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H E RMANN RAH N
July 5, 1912-June 23, 1990
BY iOHN PAPPENHEIMER
I do not wish to be a professor and just sit and count the veins of a butter-
fly (wing) and then try to determine their names. I could not stand that for
long. I am more interested in the whole animal world.... My ideal is not a
professor but a man like Ernest Thomson Seton. . . by reading his books I
try to learn how to live in the woods, how one can sleep warm there in the
winter at below freezing temperatures, what wood burns best, how to make
a fire in the rain, what provisions one takes on a long trip when you are all
alone.
SO WROTE THE SIXTEEN-YEAR-QED Hermann Rahn to his clos-
est frienc! en c! schoolmate, Wolf Tischier, in Germany.
Despite these youthful sentiments, Hermann clic! become a
professor en c! one of the foremost physiologists of his gen-
eration. Only a few years later, while still a student, he again
wrote to his friend, Wolf, this time saying:
Natural History is not the problem of today, it is merely a good and inter-
esting basis. I believe that experimental zoology with its "cause and effects"
has gotten hold of me.... I was torn between two worlds. Both afford
interesting problems, but I never found a connection between them. You
work either systemically and anatomically or on the other hand as physiolo-
gist and experimental zoologist. According to my mind the latter offers the
greater problems.... After being torn between these two worlds it is a
good feeling at last to know my way.
These were serious thoughts for a schoolboy, en c! they
243
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244
B I O G RA P H I C A L
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reveal Hermann's commitment to research in biology at an
early age. In the enc! he fount! the connection between the
two worIcis, en c! incleec! his abiding love of natural history
en c! the call of the wiTc! cleterminec! the course of much of
his experimental work on respiratory en c! comparative physi-
ology.
Hermann was brought up in an academic milieu. His
father, Otto Rahn, was professor of bacteriology en c! dairy
chemistry, first at Kiel in Germany en c! later at Cornell
University in Ithaca, New York. His mother knee Bell FarrancI,
ISS3) was a fourth-generation native of Lansing, Michigan,
en c! she was Otto's research assistant at the Agricultural
College in Lansing prior to their marriage. Hermann was
the elclest of their four chilciren, en c! most of his formative
years were spent in a happy, stable, and intellectually stimu-
lating family environment in Kiel, Germany, en c! Ithaca,
New York. Nevertheless, the first few years of his life were
unusually turbulent, no account of his life wouIc! be com-
plete without providing brief biographies of his parents and
the history of their early marries! life.
Otto Rahn was one of eleven chilciren born to uneclu-
catec! Mennonite parents in the little town of Tiegenshof in
East Prussia. He was a star student at the town school, and,
with the sympathetic support of his father and his math-
ematics teacher, he was able to attenc! the University of
Gottingen, where he stucliec! physical chemistry with W.
Nernst en c! organic chemistry with Professor WalIach (later
to become Nobel laureate in chemistry).
Otto obtainer! his Ph.D. in 1902 at the age of twenty-one,
but academic positions in chemistry were scarce, en c! he
acceptec! a position in the Department of Dairy Science.
His acivancec! training in organic chemistry en c! mathemat-
ics was unusual in this fielcI, en c! he soon fount! applica-
tions to bacterial metabolism that brought him international
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HERMANN RAHN
245
renown. In 1906 he was offerer! a faculty position in bacte-
riology at Michigan State and 1907 found him struggling
with the English language while teaching chemistry en c!
bacteriology to students in the agricultural college at Lan-
slng.
In the same year, Bell Farranc! (Hermann's mother-to-
be) gracluatec! from Michigan State en c! acceptec! a job as
assistant to Professor Clinton Dewitt Smith, who was about
to become president of a small agricultural college in
Perchicaba, Brazil, six-hours travel west of San PauTo. That
was an extraordinary adventure for a well-brought-up young
lady from Michigan in 1907, but in her own words, "My
nature has always been a bit on the romantic sicle en c! hav-
ing a young en c! adventurous heart I just conic! not turn
clown such an offer to see the worIcI, even though I clic! not
like Mr. Smith." Her romantic nature was to be transmitter!
en c! amplifier! in her son Hermann, whose interests in envi-
ronmental and comparative physiology led him to roam the
woricI. Bell returnee! to Lansing in 1908 to join the Depart-
ment of Bacteriology uncler Professor Marshall, but she was
soon assignee! to Otto Rahn as a research en c! teaching
assistant. They were marries! in 1911, en c! Hermann was
born on July 5, 1912.
