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WALLACE OSGOOD PENN
Al ugust 27, 1893-September 20, 1971
BY HERMANN RAHN
WALLACE OSGOOD FENN was born in Lanesboro (Berkshire
County), Massachusetts and died in Rochester, New York
in his seventy-ninth year, after a brief illness. He is survived by
his widows, Clara Bryce (Comstock) Fenn; his children, William
Wallace Fenn, Ruth (Fenn) Starman, Priscilla (Fenn) Roslansky,
and David Bryce Fenn; and ten grandchildren. He led a most
vigorous life and up to his very last days was working in the
laboratory; during his last three years he shaped the new direc-
tions of the International Union of Physiological Sciences as its
President. To many of his colleagues he was the Dean of Physio-
logical Sciences, the last Renaissance Man, whose basic contri-
butions covered so many areas and who had a remarkable per-
spective on the whole field of biology.
His forefathers settled in New England in the seventeenth
century. William Wallace Fenn, his father, was a Unitarian
minister who had married Faith Huntington Fisher, also from
New England. Later his father became the Bussey Professor of
Theology at Harvard and Dean of the Divinity School. Thus
Wallace Fenn's childhood was spent in Cambridge, where he
attended the Cambridge Latin School and entered Harvard with
the goal of preparing himself for the ministry. However, when
he started cutting his father's lectures to attend plant physi-
ologist W. J. V. Osterhout's classes in biology, the foundations
141
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BIOGRAPHICAL MEMOIRS
were laid for a career in physiology that was to span more than
half a century.
Fenn graduated in 1914. His graduate work at Harvard with
Osterhout was interrupted by World War I, during which he
served in the Sanitary Corps of the U.S. Army and was com-
missioned a Second Lieutenant. Upon discharge in 1919, he fin-
ished his doctoral thesis in Curie, married Clara Bryce Comstock
in September, and began his appointment as an Instructor in
Applied Physiology in the Department of Industrial Hygiene at
the Harvard Medical School under Cecil K. Drinker. Here
began his classical studies of phagocytosis of solid particles by
white blood corpuscles.
In 1922 he accepted a Rockefeller Travel Fellowship and
was the first American to work in A. V. Hill's laboratory in
London, England. This was followed by a six-month stay in
H. H. Dale's laboratory at the National Institute for Medical
Research in London. Returning to this country in 1924, he
accepted the Chair of Physiology at the newly formed Medical
School at the University of Rochester, New York. This position
he filled for thirty-five years. In 1961 he was named Distinguished
Professor of Physiology, a post he occupied until his death in
1971.
THE SCIENTIST
Fenn's first paper was published in 1916 in the Proceedings
of the National A cademy of Sciences. It was entitled "Salt
Antagonism in Gelatin." His last paper, "Partial Pressure of
Gases Dissolved at Great Depth," was published posthumously
in Science in 1972. During the intervening half-century his 267
publications can be conveniently divided into four general areas:
the physiology of muscle, electrolytes, respiration, and high
pressure. In each area he laid foundations of new concepts, and
when he was satisfied that he had made new basic contributions,
moved on to explore new fields.
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WALLACE OSGOOD FENN
CONTRIBUTIONS TO MUSCLE AND
ELECTROLYTE PHYSIOLOGY
143
The work that brought Fenn his first recognition was his
study on the heat production of muscle, which he started in
A. V. Hill's laboratory in 1922-1923. Fenn wrote: "In particular
it can now be shown that there is a fairly good quantitative
relation between the heat production of muscles and the work
which they perform, and that a muscle which does work liberates,
ipso facto, an extra supply of energy which does not appear in an
isometric contraction."3t It was A. V. Hill who referred to this
as the Fenn Effect, and so it has been known ever since.
Fenn's heat data showed first of all that if a muscle shortens,
no matter how little and no matter how lightly loaded, it pro-
duces more heat than during an isometric contraction over the
same time period. He then showed that this extra heat produc-
tion was proportional to the external work done by the muscle.
