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CONRAD ARNOLD ELVEH]EM
May 27, 1901 - July 27, 1962
BY R. H. BURRIS, C. A. BAUMANN,
AND VAN R. POTTER
THE WORK OF CONRAD ELVEH]EM a major contributor
to the golden era of nutritional research touched most
aspects of animal nutrition, advancing, in particular, our
unclerstanding of the B vitamins, the phenomenon of amino
acid imbalance, and the identification of trace minerals
needed in the diet. EIvehjem macle the major discovery that
nicotinic acid functions as the antipelIagra vitamin.
EIvehjem was also a superb administrator, an efficient
man who channeled his great energy with seemingly little
effort. On the local scene he served the University of Wiscon-
sin as chairman of the Department of Biochemistry, clean of
the Graduate School, and, finally, as president of the Univer-
sity. On the national level he helped make policy decisions
concerning the level of vitamins and other nutrients required
for health. The implementation of his cure for pelIagra was
· . .
1nternatlona In scope.
EARLY YEARS
Conrac! EIvehjem was born in 1901 to Ole Johnson
EIvehjem and Christine Lewis EIvehjem on a moclest farm
near McFarland, Wisconsin. In this primarily Norwegian
area (May 17 is still celebrated as Norwegian Inclependence
Day in Stoughton), EIvehjem grew up and attendee! high
135
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136
BIOGRAPHICAL MEMOIRS
school. He would spend his adult life within a few miles of
his birth place, for Madison's capitol building is visible from
the farm.
The EIvehjem children were expected to do their share
of the farm chores, and while there was little time for non-
sense education was encouraged. In those clays Wisconsin
farm boys usually did not go to high school and college, but
his family macle sure he was able to clo so. In 1919, EIvehjem
enrolled in the University of Wisconsin's College of Agricul-
ture, aIreacly recognized for its research in agricultural chem-
istry, genetics, plant pathology, and bacteriology. EIvehjem
majored in agricultural chemistry, a field] in which Babcock,
Hart, Steenbock, McCollum, and Peterson tract all done, or
were doing, meritorious work at Madison. He did his under-
graduate research under the direction of Harry Steenbock
and wrote his senior thesis jointly with W. P. Elmslie on "buck-
wheat itch," a light-inducec! disturbance in animals.
As to EIvehjem's early motivation in the choice of his ca-
reer, in 1957 he answered a thirteen-year-old boy who had
questioned him on this subject as follows: "l chose the field
of biochemistry because as a youngster I was interested in
what made plants grow and clevelop. ~ was very intrigued by
the rapic! growth of the corn plant, and ~ was interested in
knowing what reactions took place within the plant to allow
such rapid growth." Of his achievements, he said: "My
achievements cover work on many of the B vitamins in-
clucling the isolation and identification of nicotinic acic! as the
antipelIagra factor, also work on a number of trace mineral
elements showing that they have specific functions in nutri-
tion and metabolism. ~ also pioneered in work demonstrating
the relationship between vitamins and enzymes. Today ~ am
more interested in amino acids in nutrition." It never oc-
curred to him, apparently, to mention his many aclministra-
tive successes!
In 1923 EIvehjem began graduate work as a teaching as-
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CONRAD ARNOLD ELVEH]EM
137
sistant uncler Professor E. B. Hart, his major professor until
he received his Ph.D. degree in 1927. In 1924 he published
his first paper with Hart ant! Steenbock on dietary factors
influencing calcium metabolism (1924,1~. But his graduate
work centered mainly on iron deficiency in rats, including a
demonstration that copper must accompany iron in the diet
to cure this type of anemia.
Hart's encouragement of EIvehjem cluring his student
days is just one example of his remarkable capacity to pick
winners. This was before the perioc! when talented students
were being attractec! to agricultural chemistry in large num-
bers, yet Hart had staffed his small department with a
remarkable group of investigators. He supported them
through administrative difficulties, had a building con-
structed for their teaching and research, and offerect them
whatever he could given the limited resources available at
that time. As long as Hart lived, he ant! EIvehjem worked
together on many joint research projects. Indeed, approxi-
mately half of EIvehjem's long list of publications contains
Hart's name as well.
On June 30, 1926, EIvehjem married Constance Waltz, a
journalism student at the University of Wisconsin and the
daughter of a Rockford, Illinois, (lentist. This was a happy
union, and the two Connies- called Mr. Connie and Mrs.
Connie by their friends complemented each other. He was
relatively quiet, while she bubbled with enthusiasm, meeting
people easily with charm and grace. She was a source of
strength to her husband throughout all stages of his career,
and most particularly when he held administrative positions.
From 1927 to 1929, after receiving his Ph.D. degree,
EIvehjem held an instructorship in agricultural biochemistry.
In 1929, he received a National Research Council Fellowship
to study in the biochemistry laboratories at Cambridge Uni-
versity, England.
This was the only substantial period in his career that
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138
BIOGRAPHICAL MEMOIRS
EIvehjem spent away from Madison, ant! he ant} Mrs. Connie
took full advantage of it. As EIvehjem himself described it,
they arrived in England, took a guided tour through Lonclon
that allowed him to spot the laboratories he wanted to visit
later, searched for housing in Cambridge, anct met Dr. ant!
