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OCR for page 386
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OCR for page 387
PER FREDRIK THORKELSSON
SCHOLANDER
November 29, ~ 905-fune 13, ~ 980
BY KNUT SCHMIDT-NIELSEN
THE MOST IMPRESSIVE ASPECT of Scholanders scien-
tific life is his versatility as a biologist and his ability to
make significant contributions in a broad range of fields. He
was first of all a physiologist, but his work always signified a
fresh approach to broader biological problems ant! prin-
ciples. He had an ability to ask the right questions, to conceive
of simple experiments and (resign the necessary equipment,
to utilize novel approaches, and to present simple and logical
answers to important questions. As a person he was warm
and enthusiastic, generous ant! kind, and utterly uncon-
cernect with the hassles of form and bureaucracy.
Scholancler was born in Sweden and grew up in a family
of talent and culture. His father was an engineer; his mother,
Norwegian born, was an accomplished professional pianist;
his grandfather was a prominent architect, as well as a writer
anc! musician, and a professor at the Royal Academy of Arts.
Scholander has related that, as a small boy, he crawlecl un-
erneath his mother's Steinway granct piano when she prac-
ticed, smothered! by the waves of emotion in the music. He
himself became an accomplished violinist, which I first ctis-
coverec! when late one night ~ found him playing in the lab-
oratory to a record that omitted the first violin.
387
OCR for page 388
388
BIOGRAPHICAL MEMOIRS
Because of the divorce of his parents, Scholancler moved
to Norway, where in 1924 he matriculated at the Faculty of
Medicine of the University of OsIo (then Christiania). At that
time mectical students started with Latin, philosophy, chem-
istry, and physics, whereupon followocl years of preclinical
work in anatomy, physiology, and so on. Although Scholan-
der was terribly borer! by much of this, he continued his med-
ical studies and completed his medical education in 1932,
within the normal time span. Scholancler has described how
his textbooks were always full of marginal notes about expla-
nations he ctict not believe, but his grades clic! not reflect his
brilliance. There was a rumor among the university stu-
dents probably untrue that he completed his final medi-
cal examinations with the unique distinction of having ob-
tained the lowest gracles ever given a passing student.
Scholancler's years as a medical student became significant
in a very cti~erent ant! unexpected] way. Walking home from
the University one day, bored with tedious coursework, he
picked some lichens off the trees along the street. At home
he found a flora for lower plants ant] soon tract the lichens
identified. He continucct collecting, and when his flora
proved inadequate, he sought further help from the profes-
sor of botany, Bernt Lynge. Lynge recognizes! the unusual
talents of the young student, who rapidly became an out-
standing lichen specialist. Lynge himself, because of severe
arthritis, was unable to go to GreenIanc! and offered Scho-
lander the opportunity to take his place as a botanist on sev-
eral arctic expeditions. As a result, Scholancler spent three
summers (1930-1932) in Greenland and Spitzbergen. His
contributions to lichenology, particularly his revision of the
family Umbilicariaceae ~ ~ 934b), placer! him among the
worIct's foremost lichenologists at that time.
The young physician who was never to practice medi-
cine-tract become a botanist. He was approached by the pro
OCR for page 389
PER FREDRIK THORKELSSON SCHOLANDER 389
fessor of systematic botany, lens Holmboe, who suggester!
that he should present himself for a doctorate in botany, and
in 1934 he was awarded the Dr. Philos. by the University of
Oslo. It might be expected that his dissertation wouIcT deal
with lichens, but this would be uncharacteristic for Scholan-
der he was awarcled the doctorate basect on a monograph
of the vascular plants of Spitzbergen (1934a).
During the arctic expeditions, Scholancler had been in
trigucct by the many seals, polar bears, and diving birds he
saw. He clearly saw that many important questions needect
answers, such as "How do diving seals get enough oxygen?"
and "Why clon't they get divers' disease as humans do after
descending to similar depths?" He obtained working space in
the basement of the University Physiological Institute, and
with the aid of the professor of anatomy, the geneticist Otto
Mohr, he was awarded a small university research fellowship
that enabler! him to pursue his interests in physiology. He
clevelopect new methods for the continuous recording of the
respiratory metabolism of diving animals, and one of his pub-
lications (1937b) caught the attention of the distinguished
Danish physiologist August Krogh, who immediately under-
stoocT Scholancler's genius for design of experimental appa-
ratus. At that time ~ was a stuclent in (Copenhagen, and when
Scholancler came to-give a lecture on his studies of diving
animals, ~ sat there completely spelIbouncl by his brilliant
presentation and the simple and logical answers he provided
to questions that long tract puzzled physiologists who contem-
platecl the mysteries of diving physiology.