It was clearly not determined by the load alone, nor by the
change in length. This was the first evidence, and remains today
the best evidence, that shortening is an active process and that
muscle is riot simply a prestretched spring shortening passively.
The Fenn Effect has emerged as the nearest thing to a law that
muscle physiologists have.
Following his pioneer work on muscle heat production, Fenn
began to measure gas exchange by nerve and by muscle. To this
end he had to invent a number of ingenious instruments to
obtain the necessary specificity and precision. In 1927 he meas-
ured for the first time the quantitative amount of oxygen
required by a nerve to conduct an impulse. Similar studies on
the metabolism of contracting muscles led him to consider the
role of electrolytes, particularly potassium, in nerve and muscle
Wallace Osgood Fenn, "A Quantitative Comparison between the Energy
Liberated and the Work Performed by the Isolated Sartorius Muscle of the
Frog," Journal of Physiology, 58(1924): 175.
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BIOGRAPHICAL MEMOIRS
activity. At the time, although it was known that muscle fibers
were rich in potassium, almost nothing was known of the mecha-
nisms by which cells accumulated and maintained a high potas-
sium content.
The work ushered in the era of electrolyte physiology. Begin-
ning in 1933 Fenn virtually created the field of potassium metab-
olism. He made the first determinations of potassium, sodium,
magnesium, and calcium in nerve. He developed a new method
for determining internal pH of muscle and nerve and obtained
values that remain acceptable today. He showed that intra-
cellular potassium was mobile, not fixed, and that muscle potas-
sium shifted in response to various environmental factors.
Most importantly, he showed that during contraction potas-
sium was lost from muscle in exchange for sodium, and that the
process was reversed in recovery. For the first time he showed
that sodium could penetrate muscle. These observations were
clearly the necessary foundation for the Hodgkin-Huxley hypo-
theses concerning initiation and propagation of nerve and
muscle impulses and the magnitude and polarity of electrical
potential differences across cell membranes. As early as 1936, at
the Cold Spring Harbor Symposium, Fenn said, "The explana-
tion of a loss of potassium from a muscle during activity is a
matter of fundamental theoretical importance. In terms of the
theory which I have been using as a guide, it is interpreted as an
increase in the permeability of the muscle membrane of sufficient
extent to permit sodium, but not chloride, to enter. Every mole-
cule of sodium which enters then displaces one molecule of
potassium." ~
Fenn showed that potassium escaped from muscle during
contraction in situ and that a large part of this potassium ap-
peared in the liver. He demonstrated that potassium uptake was
#Wallace Osgood Fenn, "Electrolytes in Muscle," Cold Spring Harbor Symp.
Quant. Viol., 4(1936):252-59.
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WALLACE OSGOOD FEN N
145
linked with carbohydrate metabolism, particularly with glycogen
deposition, and developed the concept that potassium tends to
follow the Cori cycle. He was always quick to seize new oppor-
tunities. When radioactive potassium became available to him
in 1939, he ingested a sample. Using himself as subject, he was
thus the first not only to study the kinetics of potassium metabo-
lism but also to demonstrate potassium incorporation into blood
cells, previously thought to be impermeable. He showed that
nearly all muscle potassium in the body is exchangeable, proving
that high intracellular potassium content is not maintained by
binding or sequestration of potassium, an idea which was con-
sonant with his notion that potassium is maintained by an active
energetic process.
CONTRIBUTIONS TO RESPIRATION PHYSIOLOGY
The entrance of Wallace 0. Fenn into the history of respira-
tory physiology can be precisely dated. It was within days after
the U.S. entry into World War II. At that time he was forty-eight
years old and had established himself as the acknowledged leader
in the physiology of muscle and electrolytes. He was to be recog-
nized in 1943 by election to the National Academy of Sciences.