Mrs. C. G. King kindrec! souls with whom they would share
many experiences. According to EIvehjem, the Biochemistry
Department at Cambridge was a lively spot in 1929 ant! 1930,
and he describes Sir Frederick Hopkins' lively welcome back
as the recently announced recipient of the Nobel Prize.
At Cambridge EIvehjem worked under the tutelage of Dr.
David Keilin, who was then busy with the cytochromes ant!
the role of iron in cytochrome c. Copper and iron in cyto-
chrome oxidase were of particular interest to EIvehjem,
whose own work had shown that animals deficient in copper
were also deficient in cytochrome oxidase. By the 1930s, a
role for copper in cytochrome oxidase was widely accepted.
At Wisconsin EIvehjem had studied nutritional anemia in
rats on a diet very low in iron (viz., milk). The aciclition of
relatively large amounts of inorganic iron salts to such milk
failed to prevent this type of anemia. Testing crude materials
protective against anemia and later the ash of those most
potent for supplementing iron in a milk diet EIvehjem,
Steenbock, Hart, and Waddell found that traces of inorganic
copper were necessary for the incorporation of iron into he-
mogIobin, even though hemoglobin contains no copper. In
this way, the idea of catalysis in life-processes was brought
forcibly to EIvehjem's attention.
EIvehjem later published two papers on his work at the
Biochemical Laboratory in Cambridge with acknowledg-
ments to Hopkins "for his interest and advice" and to Keilin
"for many helpful suggestions." The first, "Factors Affecting
the Catalytic Action of Copper in the Oxidation of Cysteine"
(1930,1), clearly derives! from a project suggester! by Keilin.
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CONRAD ARNOLD ELVEHJEM
139
The second, "The Role of Iron and Copper in the Growth
arid Metabolism of Yeast" (193l,1), contained observations
that, though yeast contains no hemoglobin, its respiration
· · · · · .
requires 1ron-contalnlng pigments; anc t eat copper Increases
the levels of cytochrome a, presumably cytochrome oxidase.
Both studies gave EIvehjem experience with manometric
techniques, and he spent ~ busy year visiting laboratories,
cloing research on several problems, ant! aiding in a labora-
tory course.
RETURN TO WISCONSIN
At Cambridge, EIvehjem in the forefront of nutrition
research—worrier! less about finding new vitamins than
about understanding how these substances functioned in the
metabolism of the living cell. He clevelopec! a new research
strategy parallel studies on respiratory enzymes and on de-
ficiency-producing diets, especially clesignec! to be assayed
for new growth factors and trace elements. This new meth-
odology would further allow him to isolate the new sub-
stances and determine their action. He was, therefore,
particularly intrigued by the Barcroft respirometer. This in-
strument permitted accurate measurements of oxiciative en-
zymatic activity with small samples of tissue, enabling re-
searchers to clefine differences in the responses of normal
versus deficient tissue and the responses of deficient tissue to
addec! compounds.
While he was away, EIvehjem maintained a correspon-
dence with Hart, and their exchange of letters concerning
salary is interesting. On April 23, 1930, Hart wrote EIvehjem:
"I understood today that the Board of Regents had passed the budget
which appoints you as an Assistant Professor at $3,000 for the academic
year. You ought to be very happy over this because it was very difficult to
get an increment of $600 for you in the present state of Wisconsin finances.
You are young, and with summer pay and gradual increments, and an
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140
BIOGRAPHICAL MEMOIRS
opportunity for research the position you will hold with us ought to be
very attractive."
EIvehjem to Hart, May ~ 6, ~ 930:
"I was glad to have your letter and to learn that there would be a job
waiting for me when I return.... I can't say that I am exceedingly happy
over the salary but we can talk about that later. What I am wondering
about now is, if you will buy a Barcroft for me. If we are going to continue
to work on the minor inorganic elements it will come in very handy. In
fact there are a thousand things to do in regard to the catalytic action of
copper before leaving it in favor of other elements."
Hart agreed to let EIvehjem purchase his crevice, and he
brought a set of respirometers back with him on his return
to Wisconsin. In Madison it soon became a treasured pos-
session, ant! each noninterchangeable flask was carefully
guarded. The Potter-EIvehjem homogenizer remains still to
remind investigators of the days when EIvehjem was actively
studying respiratory enzymes. EIvehjem immediately put his
Barcroft respirometer to good use stuclying the respiration
of minced tissues from normal and from vitamin-deficient
experimental animals. He also continued his joint researches
with Professor Hart and a number of students on the mineral
requirements zinc, manganese, and molybdenum in the
rat, chicken, clog, and pig and began a large program on the
vitamin B complex, a relatively neglected area at Wisconsin
at that time.
.
In the early 1930s techniques available for nutritional
studies left much to be desire(l. Deficient diets were usually
lacking to varying degrees in more than one essential, and
curative preparations contained a number of clifferent vita-
mins. A typically crude (but useful) method! of producing
deficiencies was to damage a mixed diet with ctry or moist
heat, destroying vitamins differentially. The sources used for
growth factors were yeast, milk, liver, or fractions of liver left
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CONRAD ARNOLD ELVEHJEM
141
over from the commercial preparation of extracts for the
treatment of pernicious anemia. "Success" meant restoration
of the growth rate—by means of a supplemented diet that
had decreased on a defective diet.