The most characteristic differences between a diving seal
ant! a human are strikingly simple. First, when a seal begins
a ctive, it exhales and clives with a minimal volume of air in
the lungs. It may seem counterproductive to dive with little
air in the lungs, but it greatly recluces the amount of nitrogen
taken up by the blooct and the tissues during the dive. This
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390
BIOGRAPHICAL MEMOIRS
seems to be the key factor in how seals and whales avoid
divers' disease, or the bencts, which is so tragically well known
from humans who ascend after dives to great depths. The
oxygen-carrying capacity of the bloocI, however, is much
greater in the seal than in humans. Its blood volume is rela-
tively larger, anct both blooct and muscles contain much
larger amounts of hemoglobin, ant! thus hold more oxygen,
than in mammals in general. A seal's most characteristic re-
sponse to an experimental dive is to slow the heart down to
a few beats per minute; the blood is diverted to the most vital
organs, notably the central nervous system and the eyes. The
muscles, which are able to function anaerobically through the
formation of lactic acid, receive no blooct ant! thus acquire
an oxygen debt that is repaid when oxygen is again available
at the termination of the dive. These important results were
. . . . . . . . .
summarized in a monograph (1940a) that remains the foun
dation for what we unclerstand today of the physiology of
diving animals.
At about the same time, Laurence Irving at Swarthmore
College had also clone distinguished work on diving physi-
ology, anct August Krogh arranger! for a Rockefeller fellow-
ship for Scholancler to continue his studies of diving physi-
ology at Swarthmore. Scholancler, however, was still in
Norway when the Second WorIct War broke out, and the
Rockefeller Foundation cancellecT all fellowship awards.
When Krogh was informer! about this, he sent an urgent
telegram to Scholancler that he should immediately leave for
the USA, and Scholancler obtained space on the last ship that
left Norway for the USA. Once lanclecI, he of course received
the cancellecI fellowship, and the studies of diving animals
were continued in collaboration with Irving.
However, as the Uniter] States became involved in the Sec-
oncl World War, Laurence Irving went to the Air Force, and
Scholancler followed him there. Both Irving and Scholancler
OCR for page 391
PER FREDRIK THORKELSSON SCHOLANDER 391
made substantial scientific and practical contributions during
their military service. Scholander became involved in the test-
ing of military equipment and survival gear. He developecl
simple and reliable field methods; for example, he stucTiecl
the conditions under which field stoves causer! carbon mon-
oxicle poisoning in tents and snow houses, he tried sleeping
bags under blizzard conditions, and tested coverer! life rafts
cluring storms in the Aleutian Islands.
be_ · ~ · · ~ 1
.
During this time he also arranged a quick and unauthor-
~zec! rescue mission to an airplane that tract crashed at the tip
of the Alaskan peninsula. One survivor had been seen, but
rescue teams had failed to reach the site over land and Scho-
lancler's camp commander hac! vetoed parachute jumps.
Going in by plane would be insubordination, but Scholancler
found a young pilot who, risking his career, volunteered. An-
other medical cloctor and a priestjoinecT them, ant! the three,
who had never iumpec! before, came clown successfully, al
.. . ..
1 ,
tnougn warn some difficulty. They found that two men had
perished, but three survivors were taken care of by the M.D.
while Scholancler took on the cooking, greatly helped by the
plane's cargo of sacramental wine for Passover, or "Hang-
over" as Scholander called it. Eventually the survivors and
the rescuers were taken out by a bush plane from Anchorage.
~ have later heard that Scholander, as a result of the suc-
cessfu! Aleutian rescue, was to be courtmartialed for insub-
orclination. As the case slowly moved through the military
channels, it became known to Detlev Bronk, who was an-
gered by the bureaucratic stupidity of punishing one of the
most elective scientists in the Air Force. By some unknown
intervention, the case was aborted at the highest level in
Washington.
After the years of military work, Irving and Scholancler,
with support from the Navy, established a research laboratory
at Point Barrow, Alaska. Among the most important results
OCR for page 392
392
BIOGRAPHICAL MEMOIRS
were measurements of the metabolic heat production ant! the
insulation neecled for warmblooded animals, mammals and
birds, to keep warm in the arctic climate. It turner! out that
larger animals, such as foxes and eskimo clogs, could sleep at
temperatures as low as - 30 or - 40°C without any increase
in heat production. Smaller animals, however, started to
shiver at temperatures well above freezing, and they neecled
increases! heat production to stay warm and maintain a nor-
mal body core temperature of 37°C. The information was in
agreement with the simple physical laws of heat exchange,
anti the studies clearly showed that below a certain temper-
ature, the lower critical temperature, the metabolic rate in-
creased linearly with the decrease in temperature. This
mode} for the responses of warmbloocled animals to low tem-
peratures has remained! the mocle} for virtually all later stud-
ies of this nature (1950c,d,e).
The arctic work incluclec] studies of coldblooded animals
and of plants, ant! also the tolerance of various organisms to
freezing. Scholancler devisect a simple and elegant method
for determining the amount of body water in a mosquito
larva that was actually frozen to ice inside the intact animal.
