton about the instrument and enclosed records of the impedance pulses. Burton replied that the impedance plethysmograph was of little account and that he could record better volume pulses from the projected shadow of a straw set in Plasticine on his toe. Bazett, when he was told of this put-down response, said that I should pay little attention; 'Alan does not know much physiology and what he does know I taught him.'"

"Richard L. Day, a pediatrician, had been at the laboratory and had developed a plethysmocalorimeter, a closed box which could act as a venous occlusion plethysmograph and whose walls acted as gradient calorimeter. (We have a painting of it in our office.) This work resulted in a number of publications on the relation of body temperature to hand and foot blood flow, but the most important was the publication of Bazett, Love, Newton, Eisenberg, Day, and Forster, in which countercurrent cooling of the arterial blood to the hand was demonstrated. (I believe this was the first experimental demonstration of this mechanism.) This appeared as the first article of the first volume of the Journal of Applied Physiology (2)."

Near the end of 1946 Forster was discharged from the Army and spent the next year as a graduate student in mathematics; he took courses in physics, mathematics, and physical chemistry in the graduate school at Harvard. When he returned in 1947 as a resident in internal medicine at the Peter Bent Brigham Hospital, he was permitted to measure the pulmonary arterial blood temperature with a wedge catheter in a patient of Lewis Dexter. He also borrowed an ear oximeter from Glenn Millikan for studies on the oxygenation of some of Samuel A. Levine's patients.

On a Life Insurance Medical Research Fund Fellowship, Forster then spent two years in the Department of Physiology at Harvard Medical School under Eugene M. Landis. He first worked on a thermostromuhr, a method for the measurement of intestinal blood flow. ("This really did not work very well.") He then moved on to measure changes in hypothalamic temperature in a chronic unanesthetized cat during changes in environmental temperature conditions. Because the hypothalamic temperature had to change nearly 0.5 degree Centigrade before the peripheral heat flow effectors were triggered, he found it to be a rather insensitive thermostat (3).

At the end of his postdoctoral fellowship Forster sought an academic post and visited several departments of medicine and physiology. (He was interviewed at Johns Hopkins Hospital under the impression he was applying for entrance to medical school.) He was offered, and he accepted, a position as assistant professor of physiology in anesthesiology under Robert Dripps and Julius Comroe in the Department of Physiology and Pharmacology of the University of Pennsylvania Graduate School of Medicine. Forster described his years in this department in these words:
"This was a very lucky choice, as I arrived at an exciting period in the short history of this department. Julius Comroe was rising to the peak of his reputation as a pulmonary physiologist and investigator of pulmonary function testing. The faculty included not only Comroe but Seymour Kety and later George Koelle, Ward S. Fowler, and Arthur DuBois. Kety and Comroe had already on order a respiratory mass spectrometer, to facilitate rapid measurements of inert respiratory gases, and an infrared meter, to measure low concentrations of carbon monoxide, which I was able to use to study the diffusing capacity of the lung. In the next few years a good deal of work was accomplished on diffusion exchanges in the lungs, including a description of the single-breath method for the measurement of the diffusing capacity of the lung, a widely used pulmonary function test (4). In collaboration with F. J. W. Roughton (5) I studied the theory and practice of measurements of pulmonary capillary blood volume (VC) and the diffusing capacity of the pulmonary membrane (DM). [During this period Forster was awarded a Lowell M. Palmer Fellowship (1954-56).]"

"Roughton introduced me also to rapid-mixing techniques for observing kinetic processes (6), and over the next several decades I was able to develop a number of different types of rapid- mixing apparatus to measure physiologically important chemical reactions and rapid exchanges of red cells, at first CO, O2, and CO2 but later anions and water. These studies have formed the foundation of my research career. In 1977 Itada and I introduced a method for the measurement of carbonic anhydrase inside intact red cells with 18O exchange (9), the only method so far reported for this purpose, because any other technique stops before significant measurements can be made because of accumulation of end products within the cell membrane. An offshoot of this interest in CO was work with Coburn and other colleagues that showed that a molecule of CO is produced for each heme group that is catabolized, at a rate of 0.4 ml/h in humans; this provides an index of the measurement of destruction of hemoglobin and other heme proteins (7)."

During this period Julius Comroe organized several courses to teach the growing body of knowledge on respiratory physiology and pulmonary function testing to physiologists from other institutions. Based on this experience, Comroe, Forster, DuBois, Briscoe, and Carlsen produced The Lung, which is just entering its third edition and has been a medical best-seller.

In 1957 Julius Comroe left the Department of Physiology and Pharmacology in the Graduate School of Medicine of the University of Pennsylvania to set up the new Cardiovascular Research Institute at the University of California School of Medicine at San Francisco. George Koelle became chairman of the Department of Physiology and Pharmacology in the School of Medicine to succeed Carl Schmidt. At this point Forster became chairman of the physiology part of the Graduate School of Medicine, the latter under John Brobeck. In 1970 Brobeck resigned in favor of Forster, and the existing departments were fused.