EIvehjem's approach was similar to that of others working
on the B vitamins except that his graduate students worked
simultaneously on different growth factors or with different
species, so that when one achieved a preparation active
against his particular deficiency, others could test a similar
preparation for those deficiencies that were their own pri-
mary concern. This insured quick determination of the ef-
fects of a given concentrate on the various deficiencies under
study.
NICOTINIC ACID
Elvehjem was particularly skillful in coordinating experi-
ments and cross-checking results, ant! he was never timid
about postulating the existence of new growth factors. One
of these, "Factor W." represented what, in addition to the
established B vitamins, remained in a liver concentrate. His
recognition of nicotinic acid as the antipellagra principle was
typical of his thoroughness ant! his ability to combine infor-
mation gleaner! from various sources, with data produced by
his own students, and of his active collaboration with aca-
demic ant! commercial colleagues.
In 1912, exactly twenty-five years before nicotinic acid's
true status as a vitamin was established, Casmir Funk in one
of the more curious twists of nutritional history—attempted
to cure a vitamin deficiency by feeding it to polyneuritic
bircis. The results were unexciting. The substance came into
its own as an important biochemical, however, in 1936, when
Warburg and Christian iclentified it as one of the components
of "coferment" (NADP). Discovery of its presence in cozy-
mase (NAD) follower! quickly. About the same time, several
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142
BIOGRAPHICAL MEMOIRS
investigators reported it to be essential for the growth of cer-
tain microorganisms. EIvehjem and Douglas Frost, feeding
nicotinic acid to rats cleficient in "Factor W." reported a slight
growth response, though much less than that obtained with
crude liver preparations.
The isolation of nicotinic acid came about primarily
through the fractionation of liver extracts. By means of suc-
cessive solvent extractions, Car! Koehn had converter! 400
grams of liver extract to 2.5 grams of a powder active against
canine black tongue. Robert Madden achieved further con-
centrations by means of adsorption on an appropriate char-
coal. EIveh~em had for some time been receiving liver ex-
tracts for these studies from the Wilson and Abbott
Laboratories. Then Dr. Rhodehamal of the Eli Lilly Com-
pany, working according to the Koehn and EIvehjem proce-
dure, furnished a concentrate from seventeen kilograms of
liver. The next big step was Frank Strong's sublimation of
this concentrate in a molecular still. Almost immediately
Wayne Woolley obtained crystals from the distillate and, on
Karl Link's microapparatus, H. Campbell determined the
percentages of C, H. and N.
The response to these crystals in deficient dogs was dra-
matic, ant! the correlation between the analytical values and
the theory for nicotinamide was close enough to lead Woolley
to take a mixed melting point and perform the appropriate
characterization reactions all in a matter of a few days. Syn-
thetic nicotinic acid and amide were then fed to other clogs
and found to be highly active.
The research community lost little time in applying these
results to human pelIagra. EIvehjem's first published notice
of his laboratory's findings appeared in September 1937, in
a "letter to the editor" of the Journal of the American Chemical
Society. Van Potter, who shared an office with him at the time,
recalls that EIvehjem sent telegrams to a number of clinical
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CONRAD ARNOLD ELVEH]EM
143
investigators interested in pellagra including Tom Spies.
Before the end of 1937, EIvehjem's results with dogs had
been confirmed by six independent investigators. By the time
his more complete paper on the subject appeared in 193S, it
was possible to add the following: "Spies has used nicotinic
acid in four cases of classical pelIagra and reports (personal
communication) that the fiery red color associated with
pelIagrous dermatitis, stomatitis, and vaginitis improved
promptly." The Wisconsin paper (193S,1) not only summa-
rized the known biochemical facts on nicotinic acid, it even
expressed concern about possible toxicity in its anolication!i
.
~ 1 1
THE B VITAMINS AND AMINO ACIDS
Nicotinic acid, however, was not the only B vitamin to
occupy EIvehjem and his research team. As his list of publ'-
cations shows, his laboratory investigated every B vitamin at
one time or another, though occasionally under a different
name until its true nature was established. The clarification
and disentanglement of the B complex occupied many in-
vestigators worldwide for years, during which the EIvehjem
group made substantial contributions to our present under-
standing. But the latter years of his laboratory career were
spent on amino acids, an interest that had grown out of the
pelIagra problem.
PelIagra occurred in areas where people consumed inad-
equate diets high in corn, and EIvehjem's studies on black
tongue in dogs also involves! a diet high in corn. The diets
used for studies of the B complex in rats and chicks, on the
~ Because of Elvehjem's generosity in disseminating his laboratory's findings
widely, the medical implications of nicotinic acid in the treatment of pellagra became
apparent almost immediately. On January 22, 1938, Tom Spies's November 5, 1937,
report of his own experiments to the Central Society for Clinical Research was the
subject of an editorial in the Journal of the American Medical Association. For his dra-
matically successful use of nicotinic acid to treat pellagra in humans, Time magazine
named Spies 1938's "Man of the Year."
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44
BIOGRAPHICAL MEMOIRS
other hand, were so-called "semisynthetic" usually basest on
casein, starch, sugar, etc. Rats fed this diet never developecI
nicotinic acid deficiency, nor die! administration of nicotinic
acid improve their growth. But when corn was used to re-
place forty percent of such a diet, growth was depressed and
could be restored by supplements of nicotinic acid or of tryp-
tophan—an amino acid that is relatively lacking in corn.