He showed that these, when they were thawed out, were still
alive and unharmed after more than 80 percent of their body
water hac! been frozen to ice (1953e).
Scholander's many ingenious methods for the analysis of
minute samples of gas became very helpful in an unexpected
area. He noticed that gas bubbles tendect to clevelop under
ice, and also that a chunk of ice chopper! from a glacier gave
off bubbles of gas when put into a drink to coo} it. Scholander
determinect that ice is virtually impermeable to gases; he
froze thin sheets of ice, 0. ! mm thick, and floated them on
cold mercury. He introduced small volumes of gases under
the ice and attempter] to measure changes in their composi-
tion. He was never able to detect any penetration, and con
OCR for page 393
PER FREDRIK THORKELSSON SCHOLANDER 393
cludect that ice at -10°C is at least 70,000 to 80,000 times
more impermeable to gases than a layer of water. This made
him wonder whether the air trapped creep in the Greenland
glaciers contains a permanent record of the composition of
the atmosphere in earlier periods (1956b,e; 195Sc,g; 1960c;
196Ic; 1962g).
After the two years of work at Barrow, Scholander left the
Arctic Research Laboratory. A. Bairc! Hastings, professor of
biological chemistry at Harvard University, was a person who
fully unclerstood Scholancler's genius, and he offered him a
position as research fellow. During two years in Hastings' lab-
oratory, Scholancler clevelope(1 a series of elegant micrometh-
o(ls for blood and gas analysis, and started applying these to
solving some of the intriguing problems of how gases are
secreted into the swimblacicler of fish. Depending on the
depth at which a fish is found, the gases in the swimblacicler
are uncler high pressure. Gases that are dissolvecI in the sur-
rounding seawater are at partial pressures close to those in
the atmosphere, and they are secreted into the swimblacIder
against pressures that in deep sea fish may amount to several
hundred atmospheres. The micromethocts Scholander hacI
clevelopect were ideal for these studies. He now demonstratect
that not only oxygen, but also nitrogen, is secreted against
tremendous concentration gradients. That this is possible for
oxygen, a relatively reactive gas, was perhaps not surprising,
but how couIct nitrogen, a presumably totally inert gas, also
be secreted actively against such concentration gradients?
The answer to both puzzles lay in the unusual vascular supply
to the swimblacIder and in physical principles that, once they
were understood, were simple to explain. The blood supply
to the swimbladder forms a counter-current system that is an
essential part of the secretion mechanism (1954a,b).
The swimbladcler studies were continued cluring the
three years of Scholancler's association with the Woods Hole
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394
BIOGRAPHICAL MEMOIRS
Oceanographic Institution as a physiologist ( 1952-19551.
The result was a series of papers that now form the foun-
dation for our unclerstancling of swimblaclcler function
955g, ~ 958f, and others).
In 1955 Scholander was callecI to the University of OsIo
as professor and director of a new Institute of Zoophysi-
ology. His three years in OsIo were very active. He continued
studies of the adaptation of humans to cold and answered
questions of how primitive humans can keep warm in cold
climates. Eskimoes, Lapps, and Australian aborigines were
comparer! with Norwegian students. The latter slept out-
doors at temperatures around 0°C, naked in a single-blanket
sleeping bag. For an unacclimatized! person this is intensely
uncomfortable and it is impossible to sleep, but after five or
six days of acclimatization the students had no trouble. The
most notable change in their physiology was a substantial in-
crease in metabolic heat production. (The motivation of the
students to go through with these uncomfortable experi-
ments seemed to be related to their being granted permission
to hunt reindeer.)
When the Australian aborigines were studiecI, it was
found that they could sleep through a coin night with a nor-
mal metabolic rate. The difference was that the aborigines
permitted the temperatures of the legs to ctrop to around
10°C. In other words, they had adapted by permitting their
peripheral temperatures to drop instead of increasing their
metabolic rate. This, of course, is a more economical ap-
proach; keeping the legs warm requires a substantial increase
in metabolic heat production.
After three highly productive years in Norway, Scholan-
cler was brought to the Scripps Institution of Oceanography
through the efforts of its director, the oceanographer Roger
Revelle, anc! he remained there until the enct of his life. With
tremendous enthusiasm he took up one problem after an-
other. Among these were some important problems of plant
OCR for page 395
PER FRED RI K TH O RKELS SO N S C H O LAN DER 395
physiology. The question of how water can be brought to the
top of the tallest trees had always intrigued him, and studies
that he hac! initiated while he was at Woods Hole and in OsIo
were continuccI ~ ~ 957a, ~ 958b, ~ 96 ~ d). The fact that suction,
in combination with the cohesion of water, can explain the
ascent of the sap lect to later studies of osmotic phenomena
in other plants, and eventually to the clevelopment of the
solvent tension theory of osmosis. Important in this connec-
tion was a series of studies of mangroves and their salt bal-
ance and the question of how they manage with their roots
in seawater. These studies started on the Cape York Peninsula
in northern Australia ~ ~ 962h) and continucct for several
years ~ ~ 965a,c; ~ 966a; ~ 96Sb).
