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WILLIAM GUMMING ROSE April 4, 1887-September 25, 1985 BY HERBERT E. CARTER AND MINOR J. COON WILLIAM GUMMING ROSE, a member of the National Acad- emy of Sciences from 1936, cried in Urbana, Illinois, at the age of ninety-eight. Thus ended the career of a clecli- catecl and inspiring teacher ant! an outstanding pioneer in biochemistry and nutritional science. He renclerec! the Uni- versity of Illinois clistinguishec! service ur~tiT his retirement in 1955. In research he clevoted his attention to the inter- mecliary metabolism of amino acids, creatine, uric acicI, and chemically related compounds and was renowned for the discovery, isolation, ant! identification of threonine. The characterization of the last of the amino acids to be found as universal components of proteins lect to his cletermina- tion of the complete essential amino acid requirements of the laboratory rat and culminates! in the establishment of the amino acic] requirements of humans. EDUCATION AND EARLY LIFE William C. Rose was born in Greenville, South Carolina, en c! spent his childhood! in small communities in the Caro- linas, where his father, John M. Rose, served as a Presbyte- rian minister. Money was scarce, particularly since the cleric's small salary was reducer! by generous gifts to religious and 253

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254 BIOGRAPHICAL MEMOIRS humanitarian causes, but somehow the necessities were pro- viclec! and the best possible education was afforded the chil- ciren in the family. Young Will attencled an assortment of local schools until the age of fourteen, when the inadequacy of the education causer! his father to remove him from school en c! tutor him at home. He was well prepared by the time he entered college and hac! aIreacly been introduced to Latin, Greek, and Hebrew en c] had acquirer! an interest in chemistry from reacting Remsen's An Introduction to the Study of Chemistry, a college text his older sister hac! used. Will wisher! to attenc! a large university, but his father thought his son, at age sixteen, was too young en c! so convincer! him to attend Davidson College in North Carolina, a school for which Will cleveloped a lifelong affection. While in graduate school at Yale University, Rose clecided on the branch of chemistry he would pursue. During his initial interview with Russell Chittenden, director of the Sheffield Scientific School, Rose mentioned an interest in foot] chemistry. This suggestion brought an introduction to Lafayette Mendel, which not only guiclec! Rose's interest en c! work towarc! biochemistry but also lecI to a strong friencI- ship that endures! until Menclel's cleath. In 1911 Rose left Yale for an instructorship in physiological chemistry at the University of Pennsylvania, a department then headed by Alonzo Taylor, which was followecl by a short period of acI- vancec! study with Professor Franz Knoop at the University of Freiburg. While in Germany, Rose receiver! a cablegram from Galveston asking him to come to the College of Mecli- cine to organize a department of biochemistry. At first Rose was hesitant about the offer because he felt obliged to go back to the University of Pennsylvania, but he was fully reassured by a cable from Taylor containing the terse single sentence, "You clarnec! fool, ~ recommended you for that job." This message came in response to his own cable in

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WILLIAM GUMMING ROSE 255 which he had turned down the offer. He then accepted the position of associate professor of biological chemistry at the University of Texas en c! quickly rose to the rank of professor and head of the department. In 1922 Rose was persuaded to move to the University of TIlinois, where he was appointed professor en c! head of the Division of Physi- ological Chemistry (the name was later changer! to Bio- chemistry) in the Chemistry Department and found a per- manent and very supportive home for his scientific career. He provided dedicates! service to that institution for the next thirty-three years. RESEARCH ACCOMPLISHMENTS In research Rose displayed a gift for meticulous experi- mentation and for thoroughness en c! clarity in preparing his results for publication. His early studies on creatine and its clehyciration product, creatinine, carried out with Mencle! at Yale and published in 1911, dealt with the role of carbo- hydrates in the metabolism of these compounds and with the effect of inanition on the creatine content of muscle. In subsequent years Rose en cl his associates macle excellent use of the experimental methods available at the time, chiefly nutritional studies, to explore the metabolic relationship of creatine to creatinine and of both to other nitrogenous substances. They observed that the ingestion of diets high in protein failed to induce the excretion of creatine in normal men and women en c! concluded that there was no exogenous source of urinary creatine in the absence of creatine in the diet. In addition, no relationship was observed in growing rats between the arginine content of the diet and total crea- tinine elimination in the urine. Furthermore, in - human subjects no evidence could be obtained that exogenous argi- nine was catabolized to creatine or creatinine. In light of