Further studies in a number of laboratories clarified the
mechanism by which tryptophan is converter! to niacin in the
belly. Working with A. E. Harper, EIvehjem carried out ex-
periments on requirements for other amino acids that pre-
sagec3 an extensive investigation of amino acid imbalance-
an investigation that ceased, however, when he became pres-
ident of the University.
The coenzyme connection to nicotinic acid (NAD ant!
NADP) was important in motivating EIvehjem and Thorfin
Hogness, of The University of Chicago, to organize a "Sym-
posium on Respiratory Enzymes" in Madison on September
Il-13, 1941, ant! one on "The Biological Action of the Vi-
tamins," held at The University of Chicago on September 15-
19. David H. Smith noted at the vitamin symposium that
EIvehjem's observations on the relation of nicotinic acid to
canine black tongue (published in the September issue of the
Journal of the American Chemical Society ~ ~ 937,4] ~ were verified
promptly and extended to human pelIagra by a number of
· .
Investigators.
Subsequent to the spectacular conquest of pelIagra,
EIvehjem was invited to Cornell University Medical School to
be interviewed for the chairmanship of the Department of
Biochemistry the only position outside of the University of
Wisconsin he ever considered. A moclest and humble man,
EIvehjem had simple tastes and more than a touch of auster-
ity. Potter recalls his dismay on entering their shared office
one Saturday shortly after the Cornell trip to find EIvehjem
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CONRAD ARNOLD ELVEH]EM
157
With R. I. Madden, F. M. Strong, and D. W. Woolley. Relation of
nicotinic acid and nicotinic acid amide to canine black tongue.
I. Am. Chem. Soc., 59:1767.
1938
With R. ]. Madden, F. M. Strong, and D. W. Woolley. The isolation
and identification of the anti-black tongue factor. I. Biol.
Chem., 123:137.
With M. A. Lipschitz and V. R. Potter. The relation of vitamin Be
to cocarboxylase. Biochem. J., 32:474.
With P. L. Pavcek and W. H. Peterson. Factors affecting the vitamin
B. content of yeast. Ind. Eng. Chem., 30:802.
With D. W. Woolley, F. M. Strong, and R. I. Madden. Anti-black
tongue activity of various pyridine derivatives. I. Biol. Chem.,
124:715.
With F. M. Strong and R. ]. Madden. The ineffectiveness of ,B-
aminopyridine in black tongue. I. Am. Chem. Soc., 60:2564.
With D. W. Woolley, H. A. Waisman, and O. Mickelsen. Some ob-
servations on the chick antidermatitis factor. I. Biol. Chem.,
125:715.
With V. R. Potter and E. B. Hart. Anemia studies with dogs. J. Biol.
Chem., 126:155.
1939
With I. I. Oleson, H. R. Bird, and E. B. Hart. Additional nutritional
factors required by the rat. J. Biol. Chem., 127:23
The vitamin B complex in practical nutrition. I. Am. Diet. Assoc.,
15:6.
With A. E. Axelrod. Effect of nicotinic acid deficiency on the co-
zymase content of tissues. Nature, 143:281.
With D. W. Woolley and H. A. Waisman. Nature and partial syn-
thesis of the chick antidermatitis factor. J. Am. Chem. Soc.,
61:977.
With A. Arnold. Influence of the composition of the diet on the
thiamin requirement of dogs. Am. J. Physiol., 126:289.
With D. W. Woolley and H. A. Waisman. Studies on the structure
of the chick antidermatitis factor. J. Biol. Chem., 129:673.
With A. E. Axelrod and R. J. Madden. The effect of a nicotinic acid
deficiency upon the coenzyme I content of animal tissues. I.
Biol. Chem., 131:85.
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158
BIOGRAPHICAL MEMOIRS
With J. M. McKibbin, R. J. Madden, and S. Black. The importance
of vitamin B6 and factor W in the nutrition of dogs. Am. J.
Physiol., 128: 102.
With H. D. Anderson and J. E. Gonce, Jr. Vitamin E deficiency in
dogs. Proc. Soc. Exp. Biol. Med., 42:750.
1940
With D. V. Frost, V. R. Potter, and E. B. Hart. Iron and copper
versus liver in treatment of hemorrhagic anemia in dogs on
milk diets. J. Nutr., 19:207.
With M. A. Lipton. Mechanism of the enzymatic phosphorylation
of thiamin. Nature, 145:226.
With E. J. Schantz and E. B. Hart. The comparative nutritive value
of butter fat and certain vegetable oils. J. Dairy Sci., 23: 181.
With A. E. Axelrod and E. S. Gordon. The relationship of the di-
etary intake of nicotinic acid to the coenzyme I content of blood.
Am. J. Med. Sci., 199:697.
With D. M. Hegsted, J.J. Oleson, and E. B. Hart. The essential
nature of a new growth factor and vitamin B6 for chicks. Poult.
Sci., 19:167.
With H. A. Sober and M. A. Lipton. The relation of thiamine to
citric acid metabolism. J. Biol. Chem., 134:605.
With E. Hove and E. B. Hart. The relation of zinc to carbonic
anhydrase. J. Biol. Chem., 136:425.
With M. I. Wegner, A. N. Booth, and E. B. Hart. Rumen synthesis
of the vitamin B complex. Proc. Soc. Exp. Biol. Med., 45:769.
1941
With L. W. Wachtal, E. Hove, and E. B. Hart. Blood uric acid and
liver uricase of zinc-deficient rats on various diets. }. Biol.