The solvent tension theory of osmosis was founded on
Scholander's experience in plant physiology. A series of ele-
gant experiments, in part carried out in collaboration with
H. T. Hammel, also of Scripps, provided easily compre-
hendect support for Scholancler's theories ~197 Ic,e). The
theories were severely criticized by several physical chemists,
who based their arguments on classical thermodynamic ar-
guments. Scholander's ingenious experiments remained] un-
challenged; only their interpretation could be disputed.
Although Scholancler's view is simple and intuitive, it is un-
doubtedly more convenient to adhere to the traditional way
of thinking about osmotic phenomena. I can only regret that
I do not have the competence to evaluate the controversy and
give proper perspective to the significance of Scholancler's
contributions in this field.
A most important development during .~cholander's
. ~. . .
.
many years at the Scripps Institution was the design and
builcling of the research vessel Alpha Helix. During his many
field expeditions, Scholancler tract understood the impor-
tance of bringing adequate equipment into the field, where
the important physiological problems are evident. He con-
ceived of a laboratory ship that should be able to take a group
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396
BIOGRAPHICAL MEMOIRS
of a dozen or so scientists to any area of the worIcI. The ship
should be equipped with modern laboratories anti an excel-
lent machine shop. This was based on Scholander's own ex-
perience; he was a skilled machinist and this hacT helped him
during his many successes in designing new methods ant!
equipment. He saw the absolute necessity of always having at
his disposal a good machine shop that could solve problems
as they arise uncler field conditions.
Support for building the ship was obtained from the Na-
tional Science Foundation, and in 1966 the ship started on
its first cruise, the BilIabong Expedition to tile Great Barrier
Reef. A National Advisory Board, initially chaired by Baird
Hastings, evaluated applications for use of the ship. Over the
fourteen years the ship remained in service as a floating phys-
iological laboratory, expeditions went to Australia, the South
Seas, the Amazon, the Antarctic, the Galapagos, the Bering
Sea, and other sites. Several hundred scientists from all over
the work! have participated actively in these expeditions, and
the records of Scripps Institution show that the work on the
Alpha Helix has resulted! in a total of 547 publications in rec-
ognized scientific journals an impressive record for the rel-
atively modest funds invested in the Alpha Helix.
Not only through the Alpha Helix, but throughout his life,
Scholander became a seminal figure for physiologists who
were concerned with the problems that animals and plants
encounter in nature. He was an immensely enthusiastic per-
son, a true naturalist who perceived interesting scientific
problems wherever he moved. He lee! a restless ant! highly
productive life. A large number of scientists, now active anti
recognized arounc] the world, have been associated with
Scholancler and have been influenced by his stimulating and
dynamic personality. In 1951 he married the daughter of
Laurence Irving, Susan, who remained his devoted compan-
ion the remainder of his life.
OCR for page 403
PER FREDRI K TH ORKELS SON S C H O LAN DER 403
b. Accurate analysis of respiratory gases in 0.5 cubic centimeter
samples. Fed. Proc. Fed. Am. Soc. Exp. Biol., 6:197-98.
Analyser for accurate estimation of respiratory gases in one-
half cubic centimer samples. i. Biol. Chem., 167:235 -50.
d. Simple syringe burette. Science, 105:581.
With S. C. Flemister and L. Irving. Microgasometric estimation
of the blood gases. V. Combined carbon dioxide and oxygen. }.
Biol. Chem., 169: 173 -81.
f. With H. }. Evans. Microanalysis of fractions of a cubic milli-
meter of gas. I. Biol. Chem., 169:551- 60.
g. With L. Irving. Micro blood gas analysis in fractions of a cubic
millimeter of blood. i. Biol. Chem., 169:561-69.
c.
1949
Volumetric respirometer for aquatic animals. Rev. Sci. Instrum.,
20:885-87.
1950
a. Volumetric plastic micro respirometer. Rev. Sci. Instrum.q
21 :378-80.
b. With H. Niemeyer and C. L. Claff. Simple calibrator for War-
burg respirometers. Science, 112 :437-38.
c. With V. Walters, R. Hock, and L. Irving. Body insulation of
some arctic tropical mammals and birds. Biol. Bull. Woods
Hole, Mass., 99:225-36.
d. With R. Hock, V. Walters, and L. Irving. Heat regulation in
some arctic and tropical mammals and birds. Biol. Bull. Woods
Hole, Mass., 99:237-58.
e. With R. Hock, V. Walters, and L. Irving. Adaptation to cold in
arctic and tropical mammals and birds in relation to body tem-
perature, insulation, and basal metabolic rate. Biol. Bull. Woods
Hole, Mass., 99: 259-71.