Wallace Fenn was drawn into respiratory physiology by his
desire to contribute to the war effort. This was to be largely a
war in the air, and from a military point of view, supremacy in
altitude tolerance meant supremacy of air power. The airplanes
of that day did not yet have pressurized cabins, but the possi-
bility occurred that the human lung might be pressurized by
application of positive pressure breathing. The question was
whether man's lungs could tolerate a sufficient amount of pres-
sure to raise the partial pressure of oxygen to a significant degree,
or would the lungs rupture, or would the circulation stop? What
were the limiting factors? What were the hazards?
What was known about respiratory physiology in general?
This can best be answered by listing some terms which did not
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146
BIOGRAPHICAL MEMOIRS
appear in the physiology textbooks of that era, but which are
commonplace today. Such terms are: positive and intermittent
pressure breathing, pressure-volume diagram, work of breath-
ing, pulmonary compliance, airway resistance, alveolar gas equa-
tion, O'-CO2 diagram, ventilation-perfusion ratio, just to name
a few.
Wallace Fenn had never worked in the field of human
respiration. The equipment in his laboratory would be regarded
as primitive by current standards. Among the more useful items
were a few assorted spirometers, two or three Haldane machines,
an equal number of Van Slykes, and several U-tube manometers.
The most sophisticated instrument was a Millikan ear oximeter,
which had been loaned to him by the Military. It carried a
security classification of a fairly high level, and since no instruc-
tion manual came with it, it took some time and a visit to Glen
Millikan himself before anyone could figure out how to use it
properly.
In addition to this modest inventory of equipment, Fenn had
three young instructors, all trained in biology departments.
They knew all about such things as how fast the drosophila can
beat its wings, how and why the rattlesnake changes color, and
how to activate or inhibit enzymes found in grasshopper eggs,
but none of them had ever blown a vital capacity; neither did
they know the difference between complemental and supplemen-
tal air. L. E. Chadwick, A. B. Otis, and H. Rahn, living with
their wives on postdoctoral stipends which were only a fraction
of what a graduate student receives today, were the most unlikely
crew to have been assembled for the unknown job that lay ahead
of them.
Neither the equipment nor the staff was very impressive,
and it seems doubtful that by present standards the project could
have qualified for a National Institutes of Health grant. How-
ever, the major asset, recognizable even then, was Wallace Fenn
himself. He was not put off by lack of ready-made equipment; he
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WALLACE OSGOOD FEN N
147
was well endowed with Yankee ingenuity, and he loved to impro-
vise. He could, with whatever components happened to be
handy, construct apparatus that would perform in a reliable and
effective fashion. Everyone associated with him has memories of
him in the laboratory surrounded by what at first sight appeared
to be an unrelated jumble of strange wires and rubber bands,
tubing, pulleys, lenses, light sources, mirrors, and other assorted
bits and pieces. A more careful examination suggested there
might be some order in the arrangement, and further observa-
tion would reveal that something of physiological interest was
actually being measured and perhaps even graphically recorded.
A relatively refined example was a device for the automatic re-
cording of blood flow through the finger and its alteration by
pressure breathing.
The high-altitude chamber was perhaps the crowning master-
piece of Fenn's ingenuity. He had received from the Committee
on Medical Research of the Office of Scientific Research and
Development a contract which provided the sum of $500 (five
hundred!) for special research equipment. From this budget he
bought a steel tank designed for the processing or transport of
beer, commandeered the tree-spraying pump from the Univer-
sity Grounds Department, reversed its valves, and connected
Dumb to tank. The result was a chamber which could go to
1 ~
. .
simulated altitudes at the rate of 5,000 feet per minute. As he
later said, "It surely was the worst high altitude chamber in the
country, but a rare atmos ?here is the same wherever you find
it."
Not only could he get the most out of primitive pieces of
equipment, but he also seemed somehow able to evoke the best
output from his staff. He did not tell people to do things. Rather,
he pointed out things that needed doing and waited for some-
#Wallace Osgood Penn, "Born Fifty Years Too Soon," Annual Review of
Physiology, 24(1962):1.