Two years later the Rahns boarclec! a ship for Germany so
that Bell could meet Otto's family and show off their two-
year-oIc! Hermann to relatives. The Archduke Ferclinanc!
was assassinates! while they were on the high seas, en c! WorIc!
War I began shortly after they clisembarkoc! in Hamburg.
Although Otto was in the process of becoming a U.S.
citizen, he en c! his American bricle were cIassifiec! in Ger-
many as German citizens, en c! they were not permitted to
return to the Uniter! States even though Bell was six months
pregnant and anxious to return home. Hermann's sister,
Marie, was born in November 1914, en c! soon thereafter
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B I O G RA P H I C A L
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Otto was ciraftec! into the army en c! assignee! menial acimin-
istrative jobs at a small airfield! in Latvia.
One of Otto's sisters owner! a bookstore in Danzig, en c!
Bell, Hermann, en c! the baby mover! into a room over the
store. For the next four years Bell was cut off from commu-
nication with her own family in Michigan, en c! it was not
until November 1918 that she learner! through the Norwe-
gian Rec! Cross of her mother's cleath. Hermann, by then
six years oIcI, hac! no shoes en c! froze his toes while waiting
in line for the tinily family ration of ~ pint of skimmer!
milk. Bell's footwear was reclucec! to her satin wocicling slip-
pers until Otto manager! to provide her with a pair of sec-
oncI-hanc! army boots he hac! traclec! for his tobacco rations.
After the war anti-German feelings remainec! high in the
Uniter! States, en c! since Otto hac! servec! in the German
army he was not invites! to return to Michigan. At the same
time, German universities were in disarray, en c! it was not
until 1920 that he obtainer! a suitable post as professor of
ciairy physics at Kiel. Hermann was then eight years oicI,
en c! so his early schooling began in Germany. At this time,
also, he en c! his frienc! TischIer (later to become professor
of ecology at Kiel) began collecting en c! identifying butter-
flies, insects, bircis, en c! fauna from the beaches of the Bal
tic Sea. In 1923 his parents were able to transfer Hermann's
savings account of $19.00 from America to Kiel, where in-
flation was such that they were able to buy him a micro-
scope and a camera, "hoping he would someday become a
scientist."
In 1925 W. A. Noyes, professor of chemistry at Illinois
(and a member of the National Academy of Sciences), made
a goodwill tour of German universities, and he became in-
terested in Otto Rahn's work on the physical properties of
milk products, noting that no comparable department of
dairy physics existed in the United States. Unbeknownst to
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HERMANN RAHN
247
her husband, Mrs. (Bell) Rahn conficlec! to Noyes that she
yearner! to return home to visit her family en c! asker! whether
it might be possible for him to arrange a lecture tour for
her husband. This iclea bore fruit, en c! in 1926 the entire
Rahn family visitor! America, the chilciren learning English
from their American cousins while Professor Rahn went coast
to coast on his lecture tour. Shortly after returning to Kie!
he receiver! an offer to come to Cornell University as ten-
urec! professor of bacteriology en c! dairy physics. Hermann
was enrollee! in the local high school in Ithaca en c! starter!
his transition to the American eclucational system en c! way
of life. The wilclerness of the Finger Lake district of upper
New York state was in contrast to the manicures! country
arounc! Kiel, en c! Hermann was fascinates! by it. Through-
out his school en c! unclergracluate years at Cornell he spent
many clays en c! sometimes weeks camping out in the wiTcis,
collecting or identifying the flora en c! fauna. During the
summers he took jobs as nature counselor at a Boy Scout
camp or as assistant in government fisheries or wiTcIlife cle-
partments. At college he grounclec! himself in the chemical
en c! physical sciences neeclec! for his planner! career in ex-
perimental zoology. After graduating from Cornell in 1933,
Hermann returnee! to Kiel for one year before enrolling as
a graduate student en c! teaching assistant in zoology at the
University of Rochester. His roots in Germany were creep,
en c! he was torn between Germany en c! the Uniter! States. It
was not until 1936, after a seconc! visit to Kiel, that he was
able to write to his frienc! Wolf, ". . . America has at last
become my real home."