1951
a. Nitrogen tension in the swimbladder of marine fishes in rela-
tion to the depth. Fed. Proc. Fed. Am. Soc. Exp. Biol., 10:121.
b. With C. L. Claff, C. T. Teng, and V. Walters. Nitrogen tension
in the swimbladder of marine fishes in relation to the depth.
Biol. Bull. Woods Hole, Mass., 101:178-93.
c. With H. Erikson and L. Irving. Apparatus for complete volu
OCR for page 404
404
BIOGRAPHICAL MEMOIRS
metric recording of the respiratory gaseous exchange in man.
Scand. i. Clin. Lab. Invest., 3:228-33.
1952
c'
a. With C. L. Claff, }. R. Andrews, and D. F. Wallach. Microvolu-
metric respirometry. I. Gen. Physiol., 35:375-95.
b. With C. L. Claff and S. L. Sveinsson. Oxygen consumption dur-
ing the cleavage of single cells. Fed. Proc. Fed. Am. Soc. Exp.
Biol.,11:141.
With C. L. Claff and S. L. Sveinsson. Respiratory studies of
single cells. I. Methods. Biol. Bull. Woods Hole, Mass., 102: 157-
77.
d. With C. L. Claff and S. L. Sveinsson. Respiratory studies of
single cells. II. Observations on the oxygen consumption in
single protozoans. Biol. Bull. Woods Hole, Mass., 102:178-84.
e. With C. L. Claff, S. L. Sveinsson, and Susan I. Scholander. Res-
piratory studies of single cells. III. Oxygen consumption during
cell division. Biol. Bull. Woods Hole, Mass., 102: 185-99.
With i. H. Kinoshita and I. P. Bunker. The use of the volumetric
respirometer in the determination of plasma carbon dioxide. I.
Lab. Clin. Med., 40:156-60.
g. With }. Wyman, fir., G. A. Edwards, and L. Irving. On the sta-
bility of gas bubbles in sea water. J. Mar. Res., 11 :47-62.
h. With W. Flagg, V. Walters, and L. Irving. Respiration in some
arctic and tropical lichens in relation to temperature. Am. I.
Bot., 39: 707-13.
i. With R. I. Hock, H. Erikson, W. Flagg, and L. Irving. Compo-
sition of the ground-level atmosphere at Point Barrow, Alaska.
i. Met., 9:441-42.
1953
a. With W. Flagg, V. Walters, and L. Irving. Climatic adaptation in
arctic and tropical poikilotherms. Physiol. Zool., 26:67-92.
b. With L. van Dam. Composition of the swimbladder gas in deep
sea fishes. Biol. Bull. Woods Hole, Mass., 104:75-86.
c. With L. van Dam. Concentration of hemoglobin in the blood of
deep sea fishes. }. Cell. Comp. Physiol., 41:522-24.
d. Studies on the physiology of frozen plants and animals in the
Arctic. Abstracts Communications, 19th Int. Physiol. Congr.,
pp. 741-42.
OCR for page 405
e.
PER FREDRIK THORKELSSON SCHOLANDER 405
With W. Flagg, R. J. Hock, and L. Irving. Studies on the phys-
iology of frozen plants and animals in the Arctic. }. Cell. Comp.
Physiol., 42, suppl. 1. 56 pp.
1954
a. With L. van Dam. Secretion of gases against high pressures in
the swimbladder of deep sea fishes. I. Oxygen dissociation in
blood. Biol. Bull. Woods Hole, Mass., 107:247-59.
b. Secretion of gases against high pressures in the swimbladder of
deep sea fishes. II. The rete mirabile. Biol. Bull. Woods Hole,
Mass., 107:260-77.
1955
a. With L. van Dam and Susan I. Scholander. Gas exchange in the
roots of mangroves. Am. }. Bot., 42 :92-98.
b. Evolution of climatic adaptation in homeotherms. Evolution,
9:15-26.
With W. E. Love and i. Id. Kanwisher. The rise of sap in tall
grapevines. Plant Physiol., 30:93-104.
d. With L. van Dam, C. L. Clam, and J. W. Kanwisher. Micro ga-
sometric determination of dissolved oxygen and nitrogen. Biol.
Bull. Woods Hole, Mass., 109:328-34.
e. With W. E. Schevill. Counter-current vascular heat exchange in
the fins of whales. J. Appl. Physiol., 8:279-82.
Hydrostatic pressure in coconuts. Plant Physiol., 30:560-61.
g. With L. van Dam and T. Enns. Secretion of inert gases and
oxygen by the swimbladder of fishes. Biol. Bull. Woods Hole,
Mass., 109:338-39.
h. With Alan C. Burton and Otto G. Edholm. Man in a cold en-
vironment. Physiological and pathological ejects of exposure
to low temperatures. Scand. }. Clin. Lab. Invest., 7:349.
c.