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148
BIOGRAPHICAL MEMOIRS
thing to happen. He worked hard himself and expected others to
do likewise, but he recognized that there were individual differ-
ences in effective work patterns and did not try to impose his
own habits on others. Although he kept rather regular working
hours himself, he apparently was not perturbed by those with
more erratic habits. Getting something done rather than com-
pulsive adherence to a fixed schedule was the important thing.
In starting a new experiment he frequently took the lead by
setting up apparatus himself rather than asking someone else to
do it. Typically, he would insist on being the first subject in a
new experimental procedure, and in experiments with pressure
breathing and in the altitude chamber he extended himself on a
number of occasions to the point of losing consciousness. He was
a pioneer in every sense, and it was a blessing that his work ante-
dated the Human Subjects Review Committee.
Fenn's intuitive approach to and logical analysis of the pres-
sure breathing problem led him to develop two powerful con-
cepts and to express them in the form of graphic relationships:
the pressure-volume diagram of the lung and thorax, and the
O2-CO2 diagram of the composition of alveolar gas.
Although the basic pressure-volume (P-V) diagram had been
previously developed by F. Rohrer, Penn conceived it independ-
ently, elaborated it further, and distilled into it some ten years of
work and thought. Like all his work, it defined physiological
boundaries, limiting values for muscle forces and the corre-
sponding volumes of gas and blood. Within these limits were
centered the normal operating range of pulmonary mechanics
and the response of the system to positive and negative pressure
breathing. It was not only a beautiful composition both artistic-
ally and scientifically, but it was also the foundation and frame-
work of respiratory mechanics that would be further embellished
by students during succeeding decades.
Fenn's second masterpiece, the O2-CO2 diagram, did for
pulmonary gas exchange what the P-V diagram did for respira-
tory mechanics. With it he could represent all parameters of the
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WALLACE OSGOOD FENN
149
alveolar gas and ventilation equations. He never claimed to have
originated these equations, but he derived them independently,
made sure they were correct, and put them in graphic form. As
somebody put it, "That's when he made them sing." On the
diagram he could show all possible compositions of alveolar gas
and the arterial blood under any specified set of conditions. He
could indicate normal ranges and limits of survival as well as
the pathways followed during hyperventilation and asphyxia
and during exposure to CO2, altitude, or hyperbaric pressures.
It could be used to demonstrate ranges of normal and impaired
performance. It was indeed a theme that could be sung with
. .
many varlatlons.
Although the P-V and O2-CO2 diagrams represent great
masterpieces of Fenn's scientific artistry, he created, inspired, or
contributed to many other works. To give a few examples: devel-
opment of the concept of an optimal breathing frequency, meas-
urement of alveolar pressure, dynamic pressure-volume curves
presented for the first time on a cathode ray oscilloscope, devel-
opment of an infrared CO2 meter, and probably the first pub-
lished continuous recording of CO2 changes during a single
breath.
Finally, one must mention two special contributions to
respiration physiology, a lasting monument to his effort in this
area: his book, ~4 Graphical Analysis of the Respiratory Gas
Exchange, which went through many reprintings, and his editor-
ship of Respiration in the Handbook of Physiology series, both
of these published by The American Physiological Society.
CONTRIBUTIONS TO THE PHYSIOLOGY OF
SPACE AND OCEANS
From the mid-1950s Fenn became greatly intrigued with two
new frontiers that began to unfold man's explorations in space
and the ocean depths. While his research continued in very basic
experiments, their application was obviously directed to filling
in the gaps of knowledge so that man could exist successfully in
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BIOGRAPHICAL MEMOIRS
these new environments. He was in great demand as a consultant
by space physiologists and tried to convince his more earth-
bound colleagues of the great new opportunities in physiology
that unfolded as man ventured into space. Every problem Fenn
"considered basic, if the investigator put some basic thinking
into it." In that sense he felt that physiology as a science had
gained immeasurably and would continue to grow as man went
forth in orbit and into the oceans thoughts that he expressed
so well in his address "Physiology in Orbit."*
Wherever man went he needed oxygen as the life-sustaining
gas, yet when it exceeded normal pressures it became poisonous.