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256 BIOGRAPHICAL MEMOIRS present knowledge, such studies did not reveal the involve- ment of this amino acid in amidine transfer in creatine biosynthesis, but they are still of fundamental interest. We will have much to say in what follows about the pioneering studies in Rose's laboratory on essential amino acids; suf- fice it to say here that creatine and creatinine were found to be incapable of replacing (lietary histi~line for the growth of rats. The knowledge available at that time on the me- tabolism of creatine and creatinine was ably summarized by Rose in 1933 and 1935 in insightful articles in the Annual Review of Biochemistry. From his own experimental studies and a summary of the literature he published in 1923 in Physiological Reviews, Rose concluded that endogenous purines may have their ultimate origin in arginine and histidine but that the ex- tent of their synthesis might be limited to the anabolic needs of the organism. And in studies similar to those referred to above involving creatine ant! creatinine, he established that neither adenine nor guanine (nor, for that matter, a mix- ture of all four of these compounds) was capable of replac- ing dietary histidine for the growth of rats. In addition to his interest in nitrogenous compounds as already summa- rized, Rose made useful and scholarly contributions to knowl- edge in several other quite diverse areas, including analyti- cal methods, the nephropathicity of dicarboxylic acids and their derivatives, the occurrence of copper in marine or- ganisms, and digestive enzymes in coelenterates, elasmo- branchs, and teleosts. However, after he developed an in- terest in amino acid metabolism and nutrition at the University of Illinois, Rose elevated himself wholeheartedly to this subject, which soon brought him international rec- . . Ognltlon. Many years later, in an article titled "How Did It Hap- pen?", Rose described the events leading up to his first

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WILLIAM GUMMING ROSE 257 report, in 1924, on an indispensable amino acid. He was aware of the much earlier work of Osborne and Mended in which zein of corn servecl as a clietary protein on which rats failer! to grow until tryptophan and lysine were incorpo- ratect into the ration. That was the first clear-cut evidence that individual amino acids might be required by animals. While at the University of Texas, Rose continued his work on creatine-creatinine metabolism, as aIreacly mentioned, anti, in order to stucly arginine as a possible precursor of these compounds, clecicleci to prepare a casein hyctrolysate and remove this amino acid as completely as possible. In view of work in another laboratory indicating that arginine and histicline were mutually interchangeable in metabolism, he changed the plan, and after preparing the hy(lrolysate by enzymatic action, followocl by miTct acic! treatment, he took it to dryness, shipped it to his new scientific home in TIlinois, anti there removed the arginine anti histicline. Rats lost weight on the treate(1 hyctrolysate but respon(le(1 im- pressively when histictine was included in their food. How- ever, arginine was totally incapable of replacing histictine for growth. Rose rightly considered the difference in re- sponse of the animals to the two amino acids to be little short of sensational. Having discoverecl that histicline is a dietary essential, he cleci(led to investigate the nutritional role of all of the other amino acids as well. Rose realizer! that work with protein hycirolysates hacI limitations but user! them in place of proteins to show that a variety of closely related synthetic compounds could not be substituted for cystine, histidine (with the exception of imidazolelactic acicI), and tryptophan (with the exception of the N-acety! com- pound, the ethyl ester, and indolelactic and indolepyruvic acids) . Rose then undertook investigations using mixtures of pure amino acids as the source of dietary nitrogen. Following