1956
a. With L. van Dam and T. Enns. Nitrogen secretion in the swim-
bladder of whitefish. Science, 123 :59-60.
b. With }. W. Kanwisher and D. C. Nutt. Gases in icebergs. Science,
123: 104-5.
c. With H. Erikson, }. Krog, and K. Lange Andersen. The critical
temperature in naked man. Acta Physiol. Scand., 37:35-39.
d. Climatic rules. Evolution, 10:339-40.
OCR for page 406
406
BIOGRAPHICAL MEMOIRS
With L. K. Coachman and E. Hemmingsen. Gas enclosures in
a temperate glacier. Tellus, 8:415 -23.
f. With L. van Dam and T. Enns. The source of oxygen secreted
into the swimbladder of cod. }. Cell. Comp. Physiol., 48:517-
22.
g. Observations on the gas gland in living fish. J. Cell. Comp. Phys-
iol., 48:523-28.
h. With L. van Dam. Micro gasometric determination of oxygen
in fish blood. }. Cell. Comp. Physiol., 48:529-32.
1957
a. With Berthe Ruud and H. Leivestad. The rise of sap in a trop-
ical liana. Plant Physiol., 32: 1-6.
b. With L. van Dam. The concentration of hemoglobin in some
cold water arctic fishes. }. Cell. Comp. Physiol., 49: 1-4.
c. With L. van Dam, i. W. Kanwisher, H. T. Hammel, and M. S.
Gordon. Supercooling and osmoregulation in arctic fish. J. Cell.
Comp. Physiol., 49:5 -24.
d. With K. Lange Andersen, }. Krog, F. Vogt Lorentzen, and }.
Steen. Critical temperature in Lapps. J. Appl. Physiol., 10:231-
34.
e. With H. T. Hammel, K. Lange Andersen, and Y. L0yning. Met-
abolic acclimation to cold in man. Fed. Proc. Fed. Am. Soc. Exp.
Biol., 16:114-15.
f. "The wonderful net." Sci. Am., 196~4) :96 -107.
g. With i. Krog. Countercurrent heat exchange and vascular
bundles in sloths. }. Appl. Physiol., 10:405-11.
h. With H. Leivestad and H. Andersen. Physiological response to
air exposure in codfish. Science, 126:505.
i. Oxygen dissociation curves in fish blood. Acta Physiol. Scand.,
41 :340-44.
J
k
.
With H. T. Andersen and H. Leivestad. "Air diving" in fishes.
Acta Physiol. Scand., 42, suppl. 145:6-7.
With H. T. Hammel, K. Lange Andersen, and Y. L0yning. Met-
abolic acclimation to cold in man. Acta Physiol. Scand., 42,
suppl. 145:63-64.
With }. Krog. Counter current vascular heat exchange, with
special reference to the arteriovenous bundles in sloths. Acta
Physiol. Scand., 42, suppl. 145: 89-90.
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PER FREDRIK THORKELS SON S C H O LAN DER 407
m. With G. Sundnes. Gas secretion in fishes lacking "rete mirabile."
Acta Physiol. Scand., 42, suppl. 145:125-26.
1958
a. With H. T. Hammel, K. Lange Andersen, and Y. L0yning. Met-
abolic acclimation to cold in man. i. Appl. Physiol., 12: 1-8.
b. The rise of sap in lianas. In: The Physiology of Forest Trees, ed.
Kenneth V. Thimann, pp. 3-17. New York: Ronald Press.
c. With L. K. Coachman, E. Hemmingsen, T. Enns, and H. de
Vries. Gases in glaciers. Science, 127:1288-89.
d. With H. Jensen. Bag spirometer. Scand. i. Clin. Lab. Invest.,
10:225-26.
e. With H. T. Hammel, I. S. Hart, D. H. LeMessurier, and I. Steen.
Cold adaptation in Australian aborigines. }. Appl. Physiol.,
13:211-18.
With G. Sundnes and T. Enns. Gas secretion in fishes lacking
rete mirabile. i. Exp. Biol., 35:671-76.
g. With L. K. Coachman and T. Enns. Gas loss from a temperate
glacier. Tellus, 10:493 -95.
h. Counter current exchange. A principle in biology. Hvalrad.
Skr., no. 44. 24 pp.
. . . .
i. Studies on man exposed to cold. Fed. Proc. Fed. Am. Soc. Exp.
Biol., 17: 1054 - 57.
With H. Leivestad and G. Sundnes. Cycling in the oxygen con
sumption of cleaving eggs. Exp. Cell. Res., 15:505-11.
k. With O. Iversen. New design of volumetric respirometer.
Scand. i. Clin. Lab. Invest., 10:429-31.
1959
a. Wave-riding dolphins: How do they do it? Science, 129:1085-
87.
b. Experimental studies on asphyxia in animals. In: Oxygen Supply
to the Human Foetus, ed. lames Walker and A. C. Turnbull, pp.
