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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 researchworrier! 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 rateby 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 historyattempted 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- tophanan 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

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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.