Fenn spent many years with his associates (R. Gerschman and
D. L. Gilbert) in trying to understand the toxic nature of oxy-
gen. Probably his most important insight was the recognition
and demonstration that oxygen poisoning and X-irradiation
effects have the same common mechanism.
He also turned his attention to the effects of high inert gas
pressures upon the metabolism of unicellular organisms and the
effects of hydraulic pressure on biological reactions. His last
benchwork emphasized the importance of partial molar volume
concepts as a tool for determining the volume that Of occupies
within the hemoglobin structure." His last research concerned
itself with the theoretical concepts of partial pressures of gases
dissolved at great depths. It was a thermodynamic interpretation
published posthumously in Science,+ where with his great mod-
esty he asked for the help of physical chemists to develop this
concept in greater detail, help which shortly appeared.§
Wallace Osgood Fenn, "Physiology in Orbit," The Physiologist, 3(1960):20-26.
~ Wallace Osgood Fenn, "Partial Molar Volumes of Oxygen and Carbon
Monoxide in Blood," Respiratory Physiology, 13~1971~ :129~0.
~ Wallace Osgood Fenn, "Partial Pressure of Gases Dissolved at Great Depth,"
Science, 176(1972):1011-12.
§ F. C. Andrews, "Gravitational Effects on Concentrations and Partial Pressures
in Solutions: A Thermodynamic Analysis," Science' 178(1972):1199-1201.
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WALLACE OSGOOD FENN
163
With M. Goettsch. Electrolytes in nutritional muscular dystrophy in
rabbits. J. Biol. Chem., 120:41-50.
Loss of potassium from stimulated frog muscle. Proc. Soc. Exp. Biol.
Med., 37:71-74.
1938
With D. M. Cobb, i. F. Manery, and W. B. Bloor. Electrolyte
changes in cat muscle during stimulation. Am. J. Physiol., 121:
595-608.
With J. H. Wills. Potassium changes in submaxillary glands during
stimulation. Am. l. Physiol., 124:72-76.
Factors affecting the loss of potassium from stimulated muscles. Am.
J. Physiol., 124:213-29.
With B. H. Carleton. Studies on respiration of muscle in the presence
of carbon monoxide. l. Cell. Comp. Physiol., 11:91-98.
The potassium and water contents of cat nerves as affected by stimu-
lation. J. Neurophysiol., 1: 1-3.
1939
The fate of potassium liberated from muscles during activity. Am. I.
Physiol., 217: 356-73.
With W. S. Wilde, R. A. Boak, and R. H. Koenen~ann. The effect of
blood flow on potassium liberation from muscle. Am. J. Physiol.,
128: 139~6.
The deposition of potassium and phosphate with glycogen in rat
livers. l. Biol. Chem., 128:297-307.
The distribution of excess potassium in cats. In: Professor Alvaro e
Miguel Ozorio de Almeida, Livro de Homenagem, pp. 197-202,
Rio de ~aneiro, Brazil.
1940
With R. H. Koenemann, B. V. Favata, and E. T. Sheridan. The role
of the lactic acid in the movements of potassium. Am. i. Physiol.,
131 :494-508.
With L. F. Haege. The deposition of glycogen with water in the
livers of cats. T. Biol. Chem., 136:87-101.
With R. H. Koenemann and E. T. Sheridan. Potassium exchange of
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164
BIOGRAPHICAL MEMOIRS
perfused frog muscle during asphyxia. J. Cell. Comp. Physiol.,
16:255-64.
The role of potassium in physiological processes. Physiol. Rev., 20:
377-415.
1941
With T. R. Noonan and L. F. Haege. The distribution of injected
radioactive potassium in rats. Am. J. Physiol., 132:474-88.
With T. R. Noonan and L. F. Haege. The effects of denervation and
of stimulation on exchange of radio-active potassium in muscle.
Am. J. Physiol., 132: 612-21.