267-74. Oxford: Blackwell Scientific Publications.
c. Supercooling and freezing in poikilotherms. In: Simposios Con-
ferencias, 21st Congr. Int. Ciencias Fisiol., Buenos Aires, pp. 77-
81.
d. Wave-riding dolphins. Science, 130: 1658.
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408
b.
BIOGRAPHICAL MEMOIRS
1960
Oxygen transport through hemoglobin solutions. Science,
131:585-90.
Oxygen diffusion. Science, 132:368.
With D. C. Nutt. Bubble pressure in Greenland icebergs. I. Gla-
ciol., 3:671-78.
d. With E. Hemmingsen. Specific transport of oxygen through
hemoglobin solutions. Science, 132: 1379-81.
Man in cold environment. Discussion. Fed. Proc. Fed. Am. Soc.
Exp. Biol., 19, suppl. 5: 8-10, 11-12.
1961
a. With H. T. Hammel, D. H. LeMessurier, E. Hemmingsen, and
W. Garey. Circulatory adjustment in pearl divers. Fed. Proc.
Fed. Am. Soc. Exp. Biol., 20:103.
b. With D. C. Nutt and L. K. Coachman. Dissolved nitrogen in
West Greenland waters. i. Mar. Res., 10:6-11.
With E. A. Hemmingsen, L. K. Coachman, and D. C. Nutt.
Composition of gas bubbles in Greenland icebergs. }. Glaciol.,
~,
3:813-22.
d. With E. Hemmingsen and W. Garey. Cohesive lift of sap in the
rattan vine. Science, 134: 1835 -38.
1962
a. With M. S. Gordon and B. H. Amdur. Freezing resistance in
some northern fishes. Biol. Bull. Woods Hole, Mass., 122:52-
62.
With H. T. Hammel, D. H. LeMessurier, E. Hemmingsen, and
W. Garey. Circulatory adjustment in pearl divers. J. Appl. Phys-
iol., 17: 184-90.
c. With C. R. Olsen and D. D. Fanestil. Some effects of breath
holding and apneic underwater diving on cardiac rhythm in
man. J. Appl. Physiol., 17:461-66.
d. With Edda Bradstreet. Microdetermination of lactic acid in
blood and tissues. }. Lab. Clin. Med., 60:164-66.
e. With Edda Bradstreet and W. F. Garey. Lactic acid response in
the grunion. Comp. Biochem. Physiol., 6:201-3.
With R. W. Elsner and E. Hemmingsen. The work of maintain-
ing flotation in sea water. Physiologist, 5: 136.
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PER FREDRIK THORKELSSON SCHOLANDER 409
g. With W. Dansgaard, D. C. Nutt, H. de Vries, L. K. Coachman,
and E. Hemmingsen. Radio-carbon age and oxygen-18 content
of Greenland icebergs. Medd. Gronl., 165(1). 26 pp.
h. With H. T. Hammel, E. Hemmingsen, and W. Garey. Salt bal
1.
ance in mangroves. Plant Physiol., 37:722-29.
With C. R. Olsen and D. D. Fanestil. Some effects of apneic
underwater diving on blood gases, lactate, and pressure in man.
J. Appl. Physiol., 17 :938-42.
i. Physiological adaption to diving in animals and man. Harvey
Lect., 57:93 -110.
1963
a. With R. W. Elsner and W. F. Garey. Selective ischemia in diving
man. Am. Heart }., 65:571-72.
b. The master switch of life. Sci. Am., 209~6~:92-106.
1964
Animals in aquatic environments: Diving mammals and birds.
In: Handbook of Physiology. Sect. 4: adaptation to the environment,
ed. D. B. Dill, pp. 729-39. Bethesda, Md.: American Physio-
logical Society.
b. With T. Enns. Oxygen transport by hemoglobin collision. Fed.
Proc. Fed. Am. Soc. Exp. Biol., 23:468.
c. With H. T. Hammel, E. A. Hemmingsen, and Edda D. Brad-
street. Hydrostatic pressure and osmotic potential in leaves of
mangroves and some other plants. Proc. Natl. Acad. Sci., USA,
52:119-25.
d. With R. W. Elsner, A. B. Craig, E. G. Dimond, L. Irving, M.
Pilson, K. Johansen, and Edda Bradstreet. A venous blood ox-
ygen reservoir in the diving elephant seal. Physiologist, 7:124.
From the frozen forest to tropical mangroves. In: Program Ab-
stracts, Pt. 2, Proc. Alaska Sci. Conf., p. 38. Washington, D.C.:
American Association for the Advancement of Science.
1965
a. With H. T. Hammel, Edda D. Bradstreet, and E. A. Hemming
sen. Sap pressure in vascular plants. Science, 148:339 - 46.
b. With T. Enns and E. D. Bradstreet. Effect of hydrostatic pres
sure on gases dissolved in water. t. Phys. Chem., 69:389-91.
