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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.
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