Forster attended his first spring meeting of the APS in 1946. He served on the Publications Committee from 1963 to 1965, was elected to Council in 1963, and was president elect in 1965. He was on the Editorial Board of the Handbook of Physiology (1973-79) and was a member of the Perkins Memorial Fund Committee (1968-70), the Daggs Award Committee (1975-78), and the Finance Committee (1978-83; chairman, 1982-83). He was also on the Editorial Board of the Journal of Clinical Investigation (1962-67). He served on the Advisory Board of the Life Insurance Medical Research Fund from 1967 until 1970, when it was unfortunately dissolved. He served in several advisory capacities to NIH: the Cardiovascular Study Section (1960-64), the General Clinical Research Center Committee (1964-67), and the National Advisory Heart Council (1967-71). He was on the Editorial Board of Annual Review of Physiology (1984-), and he was a member of the U.S. National Committee of IUPS (1976-83).

Many colleagues and associates from abroad have visited in Forster's laboratory, and he has returned the visits on many occasions. Although he has never taken a sabbatical year's leave, he did spend a long vacation at Trinity College with F. J. W. Roughton in 1954. Forster has long been interested in the application of physiology to underwater biology and to high-altitude and space conditions. In 1966 he was chairman of the NAS-NRC Space Science Board Summer Study Group on Respiratory Physiology that concluded that an atmosphere of pure O2 was dangerous and a first hazard in spacecraft. The report of this group appeared almost simultaneously with the tragic fire on the launching pad at Cap Canaveral, where several astronauts lost their lives. Forster wrote that he consequently experienced a 'transient notoriety."

One of the more notable events of Forster's association with APS was the fall meeting in 1976, the Bicentennial Year of the Declaration of Independence, held in Philadelphia and principally at the University of Pennsylvania. Brobeck happened to be the one who extended the invitation, but Forster was both officially and unofficially the host. The meeting was judged to be a grand success. Especially gratifying to Forster was the fact that it was "the first Fall Meeting in many years that did not require any subsidization by the American Physiological Society."

Selected Publications

1. Forster, R. E. The medical use of thiocyanates in the treatment of arterial hypertension. Am. J. Med. Sci. 206: 668-686, 1943.

2. Bazett, H. C., L. Love, M. Newton, L. Eisenberg, R. Day, and R. E. Forster. Temperature changes in blood flowing in arteries and veins in man. J. Appl. Physiol. 1: 3-19, 1948.

3. Forster, R. E>, and T. B. Ferguson. The relationship between hypothalamic temperature and thermoregulatory effectors in unanesthetized cat. Am. J. Physiol. 169: 255-269, 1952.

4. Ogilvie, C. M., R. E. Forster, W. S. Blakemore, and J. W. Morton. A standardized breath holding technique for the clinical measurement of the diffusing capacity of the lung for carbon monoxide. J. Clin. Invest. 36: 1-17, 1957. (Abstr. Federation Proc. 14: 108, 1955 and J. Clin. Invest. 34: 917, 1955.)

5. Roughton, F. J. W., and R. E. Forster. Relative importance of diffusion and chemical reaction rates in determining rate of exchange of gases in the human lung, with special reference to the true diffusing capacity of pulmonary membrane and volume of blood in the lung capillaries. J. Appl. Physiol. 11: 290-302, 1957.(Abstr. Am. J. Physiol. 183: 615-616, 1955.)

6. Roughton, F. J. W., R. E. Forster, and L. Cander. Rate at which carbon monoxide replaces oxygen from combination with human hemoglobin in solution and in the red cell. J. Appl. Physiol. 11: 269-276, 1957.

7. Coburn, R. F., W. J. Williams, and R. E. Forster. Effect of erythrocyte destruction of carbon monoxide production in man. J. Clin. Invest. 43: 1098-1103, 1964.

8. Constantine, H. P., M. R. Craw, and R. E. Forster. Rate of the reaction of carbon dioxide with human red blood cells. Am. J. Physiol. 208: 801-811, 1965.

9. Itada, N., and R. E. Forster. Carbonic anhydrase activity in intact red blood cells measured with 18O exchange. J. Biol. Chem. 252: 3881-3890, 1977.

10. Dodgson, S. J., R. E. Forster II, B. T. Storey, and L. Mela. Mitochondrial carbonic anhydrase. Proc. Natl. Acad. Sci. USA 77: 5562-5566, 1980.

11. Dodgson, S. J., R. E. Forster II, D. E. Schwed, and B. T. Storey. Contribution of matrix carbonic anhydrase to citrulline synthesis in isolated guinea pig liver mitochondria. J. Biol. Chem. 258: 7696-7701, 1983.