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410
BIOGRAPHICAL MEMOIRS
c. With H. T. Hammel, E. D. Bradstreet, and E. A. Hemmingsen.
Sap pressure in plants. Science, 149:920 - 21.
d. Tension gradients accompanying accelerated oxygen transport
in a membrane. Science, 149:876-77.
e.
Reverse osmosis and sap pressure in vascular plants. Science,
150:384.
f. With R. Elsner. Circulatory adaptations to diving in animals and
Nan. In: Physiology of Breath-Hold Diving and the Ama of Japan,
pp. 281-94. Washington, D.C.: National Academy of Sciences
Publ. 131.
1966
a. With E. D. Bradstreet, H. T. Hammel, and E. A. Hemmingsen.
Sap concentrations in halophytes and some other plants. Plant
Physiol., 41:529-32.
b. The role of solvent pressure in osmotic systems. Proc. Natl.
Acad. Sci. USA, 55:1407-14.
1967
a. Osmotic mechanism and negative pressure. Science, 156:67-
69.
b. Negative pressure in plants and osmotic mechanism. Science,
156:541.
c. Osmotic pressure. Science, 158: 1212.
d. With T. Enns and E. Douglas. Role of the swimbladder rete of
fish in secretion of inert gas and oxygen. Adv. Biol. Med. Phys.,
2:231-44.
1968
a. With R. W. Elsner. A comparative view of cardiovascular de-
fense against acute asphyxia. Proc. 2nd Int. Symp. Emergency
Resuscitation, Oslo. Acta Anaesthesiol. Scand., suppl. 29:15-
33.
b. How mangroves desalinate seawater. Physiol. Plant,21:251-61.
c. With A. R. Hargens and S. L. Miller. Negative pressure in the
interstitial fluid of animals. Science, 161:321-28.
d. With R. S. Bandurski and E. Bradstreet. Metabolic changes in
the mud-skipper during asphyxia or exercise. Comp. Biochem.
Physiol., 24:271-74.
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PER FREDRIK THORKELSSON SCHOLANDER 411
e. With M. de Oliveira Perez. Sap tension in flooded trees and
bushes -of the Amazon. Plant Physiol., 43: 1870-73.
1969
a. With L. Irving, E. A. Hemmingsen, and E. Bradstreet. Ultra-
violet absorption in the cornea of arctic and alpine animals. In:
The Biologic Elects of Ultraviolet Radiation, ed. F. Urbach, pp.
469-71. London: Pergamon Press.
b. With S. B. Stromme and i. E. Maggert. Interstitial fluid pres-
sure in terrestrial and semiterrestrial animals. I. Appl. Physiol.,
27: 123-26.
c. With A. R. Hargens. Stretch mounting for osmotic membranes.
Microvasc. Res., 1 :417-19.
1971
a. State of water in osmotic processes. Microvasc. Res., 3:215-32.
b. Imbibition and osmosis in plants. In: Topics in the Study oiLife:
The Bio Source Book, ed. Amy Kramer, pp. 138-47. New York:
Harper & Row.
With M. Perez. Experiments on osmosis with magnetic fluid.
Proc. Natl. Acad. Sci. USA, 68:1093-94.
d. With i. E. Maggert. Supercooling and ice propagation in blood
from arctic fishes. Cryobiology, 8:371-74.
With M. Perez. Effect of gravity on osmotic equilibria. Proc.
Natl. Acad. Sci. USA, 68:1569-71.
c.
1972
a. With M. Perez. Molecular buoyancy and osmotic equilibrium.
Proc. Natl. Acad. Sci. USA, 69:301-2.
b. Tensile water. Am. Sci., 60:584-90.
1973
With H. T. Hammel. Thermal motion and forced migration of
colloidal particles generate hydrostatic pressure in solvent.
Proc. Natl. Acad. Sci. USA, 70:124-28.
1975
Water states and water gates in osmotic processes, and the inop-
erative concept of molfraction of water. l. Exp. Zool., 194:241-
48.
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412 BIOGRAPHICAL MEMOIRS
1976
With H. T. Hammel. Osmosis and Tensile Solvent. Berlin: Springer
Verlag.
1978
c.
a. Rhapsody in science. Annul Rev. Physiol., 40:1-17.
b. Water under tension, its fundamental role in capillarity, osmosis
and colligative properties. In: Frontiers Or Human Knowledge,
Skrifter rorande Uppsala universitet, C:38, Acta Univ. Ups.
Nova Acta Regiae Soc. Sci. Ups. Ser. VC, pp. 297-308.
With A. R. Hargens and W. L. Orris. Positive tissue fluid pres-
sure in the feet of antarctic birds. Microvasc. Res., 15:239-44.
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Representative terms from entire chapter:
thorkelsson scholander