With L. J. Mullins, T. R. Noonan, and L. F. Haege. Permeability of
erythrocytes to radioactive potassium. Am. J. Physiol., 135:
93-101.
With T. R. Noonan, L. l. Mullins, and L. F. Haege. The exchange
of radioactive potassium with body potassium. Am. J. Physiol.,
135: 149-163.
Muscle. Annul Rev. Physiol., 3:209-32.
Preface. In: Biological Symposia, ed. l. Cattell, 3:vii-ix. Lancaster,
Pa.: Cattell Press.
Introduction to muscle physiology. In: Biological Symposia, ed.
I. Cattell, 3: 1-8. Lancaster, Pa.: Cattell Press.
With R. B. Dean, T. R. Noonan, and L. F. Haege. Permeability of
erythrocytes to radioactive potassium. J. Gen. Physiol., 24:353-65.
1942
With L. F. Haege. The penetration of magnesium into frog muscle.
J. Cell. Comp. Physiol., 19:37-46.
With W. F. Bale and L. J. Mullins. The radioactivity of potassium
from human sources. l. Gen. Physiol., 25:345-53.
1944
With L. F. Haege, E. Sheridan, and l. B. Flick. The penetration of
ammonia into frog muscle. l. Gen. Physiol., 28: 53-77.
1945
Muscles. In: Physical Chemistry of Cells and Tissues, ed. R. Hober.
Philadelphia: Blakiston.
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WALLACE OSGOOD FEN N
1946
16S
With H. Rahn, A. B. Otis, and L. E. Chadwick. The pressure volume
diagram of the thorax and lung. Am. l. Physiol., 146: 161-78.
With A. B. Otis, H. Rahn, and M. A. Epstein. Performance as related
to composition of alveolar air. Am. I. Physiol., 146:207-21.
With A. B. Otis and H. Rahn. Venous pressure changes associated
with positive intra-pulmonary pressures: their relationship to the
distensibility of the lung. Am. J. Physiol., 146:307-17.
With H. Rahn and A. B. Otis. A theoretical study of the composition
of the alveolar air at altitude. Am. I. Physiol., 146:637-53.
With H. Rahn, A. B. Otis, M. Hodge, M. A. Epstein, and S. W.
Hunter. The effects of hypocapnia on performance. l. Aviat.
Med., 17: 164-72.
With H. Rahn, I. Mohney, and A. B. Otis. A method for the con-
tinuous analysis of alveolar air. i. Aviat. Med., 17: 173-78.
With A. B. Otis, H. Rahn, M. Brontman, and L. I. Mullins. Ballisto-
cardiographic study of changes in cardiac output due to respira-
tion. J. Clin. Invest., 25:413-21.
1947
With A. B. Otis, H. Rahn, L. E. Chadwick, and A. H. Hegnauer.
Displacement of blood from the lungs by pressure breathing. Am.
J Physiol., 151 :258-69.
With L. E. Chadwick. Effect of pressure breathing on blood flow
through the fingers. Am. J. Physiol., 151 :270-75.
With A. L. Barach, E. B. Ferris, and C. F. Schmidt. The physiology
of pressure breathing: a brief review of its present status. J.
Aviat. Med., 18: 73-87.
With R. J. Dern. The effect of varying pulmonary pressure on the
arterial pressures in men and anesthetized cats. J. Clin. Invest.,
26:460-67.
1948
With A. B. Otis and H. Rahn. Alveolar gas changes during breath
holding. Am. .T- Physiol., 152:674-86.
Physiology of exposures to abnormal concentrations of the respira-
tory gases. Proc. Am. Philos. Soc., 92: 145-54.
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166
BIOGRAPHICAL MEMOIRS
1949
Physiology on horseback. Am. J. Physiol., 159:~51-55.
With R. Galambos, A. B. Otis, and H. Rahn. Corneoretinal potential
in anoxia and acapnia. J. Appl. Physiol., 1:710-16.