CONTRIBUTIONS TO REPRODUCTIVE PHYSIOLOGY OF
SNAKES AND BIRDS, 1937-43
Hermann's first publications, based on his independent
work as a graduate student, were concernec! with the repro
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B I O G RA P H I C A L
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cluctive physiology of viviparous snakes en c! on the clevelop-
ment of the pituitary glanc! in bircis. His discovery in 1937
that viviparous snakes develop a primitive placenta analo-
gous to the mammalian organ won him a National Research
Council postcloctoral fellowship to work in reproductive
physiology with Frederick Hisaw at the Harvarc! Biological
Laboratories. His year at Harvarc! (1938-39) was eviclently a
productive one for it lee! to a series of eight papers on the
structure en c! function of the pituitary in bircis en c! snakes.
With Louis Kleinho~z he clevelopec! a biological assay for
the melanophore-stimulating hormone ("intermeclin") of
the pars intermeclia en c! cleterminec! its activity in a variety
of mammalian species. At the same time, he completec! a
cletailec! histological stucly of cell types in the pars anterior
of eighteen species of bircis en c! shower! that all these spe-
cies lackey! an intermediate lobe. In the same year he fount!
that female garter snakes, collectec! from Penekese Islanc!
off Cape CocI, conic! store viable sperm in utero for at least
one month following insemination, so that the exact time
of fertilization of ova and the gestation period were inde-
terminate. Finally, he fount! time to court en c! marry
Katherine (Kay) Wilson, a student at the Graduate School
of Landscape Architecture.
In September 1939 Hermann mover! to his first academic
post as instructor in zoology at the University of Wyoming
at Laramie. There he macle goof! use of the mountains en c!
prairies to combine his love of nature en c! natural history
with his interests in the reproductive behavior of reptiles.
He fount! that rattlesnakes living at an altitucle of 6,000
feet, where the summers are short, have a two-year repro-
ductive cycle, a phenomenon made possible by storage of
viable sperm over the winter in a special pocket of the
uterus. During the winter hibernation period, also, mature
ova were retainer! in the ovaries en c! not clischargec! to meet
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HERMANN RAHN
249
the sperm until spring. This bizarre schecluTe meant that at
any one time cluring the summer approximately 50 percent
of the aclult females were gravicI. At lower altitucles, where
the summers are longer en c! warmer, the same species of
rattler has a one-year reproductive cycle, en c! almost all the
aclult females are gravicI. Rahn's principal paper describing
this work (publisher! in Copeia, 1942) is as convincing as it
is interesting, en c! the simple but original style of this early
publication foreshadowed the elegant simplicity of exposi-
tion for which he later became noted. His sojourns on the
prairies also lee! him to stucly reproductive behavior en c!
sexual dimorphism of the sage grouse, when Hermann spoke
of the elaborate courtship ciances of these bircis, you lis-
tenec! to sheer poetry.
PULMONARY MECHANICS AND BLOOD-GAS EXCHANGE;
YEARS WITH WALLACE FENN, 1941-56
WorIc! War II en c! chance events abruptly alterec! the course
of Rahn's career en c! the direction of his research. Of the
chance events, uncloubtecIly the most important was his
meeting with Wallace Fenn in Rochester in the summer of
1941. This meeting occurrec! when Hermann, coming east
from Wyoming for a visit to his parents, stopper! briefly in
Rochester to visit friends before proceeding to Ithaca. He
caller! on Fenn, whom he greatly acimirecI, en c! before their
conversation was over Fenn offered him a job as instructor
in physiology, an offer that was acceptec! on the spot. Fenn
was already one of the most distinguished general physiolo-
gists in the country, having macle pioneer contributions to
the mechanism of phagocytosis, the heat production of con-
tracting muscle, the metabolism of active nerve, en c! the
exchange of electrolytes in excitable tissues.1 Hermann was
to become Fenn's closest colleague, confidant, and scien-
tific protege. Shortly after the United States entered World
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B I O G RA P H I C A L
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War II, the National Research Council asker! Fenn to inves-
tigate the possibility that the operational altitucle of Air
Force personnel might be increasec! by breathing oxygen
uncler pressure (positive pressure breathing). Neither Fenn
nor his junior colleagues hac! ever worker! in the field! of
human respiration, but they acceptec! the challenge en c! in
the perioc! ~941-45 clevelopec! funciamental new approaches
to pulmonary mechanics en c! respiratory gas exchange-
concepts that helpec! to introduce a goIclen age of theoreti-
cal and applied respiratory physiology in the decade follow-
ing WorIc! War II. In this clevelopment, Hermann Rahn playact
a central role, although his association with Fenn en c! other
colleagues was so close that it is clifficult for a biographer
to separate the relative contributions macle by each incli-
viclual. It is reasonable to suppose, however, that Fenn's
quantitative biophysical approach awakened latent talents
in Hermann Rahn that macle him a full partner in the
enterprise en c! shaper! his own approach to biological prob-
lems cluring the next forty years.