Chem., 138:361.
With D. M. Hegsted, R. C. Mills, and E. B. Hart. Choline in the
nutrition of chicks. l. Biol. Chem., 138 :459.
With T. W. Conger. The biological estimation of pyridoxine (vita-
min B61. J. Biol. Chem., 138:555.
With H. A. Waisman. Chemical estimation of nicotinic acid and
vitamin B6. Ind. Eng. Chem., 13:221.
With M. I. Wegner, A. N. Booth, and E. B. Hart. Rumen synthesis
OCR for page 159
CONRAD ARNOLD ELVEH]EM
159
of the vitamin B complex on natural rations. Proc. Soc. Exp.
Biol. Med., 47:90.
With S. Black and I. M. McKibbin. Use of sulfaguanidine in nutri-
tion experiments. Proc. Soc. Exp. Biol. Med., 47:308.
With E. Nielsen. Cure of spectacle eye condition in rats with biotin
concentrates. Proc. Soc. Exp. Biol. Med., 48:349.
1942
With L. M. Henderson, I. M. McIntire, and H. A. Waisman. Pan-
tothenic acid in the nutrition of the rat. }. Nutr., 23:47.
With A. E. Axelrod and V. R. Potter. The succinoxidase system in
riboflavin-deficient rats. I. Biol. Chem., 142:85.
With D. M. Hegsted, R. C. Mills, G. M. Briggs, and E. B. Hart.
Biotin in chick nutrition. l. Nutr., 23:175.
With L. J. Teply and F. M. Strong. Nicotinic acid, pantothenic acid
and pyridoxine in wheat and wheat products. I. Nutr., 24: 167.
With S. Black, R. S. Overman, and K. P. Link. The effect of sulfa-
guanidine on rat growth and plasma prothrombin. I. Biol.
Chem., 145:137.
With D. Orsini and H. A. Waisman. Effect of vitamin deficiencies
on basal metabolism and respiratory quotient in rats. Proc. Soc.
Exp. Biol. Med., 51:99.
With I. D. Teresi and E. B. Hart. Molybdenum in the nutrition of
the rat. Am. I. Physiol., 137:504.
1943
With G. M. Briggs, fir., T. D. Luckey, R. C. Mills, and E. B. Hart.
Effect of p-aminobenzoic acid when added to purified chick
diets deficient in unknown vitamins. Proc. Soc. Exp. Biol. Med.,
52:7.
With O. K. Gant, B. Ransone, and E. McCoy. Intestinal flora of
rats on purified diets containing sulfonamides. Proc. Soc. Exp.
Biol. Med., 52:276.
With I. B. Field and C. Juday. A study of the blood constituents of
carp and trout. I Biol. Chem., 148:261.
With H. A. Waisman, A. F. Rasmussen, fir., and P. F. Clark. Studies
on the nutritional requirements of the rhesus monkey. l. Nutr.,
26:205.
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160
BIOGRAPHICAL MEMOIRS
With L. W. Wachtel and E. B. Hart. Studies on the physiology of
manganese in the rat. Am. I. Physiol., 140:72.
1944
With L. J. Teply. Use of germicidal quaternary ammonium salt in
nutritional studies. Proc. Soc. Exp. Biol. Med., 55:59.
With H. C. Lichstein, H. A. Waisman, and P. F. Clark. Influence of
pantothenic acid deficiency on resistance of mice to experimen-
tal poliomyelitis. Proc. Soc. Exp. Biol. and Med., 56:3.
With B. S. Schweigert, I. M. Mclntire, and F. M. Strong. The direct
determination of valine and leucine in fresh animal tissues. l.
Biol. Chem., 155: 183.
With J. H. Shaw, B. S. Schweigert, J. M. McIntire, and P. H. Phillips.
Dental caries in the cotton rat. II. Methods of study and prelim-
inary nutritional experiments. I. Nutr., 28: 333.
With S. R. Ames. Inhibition of the succinoxidase system by cysteine
and cystine. Arch. of Biochem., 5:191.
With W. A. Krehl and F. M. Strong. The biological activity of a
precursor of nicotinic acid in cereal products. I. Biol. Chem..
156:13.
1945
With H. A. Waisman and K. B. McCall. Acute and chronic biotin
deficiencies in the monkey (Macaca mulatto). ]. Nutr., 29:1.
With L. I. Teply. The titrimetric determination of "Lactobacillus
Casei factor" and "Folic acid." J. Biol. Chem., 157:303.
With W. A. Krehl and L. I. Teply. Corn as an etiological factor in
the production of a nicotinic acid deficiency in the rat. Science,
101:283.
With James H. Shaw, B. S. Schweigert, and Paul H. Phillips. Dental
caries in the cotton rat. II. Production and description of the
carious lesions. I. Dent. Res., 23:417.
With B. S. Schweigert and I. E. Tatman. The leucine, valine, and
isoleucine content of meats. Arch. Biochem., 6:177.
With W. A. Krehl, L. I. Teply, and P. S. Sarma. Growth-retarding
effect of corn in nicotinic-acid-low rations and its counteraction
by tryptophane. Science, 101:489.
With B. S. Schweigert, J. M. McIntire, and L. M. Henderson. In-
testinal synthesis of B vitamins by the rat. Arch. Biochem.,
6:403.