With H. Rahn and A. B. Otis. Daily variations of vital capacity,
residual air, an expiratory reserve, including a study of the resid-
ual air method. J. Appl. Physiol., 1: 725-36.
With H. Rahn, A. B. Otis, and L. E. Chadwick. Voluntary pressure
breathing at high altitudes. l. Appl. Physiol., 1:752-72.
With H. Rahn, A. B. Otis, and L. E. Chadwick. Physiological obser-
vations on hyperventilation at altitude with intermittent pressure
breathing by the pneumolator. i. Appl. Physiol., 1: 773-89.
With R. T. Clark, Jr., and J. N. Stannard. Evidence for the conver-
sion of carbon monoxide to carbon dioxide by the intact animal.
Science, 109:615-16.
Potassium. Sci. Amer., 181 (No. 2:~16-21.
1950
With R. T. Clark, in and l. N. Stannard. The burning of CO to CO2
by isolated tissues as shown by the use of radioactive carbon. Am.
J. Physiol., 161:40~6.
With H. Rahn and A. B. Otis. Respiratory system. Annul Rev.
Physiol., 7: 179-204.
With A. B. Otis, W. O. Fenn, and H. Rahn. Mechanics of breathing
in man. J. Appl. Physiol., 2:592-607.
With R. Gerschman. The loss of potassium from frog nerves in
anoxia and other conditions. l. Gen. Physiol., 33: 195-203.
Department of physiology and vital economics. In: The Quarter
Century 1925-50, pp. 53-60. Rochester, N.Y.: The University of
Rochester.
1951
Mechanics of respiration. American journal of Medicine, 10:77-90.
With R. Gerschman, G. Fischer, J. Lacy, M. Bailly, and J. L. Wright.
Experiments on the role of potassium in the blocking of neuro-
muscular transmission by curare and other drugs. J. Gen. Physiol.,
34:~.
Medical aspects of military manpower selection. In: The Selection of
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WALLACE OSGOOD FENN
167
Military Manpower, ed. L. Carmichael and L. C. Mead, pp. 28-
37, National Academy of Sciences Publ. No. 209. Wash., D.C.:
National Academy of Sciences.
Medical aspects of military manpower selection. Sci. Mon., 73:
209-12.
With A. B. Otis and H. Rahn. Studies in respiratory physiology. Air
Force Technical Report No. 6528. U.S. Air Force, Wright Air
Development Center, August.
lg52
With A. B. DuBois, R. C. Fowler, and A. Softer. Alveolar CO2
measured by expiration into the rapid infra-red gas analyzer. J.
Appl. Physiol., 4:526-34.
With A. B. DuBois and A. G. Britt. Alveolar CO2 during the respira-
tory cycle. J. Appl. Physiol., 4:535~8.
With A. B. DuBois and A. G. Britt. COP dissociation curve of lung
tissue. J. Appl. Physiol., 5: 13-16.
Cost of medical education. Bull. Monroe County Med. Soc. Roches-
ter Acad. Med., 10:49-52.
The great goldrush in medical research. University Tennessee Rec-
ord, 55:78-89.
lg53
With F. H. Freeman. Changes in carbon dioxide stores of rats due to
atmosphere low in oxygen or high in carbon dioxide. Am. J.
Physiol., 174:422-30.
Acute and sustained high energy output. Symposium on stress, Army
Medical Services Graduate School, Walter Reed Army Medical
Center, Wash., D.C., March.
lg54
With A. B. Otis and M. Suskind. The accumulation of carbon di-
oxide in apneic dogs during intermittent oxygen insulation.
Am. l. Physiol. Med., 33:299-312.
With R. Gerschman. Ascorbic acid content of adrenal glands of rat
in oxygen poisoning. Am. J. Physiol., 176:6-8.
With R. Gerschman, D. L. Gilbert, S. W. Nye, and P. W. Nadig.
Role of adrenalectomy and adrenal-cortical hormones in oxygen
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Representative terms from entire chapter:
biographical memoirs