Two major contributions to respiratory physiology emerged
from the 1941-45 work on positive pressure breathing by
the Rochester team, en c! both were publisher! in 1946 in
the American Journal of Physiology. The first was titles! "The
Pressure-Volume Diagram of the Thorax en c! Lung" with
Rahn as senior author, the seconc! was "A Theoretical Stucly
of the Composition of Alveolar Air at Altitude" with Fenn
as senior author. The first paper became the starting point
for research on pulmonary mechanics in many physiologi-
cal en c! clinical laboratories. The seconc! paper proviclec! a
graphical solution to equations describing the partial pres-
sures of oxygen en c! carbon clioxicle in alveolar gas as a
function of barometric pressure (altitucle), inspirer! gas com-
position, en c! respiratory exchange ratio. While most of the
equations underlying this analysis had been derived inde
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HERMANN RAHN
251
penclently by others, their representation in a graphical form
that conic! easily be unclerstooc! en c! applier! to a variety of
problems was a major contribution comparable to the cIas-
sic nomographic analysis of blooc! chemistry by L. l.
Henclerson.2 IncleecI, the next important step was to extent!
the analysis to the bloocI-gas exchange, en c! in 1949 Rahn
published his now classic paper titled "A Concept of Mean
Alveolar Air en c! the Ventilation-Blooc! Flow Relationships
During Pulmonary Gas Exchange." In this paper Rahn shower!
how regional differences in the ratio of alveolar ventilation
to alveolar blooc! perfusion (VA/Q) give rise to oxygen pres-
sure differences between mean alveolar gas en c! blooc! leav-
ing the lungs. His analysis was presented in a clear graphi-
cal form that has been user! by many subsequent investigators.
At the time of this work, there were no experimental meth-
ocis for determining regional pulmonary blooc! flow or ven-
tilation, en c! Rahn hac! to assume normal Gaussian clistribu-
tions in order to provicle numerical solutions in graphical
form. More than ten years later, when methods for cleter-
mining regional ventilation en c! perfusion using raclioac-
tive gases hac! been clevelopec! by I. B. West3 en c! others, it
was fount! that the distribution of VA/Q was far from Gaussian.
Nevertheless, the new experimental ciata were easily incor-
poratec! into Rahn's theoretical analysis, which continues
to be the preferred means of presenting the data. Abnor-
malities of VA/Q, rather than diffusion capacity, prover! to
be the most common cause of poor oxygenation of arterial
blooc! in a variety of pulmonary diseases, en c! Rahn's analy-
sis provides the theoretical basis for clinical tests of im-
paired gas exchange.
The techniques en c! concepts clevelopec! to investigate
respiratory gas exchange during acute exposure to low baro-
metric pressures (altitucle) were well suites! to studies of
other perturbations of the respiratory environment, includ
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B I O G RA P H I C A L
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ing the inhalation of CO2, hyperventilation, breath-hoicI-
ing, cliving, en c! acclimatization to altitucle. All of these per-
turbations were stucliec! by the Rochester team, but for Rahn
the lure of the mountains was not to be cleniecI, en c! cluring
the immediate postwar years he organizer! three expecli-
tions to high altitucles in Wyoming, Coloraclo, en c! the Pe-
ruvian Ancles. He en c! his colleagues were first to show that
respiratory acclimatization ant! cleacclimatization to altitucle,
measurer! in terms of alveolar gas composition, occurs ex-
ponentially with a half-time of about twelve hours. The re-
sults were clearly clelineatec! as a hysteresis loop on the
Fenn-Rahn O2-CO2 diagram, en c! they proviclec! the starting
point for subsequent studies by Severinghaus4 en c! others
showing that the time course of acclimatization is cleter-
minec! by changes in composition of cerebral fluicis bathing
meclullary chemoreceptors.