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CONRAD ARNOLD ELVEH]EM
161
With B. Schweigert, I. H. Shaw, and P. H. Phillips. Dental caries in
the cotton rat. III. Effect of different dietary carbohydrates on
the incidence and extent of dental caries. l. Nutr., 29:405.
With l. H. Shaw, B. S. Schweigert, and P. H. Phillips. Dental caries
in the cotton rat. IV. Inhibitory effect of fluorine additions to
the ration. Proc. Soc. Exp. Biol. Med., 59:89.
With H. C. Lichstein, H. A. Waisman, K. B. McCall, and P. F. Clark.
Influence of pyridoxine, inositol, and biotin on susceptibility of
Swiss mice to experimental poliomyelitis. Proc. Soc. Exp. Biol.
Med., 60:279.
1946
With W. A. Krehl, P. S. Sarma, and L. I. Teply. Factors affecting the
dietary niacin and tryptophane requirement of the growing rat.
I. Nutr., 31:85.
With W. H. Ruegamer and E. B. Hart. Potassium deficiency in the
dog. Proc. Soc. Exp. Biol. Med., 61:234.
With A. Evenson, Elizabeth McCoy, and B. R. Geyer. The cecal
flora of white rats on a purified diet and its modification by
succinylsulfathiazole. I. Bacterial., 5 1 :5 13.
With A. E. Schaefer and C. K. Whitehair. Purified rations and the
importance of folic acid in mink nutrition. Proc. Soc. Exp. Biol.
Med., 62:169.
With B. A. McLaren and E. F. Herman. Nutrition of rainbow trout;
studies with purified rations. Arch. Biochem., 10:433.
With S. R. Ames. Enzymatic oxidation of glutathione II. Studies
on the addition of several cofactors. Arch. Biochem., 10:443.
With P. S. Sarma and E. E. Snell. The vitamin B6 group. VIII.
Biological assay of pyridoxal, pyridoxamine, and pyridoxine. J.
Biol. Chem., 165:55.
With S. P. Ames and A. I. Ziegenhagen. Studies on the inhibition
of enzyme systems involving cytochrome c. J. Biol. Chem.,
165:81.
With S. R. Ames. Determination of aspartic-glutamic transaminase
in tissue homogenates. J. Biol. Chem., 166:81.
With W. A. Krehl, L. M. Henderson, and }. de la Huerga. Relation
of amino acid imbalance to niacin-tryptophane deficiency in
growing rats. J. Biol. Chem., 166:531.
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162
BIOGRAPHICAL MEMOIRS
1947
With L. J. Teply and W. A. Krehl. The intestinal synthesis of niacin
and folic acid in the rat. Am. }. Physiol., 148:91.
With S. R. Ames and P. S. Sarma. Transaminase and pyridoxine
deficiency. }. Biol. Chem., 167:135.
With T. D. Luckey, P. R. Moore, and E. B. Hart. Growth of chicks
on purified and synthetic diets containing amino acids. Proc.
Soc. Exp. Biol. Med., 64:423.
With B. A. McLaren and E. F. Herman. Nutrition of trout: Studies
with practical diets. Proc. Soc. Exp. Biol. Med., 65:97.
With A. E. Schaefer and C. K. Whitehair. The importance of ri-
boflavin, pantothenic acid, niacin and pyridoxine in the nutri-
tion of foxes. l. Nutr., 34: 131.
With H. A. Lardy and R. L. Potter. The role of biotin in bicarbonate
utilization by bacteria. }. Biol. Chem., 169:451.
With G. W. Newell, T. C. Erickson, W. E. Gilson, and S. N. Gershoff.
Role of "agonized" flour in the production of running fits. }.
Am. Med. Assoc., 135:760.
1948
With A. E. Schaefer, S. B. Tove, and C. K. Whitehair. The require-
ment of unidentified factors for mink. I. Nutr., 35:157.
With V. H. Barki, H. Nath, and E. B. Hart. Production of essential
fatty acid deficiency symptoms in the mature rat. Proc. Soc.
Exp. Biol. Med., 66:474.
With E. l. Wakeman, i. K. Smith, W. B. Sarles, and P. H. Phillips.
A method for quantitative determinations of microorganisms
in carious and noncarious teeth of the cotton rat. }. Dent. Res.,
27:41.
With E. M. Sporn and W. R. Ruegamer. Studies with monkeys fed
army combat rations. }. Nutr., 35:559.
With O. E. Olson, E. E. C. Fager, and R. H. Burris. The use of a
hog kidney conjugase in the assay of plant materials for folic
acid. Arch. Biochem., 18:261.
With L. V. Hankes, L. M. Henderson, and W. L. Brickson. Effect
of amino acids on the growth of rats on niacin-tryptophan-de-
ficient rations. J. Biol. Chem., 174:873.
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CONRAD ARNOLD ELVEH3EM
1949
163
With A. Sreenivasan and A. E. Harper. The use of conjugase prep-
arations in the microbiological assay of folic acid. I. Biol. Chem.,
177:117.
With C. A. Nichol, L. S. Dietrich, and W. W. Cravens. Activity of
vitamin BE in the growth of chicks. Proc. Soc. Exp. Biol. Med.,
70:40.
With K. H. Maddy. Studies on growth of mice fed rations contain-
ing free amino acids. }. Biol. Chem., 177:577.