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258 BIOGRAPHICAL MEMOIRS the extremely important observation that (liets containing nineteen highly purifier] amino acids wouic! not support growth, his laboratory made a painstaking effort to identify the missing growth essential in proteins. After several years they finally succeeded in obtaining the unknown compound in pure crystalline form, as clescribed in the 1935 lancimark paper by McCoy, Meyer, and Rose. The structure was estab- lished as oc-amino-13-hydroxy-n-butyric acid, ant! the purified amino acic! was clemonstratect to incluce maximum growth, thus constituting the first successful attempt to rear ani- mals on a ration containing purified amino acids as the sole source of nitrogen. In a 1979 symposium on earlier nutritional discoveries, H. E. Carter clescribect how he became a faculty member in biochemistry at Illinois and was given what he consiclerec3 a wonclerful- opportunity of participating in the threonine story. He accomplished the chemical synthesis of the four isomers of oc-amino-~-hyciroxy-n-butyric acid and clemonstratecl that only one form wouIcl support the growth of rats. This isomer, analogous in structure to D-threose and having a steric relationship to that of the other naturally occurring [-amino acids, was designatecI [-threonine. Thus, the way was paved to classify other amino acids in the essential or nonessential category as judged by the main- tenance of normal growth in the rat. Thorough experiments extending over the next two clecacles lecT to many impor- tant conclusions. Only ten of the twenty-two amino acids known to exist in proteins are inclispensable dietary com- ponents. These are histicTine, isoleucine, leucine, threonine, Tysine, methionine, phenylalanine, tryptophan, valine, en cl arginine. With the exception of arginine, the removal of any one of these from the food of growing rats leacis to profound nutritive failure, accompanied by a rapid decline in weight, Toss of appetite, and eventual cleath. However,

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WILLIAM GUMMING ROSE 259 without arginine the animals continue to gain weight but at a suboptimal rate, thus indicating that this amino acid can be manufactured by the body but only slowly. In a study of the significance of the amino acids in canine nutrition, it was found that those amino acids that are dispensable for the growing rat are also dispensable for the adult dog, as . . . . . - judged by the maintenance of nitrogen equilibrium. Not unexpectedly, arginine is not needed, presumably because . ~ . ~ . ~ _ _1 ~ 1_ _ _ 1 _ _ 1 . _ ~ _ ~ ~ 1 T] ~ ~ ~ the rate ot synthesis IS adequate in tne actull animal. nose and his associates then determined the quantitative amino acid requirements by establishing the minimum amount needed to support optimal growth in the laboratory rat. In addition, many other interesting findings were made with this species using diets containing purified amino acids. ~ . . . ~ 1 for example, cystlne was found to stimulate growth only when methionine was fed in suboptimal amounts and, simi- larly, the phenylalanine requirement could be partially re- placed by tyrosine. In other attempts at substitution by re- lated compounds, argininic acid was shown to be a poor - substitute for arginine, and the ~isomers of phenylalanine and methionine were found to be active, whereas those of tryptophan and valine were only partly effective. Glycine, glutamate, urea, or ammonium salts were found to serve as a source of nitrogen for synthesis of the nonessential amino acids, and the effect of urea was confirmed with the ~5N- labeled compound. In other studies that contributed sig- nificantly to knowledge of amino acid metabolism, 5-~4C- labeled glutamate was observed to lead to labeled praline and arginine, thus establishing the reversibility of the known reactions in which glutamate is formed from the other two compounds. In addition, the fate of valine was investigated in the phlorhizinized dog, and three of the carbon atoms were found to yield glucose. The experiments described briefly above provided highly

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260 BIOGRAPHICAL MEMOIRS important information about synthetic reactions that ani- mals couicl ant! could not accomplish, but the ultimate ob- jective of Rose's investigations was establishment of the amino acid requirements of the human species. In 1942 he took on this research challenge, with healthy mate graduate stu- dents as the experimental subjects.2 The diets consisted of corn starch, sucrose, butter fat from which the protein hacI been removed, corn oil, inorganic salts, the known vita- mins, en c! mixtures of highly purif~ec3 amino acids. The only variables allowecI, other than when changes were pur- posely macle in the amino acicis consumed, were clistillecI water and celluflour, a product that proviclecI bulk but had no nutritive value, nitrogen, or flavor. The only unusual component of this otherwise blancI cliet was a large brown c ancly containing a concentrated liver extract as a possible source of unknown vitamins, sweetened with sugar and fla- vore(1 with peppermint oil, which provicle(1 a never-to-be- forgotten taste. Total urinary and fecal nitrogen were de- terminecI, and by the criterion of nitrogen equilibrium it was established that the twelve amino acids previously shown to be clispensable for animals were also clispensable for hu- mans. The remaining ten amino acicis were then remover! from the diet one at a time; a pronouncer] negative nitro- gen balance ensued in the case of isoleucine, leucine, tryp- tophan, Tysine, methionine, phenylalanine, threonine, ancI valine. In contrast, the removal of arginine hac3 no effect, a fancying that was not surprising inasmuch as animals have a limited ability to synthesize this compound, as aIreacly stated. The results obtained with histicline, however, were most un- expectec3, since on a diet lacking this amino acic! the sub- jects all maintained normal nitrogen equilibrium. Thus, only eight amino acids are essential clietary components for the aclult human. However, as Rose was careful to point out, certain amino acids not necessary for nitrogen equilibrium