BLOOD-GAS EXCHANGE AT HIGH AND LOW PRESSURES;
PHYSIOLOGY OF DIVING IN THE AMA (DIVING WOMEN)
OF KOREA AND JAPAN, 1956-68
In 1956 Rahn mover! to the University of Buffalo Meclical
School as chairman of the Department of Physiology en c!
with him mover! the center of gravity of the Rochester school
of respiratory physiology. In the years to come he was to
attract more than 100 collaborators from some twelve coun-
tries to work on such diverse topics as respiratory gas ex-
change in cliving insects, the regulation of pH in polkilo-
therms, the role of nitrogen in the absorption of gas pockets
in animals en c! humans, distribution of ventilation en c! per
fusion in health en c! disease, respiratory gas equations as
applied to gill breathing, the physiology of diving in the
Ama sea-women of Korea en c! Japan, en c! allometric studies
of gas exchange through the eggshells of cleveloping birc!
embryos ranging from hummingbirds to ostriches. Although
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HERMANN RAHN
-
~ 100
Lo
~ 10
In
CE
J
J
- 1
UJ
an
o .1
as
lo
o
.01
~100
OXYGEN CONDUCTANCE
1- ~,
|~-WITH
. 1 1 1
1 1 0 1 00 1 ,000
EGG MASS (GRAMS)
257
In
10 ~
111
3
I
CD
I
J
llJ
o
.1
.01
Figure 2 Pore length and oxygen conductance increase at different
rates with increasing egg mass, as is shown in this graph encompass-
ing data from the eggs of some ninety species from different parts of
the world. For every tenfold increase in mass, the oxygen conduc-
tance of the eggshell increases 6.5 times, but the pore length in-
creases only 2.7 times. Pore length probably increases slower be-
cause the eggshell must be thin enough for the embryo to hatch.
the heartbeat of developing bird embryos by means of a
microphone placer! in a sealer! chamber containing the egg.
SERVICE TO SCIENCE: NATIONAL AND INTERNATIONAL
Rahn was a member of numerous scientific societies, en c!
he received distinguished service awards from several of
them. However, his primary allegiance was to the American
Physiological Society (APS) en c! to the International Union
of Physiological Sciences (IUPS). He was president of APS
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B I O G RA P H I C A L
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(1963-64) and served for many years as scientific editor and
boars! member of its publications, inclucling the compre-
hensive en c! scholarly two-volume Handbook of Respiratory
Physiology, which crownec! more than a clecacle of major acI-
vances in this fielcI. He also gave generously of his time to
advisory panels of the National Research Council, the Na-
tional Institutes of Health, the National Aeronautics en c!
Space Administration, en c! the American Institute of Bio-
logical Sciences.
Travel en c! international aspects of physiology playact a
major role in Rahn's life. His roots in Europe, his colIabo-
rative research with physiologists from many countries, en c!
his sensitivity to different cultures gave him a strong voice
in the International Union of Physiological Sciences. He
server! on its council from 1965 to 1971 en c! subsequently
as its vice-presiclent. From Wallace Fenn he absorber! a strong
tradition of loyalty to the triennial international congresses
of physiology, en c! he server! on the executive committee of
the large en c! successful XXIVth Congress hell! in Washing-
ton, D.C., in 1968. On several occasions he server! as resi-
clent visiting professor at foreign universities.
TEACHER, SCHOLAR, AND GENTLEMAN
Rahn grew up in a prewar academic environment in which
research was regarclec! as a joyous, spare-time privilege of a
university teacher rather than a ciriving professional career.
This point of view changer! rapicIly after the war, when large-
scaTe government support for research macle it possible for
young scientists to create incliviclual research empires with-
out regarc! for teaching or other traclitional academic re-
sponsibilities. Rahn was especially vulnerable to this devel-
opment because his research on gas exchange at high en c!
low barometric pressures had important applications to both
clinical and military problems. He was well supported by
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HERMANN RAHN
259
contracts from the Air Force en c! the Office of Naval Re-
search as well as from the National Science Foundation en c!
the National Institutes of Health, it wouIc! have been easy
for him to neglect his teaching en c! university responsibili-
ties, but to this temptation he never succumbed. Instead,
he remained true to his principles, namely, that the pri-
mary responsibility of university professors is to their stu-
clents en c! departments. At the time he mover! to Buffalo in
1956, he organizer! a comprehensive course in human physi-
ology for meclical students, en c! he continues! to play a ma-
jor role in teaching it throughout his tenure as heat! of the
department. At the same time, he createc! a stimulating
research environment for all members of his staff as well as
for the continual stream of postcloctoral fellows, many from
abroad, who came to work with him. He was a magnetic
source of icleas, drawing in all those arounc! him, en c! over
the years he colIaboratec! en c! publisher! with all fourteen
of his permanent staff members. As one staff member put
it, "tRahn] hac! a way of sharing his excitement over a new
iclea en c! before you knew it both of you were in the lab
trying it out."