With C. A. Nichol and A. E. Harper. Effect of folic acid, liver ex-
tract, and vitamin BE on hemoglobin regeneration in chicks.
Proc. Soc. Exp. Biol. Med., 71:34.
With G. B. Ramasarma and L. M. Henderson. Purified amino acids
as a source of nitrogen for the growing rat. I. Nutr., 38:177.
With V. H. Barki, R. A. Collins, and E. B. Hart. Relation of fat
deficiency symptoms to the polyunsaturated fatty acid content
of the tissues of the mature rat. Proc. Soc. Exp. Biol. Med.,
71:694.
With V. H. Barki, P. Feigelson, R. A. Collins, and E. B. Hart. Factors
influencing galactose utilization. I. Biol. Chem., 181 :565.
1950
With H. T. Thompson, P. E. Schurr, and L. M. Henderson. The
influence of fasting and nitrogen deprivation on the concentra-
tion of free amino acids in rat tissues. J. Biol. Chem., 182:47.
With P. Roine. Significance of the intestinal flora in nutrition of the
guinea pig. Proc. Soc. Exp. Biol. Med., 73:308.
With J. N. Williams, Jr., and P. Feigelson. A study of xanthine me-
tabolism in the rat. l. Biol. Chem., 185:887.
With A. E. Denton and I. N. Williams, Jr. The influence of methi-
onine deficiency on amino acid metabolism in the rat. I. Biol.
Chem., 186:377.
With L. S. Dietrich and W. }. Monson. Effect of sulfasuxidine on
the interrelation of folic acid, vitamin Be and vitamin C. Proc.
Soc. Exp. Biol. Med., 75:130.
With L. V. Hankes and R. L. Lyman. Effect of niacin precursors on
growth of rats fed tryptophan-low rations. I. Biol. Chem.,
187:547.
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164
BIOGRAPHICAL MEMOIRS
1951
With I. N. Williams, Jr., P. Feigelson, and S. S. Shahinian. Interre-
lationships of vitamin Be, niacin, and tryptophan in pyridine
nucleotide formation. Proc. Soc. Exp. Biol. Med., 76:441.
With P. Feigelson and I. N. Williams, Jr. Inhibition of diphospho-
pyridine nucleotide-requiring enzymes by nicotinamide. I. Biol.
Chem., 189:361.
With R. I. Sirny and L. T. Cheng. An arginine-proline interdepen-
dence in Leuconostoc mesenteroides P-60. I. Biol. Chem., 190:547.
With I. N. Williams, Jr., and G. Litwack. Studies on rat liver choline
oxidase: an assay method. I. Biol. Chem., 192:73.
With S. N. Gershoff. Studies of the biological effects of methionine
sulfoximine. I. Biol. Chem., 192:569.
With R. L. Lyman. Further studies on amino acid imbalance pro-
duced by gelatin in rats on niacin-tryptophan-low ration. I.
Nutr., 45:101.
1952
.
With A. R. Taborda, L. C. Taborda, and T. N. Williams, Jr. A study
of the ribonuclease activity of snake venoms. I. Biol. Chem.,
194:227.
With D. V. Tappan, U. I. Lewis, and U. D. Register. Niacin defi-
ciency in the rhesus monkey. l. Nutr., 46:75.
With M. Constant and P. H. Phillips. Dental caries in the cotton rat.
XIII. The effect of whole grain and processed cereals on dental
caries procluction. I. Nutr., 46:271.
With I. P. Kring, K. Ebisuzaki, and }. N. Williams, Tr. The influence
of vitamin B6 on the formation of liver pyridine nucleotides. J.
Biol. Chem., 195:591.
With S. S. Shahinian, K. Ebisuzaki, l. P. Kring, and I. N. Williams,
fir. The action of threonine in inducing an amino acid imbal-
ance. Proc. Soc. Exp. Biol. Med., 80:146.
With L. S. Dietrich and W. J. Monson. Utilization of pteroylglu-
tamic acid conjugates in the in vitro synthesis of L. citrovorum
activity. I. Am. Chem. Soc., 74:3705.
With W. L. Davies, W. L. Pond, S. C. Smith, A. F. Rasmussen, fir.,
and P. F. Clark. The effect of certain amino acid deficiencies on
Lansing poliomyelitis in mice. J. Bacterial., 64:571.
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CONRAD ARNOLD ELVEHJEM
1953
165
With H. Nino-Herrera, M. Schreiber, and R. A. Collins. Dermatosis
in weanling rats fed lactose diets. II. Histological studies. I.
Nutr., 49:99.
With A. E. Denton. Enzymatic liberation of amino acids from dif-
ferent proteins. I. Nutr., 49:221.
With G. Litwack and I. N. Williams, fir. The roles of essential and
nonessential amino acids in maintaining liver xanthine oxidase.
I. Biol. Chem., 201:261.
With S. C. Smith, A. F. Rasmussen, fir., and P. F. Clark. Influence
of hyper- and hypothyroidism on susceptibility of mice to in-
fection with Lansing poliomyelitis virus. Proc. Soc. Exp. Biol.
Med., 82:269.
With I. N. Williams, Jr., A. Sreenivasan, and S. C. Sung. Relation-
ship of the deposition of folic and folinic acids to choline oxi-
dase of isolated mitochondria. I. Biol. Chem., 202:233.