Hermann Rahn was equally at home in the wiTclerness
en c! in the most formal settings. In civilizer! society he usu-
ally ciressec! impeccably, en c! his innate courtesy, modesty,
and sensitivity to others (perhaps best described as "courtTi-
ness") allowed him to fit in with all social situations, how-
ever foreign or sophisticated. His concern for others en c!
his willingness to take on responsibility enclearec! him to all
those who hac! the privilege of working with him. I have a
vivic! memory of an exhaustec! Hermann after he hostel! a
three-day meeting of 800 members of the American Physi-
ological Society in Buffalo, he was walking back to the lab
with drooping shoulders, laden with shopping bags full of
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B I O G RA P H I C A L
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presents for the secretaries en c! others who hac! helpec! him
organize the meeting.
In 1973 Rahn retiree! as chairman of the department en c!
became distinguished service professor of physiology. The
International renown he brought to this university was fur-
ther recognizec! by various awards, including the Stockton
Kimball Awarc! in 1969 en c! the Chancellor Norton Mecial
in 1981. In 1985 the experimental cliving laboratory that
hac! been constructor! by the Office of Naval Research for
Rahn's studies of underwater physiology was renames! "The
Hermann Rahn Laboratory of Environmental Physiology."
In the spring of 1990 Rahn learner! that he hac! incur-
able pancreatic cancer, but he continues! to work in the lab
as long as physically possible, en c! he was working on a
manuscript in bee! at home a few clays before the anti. In
one of his last letters to his lifelong frienc! Wolf Tischier he
commented! on his life:
. . . the general maturing of a happy child with his insect collecting, his
love with all nature, his wonders and aspirations . . . to the mature student,
the young investigator and finally the reflecting scientist . . . I am happy to
have stayed a romantic in science. Today my colleagues have become busi-
ness scientists and I am sure your colleagues have to do the same in order
to survive as researchers. So we have both been most fortunate because we
are both in a sense still children, with our youthful enthusiasm to explore
and search for answers.
ACKNOWLEDGMENTS ARE DUE PROFESSOR Wolfgang Tischler for reading
the manuscript of this memoir and for giving permission to quote
from his correspondence with Hermann; Hermann's sister, Marie
Wohlmann, and his son, Robert, for allowing me to read and quote
from the typewritten autobiographies left to them by Otto and Bell
Rahn; and members of Rahn's staff at Buffalo, especially Charles
Paganelli, R. Blake Reeves, and Augusta Dustan for their comments.
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HERMANN RAHN
NOTES
261
1. See "Wallace O. Penn," in Biographical Memoirs, vol. 50, pp.
141-73. Washington, D.C.: National Academy of Sciences, 1979.
2. L. T. Henderson. Blood: A Study in General Physiology. New Ha-
ven, Conn.: Yale University Press, 1928.
3. T. B. West. British Medical Bulletin, 19~1963~:53-60.
4. T. W. Severinghaus, et al. Journal of Applied Physiology, 18~1963~:1155
56.
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262
S E L E C T E D
1938-39
1960
1963-64
1964
1965
1966
1968
1971
1971-74
1973
1977
1980
1981
1981
1985
B I O G RA P H I C A L
EMOIRS
AWARDS AND DISTINCTIONS
National Research Council fellow
Harvey Society lecturer
President, American Physiological Society
Doctor of medicine, Honoris Causa, University of Paris
Honorary LL.D., Yonsei University, Seoul
American Academy of Arts and Sciences
National Academy of Sciences
Institute of Medicine
Vice-president, International Union of Physiological
Sciences
Albert Behnke Award, Undersea Medical Society
Honorary D.Sc., University of Rochester
Distinguished professor, State University of New York at
Buffalo
1976-77 Alexander von Humboldt Award and visiting professor,
University of Gottingen
Painton Award, Cooper Ornithological Society
Profesor honoraria, Universidad Peruana, Lima
Doctor of medicine, Honoris Causa, University of Berne
Elliott Cones Award, American Ornithological Union
Chancellor Norton Medal, State University of New York
at Buffalo
Dedication of the Hermann Rahn Laboratory for
Environmental Physiology, State University of New York
at Buffalo
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