With I. N. Williams, Jr., W. I. Monson, A. Sreenivasan, L. S. Die-
trich, and A. E. Harper. Effects of a vitamin Be deficiency on
liver enzymes in the rat. J. Biol. Chem., 202:151.
With A. E. Harper, W. I. Monson, and D. A. Benton. The influence
of protein and certain amino acids, particularly threonine, on
the deposition of fat in the liver of the rat. I. Nutr., 50:383.
1954
With A. E. Denton. Amino acid concentration in the portal vein
after ingestion of amino acid. T. Biol. Chem., 206:455.
With W. L. Loeschke. Prevention of urinary calculi formation in
mink by alteration of urinary pH. Proc. Soc. Exp. Biol. Med.,
85:42.
With W. I. Monson, A. E. Harper, and M. E. Winje. A mechanism
of the vitamin-sparing effect of antibiotics. J. Nutr., 52:627.
With A. E. Harper, D. A. Benton, and M. E. Wince. Leucine-
isoleucine antagonism in the rat. Arch. Biochem. Bionhvs..
51:523.
With M. E. Winje, A. E. Harper, D. A. Benton, and R. E. Boldt.
Effect of dietary amino acid balance on fat deposition in the
livers of rats fed low protein diets. I. Nutr., 54: 155.
~ , ,
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166
BIOGRAPHICAL MEMOIRS
1955
With L.-E. Ericson and }. N. Williams, Jr. Studies on partially pur-
ified betaine-homocysteine transmethylase of liver. I. Biol.
Chem., 212:537.
With D. A. Benton and A. E. Harper. Effect of isoleucine supple-
mentation on the growth of rats fed zein or corn diets. Arch.
Biochem. Biophys., 57:13.
With Selma Hayman, S. S. Shahinian, and I. N. Williams, [r. Effect
of 3-acetylpyridine on pyridine nucleotide formation from
tryptophan and niacin. J. Biol. Chem., 217:225.
With P. D. Deshpande, A. E. Harper, and Felipe Quiros-Perez. Fur-
ther observations on the improvement of polished rice with
protein and amino acid supplements. J. Nutr., 57:415.
With H. R. Heinicke and A. E. Harper. Protein and amino acid
requirements of the guinea pig. I. Effect of carbohydrate, pro-
tein level and amino acid supplementation. I. Nutr., 57:483.
1956
With D. A. Benton, A. E. Harper, and H. E. Spivey. Leucine, iso-
leucine, and valine relationships in the rat. Arch. Biochem. Bio-
phys., 60:147.
With D. A. Benton and A. E. Harper. The effect of different dietary
fats on liver fat deposition. J. Biol. Chem., 218:693.
With L. E. Ericson, A. E. Harper, and [. N. Williams, Jr. Effect of
diet on the betaine-homocysteine transmethylase activity of rat
liver. I. Biol. Chem., 219:59.
With A. E. Harper, L. E. Ericson, and R. E. Boldt. Effect of thyroid-
active substances on the betaine-homocysteine transmethylase
activity of rat liver. Am. I. Physiol., 184:457.
With R. F. Wiseman, W. B. Sarles, D. A. Benton, and A. E. Harper.
Effects of dietary antibiotics upon numbers and kinds of intes-
tinal bacteria used in chicks. J. Bacterial., 72:723.
1957
With P. D. Deshpande, A. E. Harper, and Macie Collins. Biological
availability of isoleucine. Arch. Biochem. Biophys., 67:341.
With P. D. Deshpande and A. E. Harper. Nutritional improvement
of white flour with protein and amino acid supplements. J.
Nutr., 62:503.
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CONRAD ARNOLD ELVEH3EM
167
With J. D. Gupta. Biological availability of tryptophan. J. Nutr.,
62:313.
With F. N. Hepburn and E. W. Lewis, fir. The amino acid content
of wheat, flour, and bread. Cereal Chem., 34:312.
1958
With M. M. Chaloupka, J. N. Williams, Jr., and May S. Reynolds.
Relative roles of niacin and tryptophan in maintaining blood
pyricline nucleotides, nitrogen balance and growth in adult rats.
I. Nutr., 63:361.
With P. D. Deshpande and A. E. Harper. Amino acid imbalance
and nitrogen retention. I. Biol. Chem., 230:335.
With I. D. Gupta, A. M. Dakroury, and A. E. Harper. Biological
availability of lysine. l. Nutr., 64:259.
With Narindar Nath and A. E. Harper. Dietary protein and serum
cholesterol. Arch. Biochem. Biophys., 77:234.
With A. Yoshida and A. E. Harper. Effect of dietary level of fat and
type of carbohydrate on growth and food intake. J. Nutr.,
66:217.
1959
With Narindar Nath, Ruta Wiener, and A. E. Harper. Diet and
cholesteremia. I. Development of a diet for the study of nutri-
tional factors affecting cholesteremia in the rat. I. Nutr., 67:289.
With A. I. Bosch and A. E. Harper. Factors affecting liver pyridine
nucleotide concentration in hyperthyroid rats. Soc. Exp. Biol.
Med., 100:774.
With W. L. Loeschke. The importance of arginine and methionine
for the growth and fur development of mink fed purified diets.
I. Nutr., 69:147.
With W. L. Loeschke. Riboflavin in the nutrition of the chinchilla.
I. Nutr., 69:214.
Representative terms from entire chapter:
biographical memoirs