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WILLIAM GUMMING ROSE 261 uncler ordinary circumstances might become indispensable cluring disease or for special needs in detoxification, repro- duction, or lactation. These investigations continued into the early 1950s, re- sulting in sixteen papers in The fournal of Biological Chemis- try, which C. Glen King, trustee of the Nutrition Founda- tion, clescribecl as a series that stands as a classic in the history of nutrition and for the benefit of humans. Impor- tantly, the studies established the quantitative as well as the qualitative amino acid neecis. Levels ranging from as Tow as 0.25 grams per day of tryptophan to as high as I.] grams per day of several other amino acids were proposed as mini- mal levels, with twice as much providing what was consi(l- erecI to be a safe margin. In further investigations with Illinois graduate students as subjects, who were grateful in those clays for the free rations, the (loliar a (lay they were paid, and the prospect of seeing their initials in print in Rose's widely read publica- tions, Rose made many other significant finclings. A higher caloric intake is needed to maintain nitrogen equilibrium on diets containing mixtures of purified amino acids as compared to casein, for reasons that are not yet well uncler- stoocI. Cystine spares part of the methionine requirement, which is of significance in those parts of the florid in which the latter appears to be the limiting amino acid in native cliets. Similarly, tyrosine spares part of the phenylalanine requirement. In studies involving the D isomers of the es- sential amino acids, only that of methionine was found to be well utilized by the human organism. Of related inter- est, acetyI-~-tryptophan is effective but not the acetyI-D form, a matter of metabolic and also practical importance in view of reports by others in the literature that the acetyI-DL prepa- ration might be fully utilized and thus be less costly as a dietary supplement than racemic unsubstitutec! tryptophan.

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262 BIOGRAPHICAL MEMOIRS Another report by other investigators that conic! not be confirmed in the Rose laboratory, much to his relief ant! that of his subjects, was the claim that arginine deficiency aclversely affects spermatogenesis. Thus, the only changes observer! in these relatively short term studies when an es- sential amino acid was removed from the cliet were negative nitrogen balance, a loss of appetite, and a sense of fatigue. The above account, though necessarily brief, may give the reader an idea of how an unexpected fincling macle in a study on the possible relationship of amino acids to creat- ine synthesis eventually led to discovery of the last of the amino acids occurring in proteins and to establishment of the qualitative and quantitative amino acid requirements of animals ant! of the human species. Incleed, no other scien- tist has hac! a comparable record in identifying and estab- lishing the quantitative requirements for so many essential nutrients. Rose's finclings have hac! many useful applications in acI- dition to their contribution to basic knowlecige. For ex- ample, they made it possible to predict the nutritional qual- ity of a protein for human diets from the amino acid composition, rather than from animal tests, and to devise highly effective mixtures of amino acids for the intravenous feeding of surgical and pediatric patients. In ~L942, partly in response to wartime nutritional problems and to the change in emphasis from acute to chronic dietary deficiencies, two important institutions were creates! the Nutrition Founcia- tion anct the Food and Nutrition Board of the National Research Council. Rose played a significant role in the cle- velopment ant! ongoing activities of both of these nongov- ernmental organizations, serving as a member of the Scien- tific Advisory Boarc! of the former from 1943 to 1956 and as a member of the latter from 1940 to 1947. With the Food and Nutrition Board, he was instrumental in advising gov-

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WILLIAM GUMMING ROSE 263 ernmental agencies on the implications of meat rationing and on minimum desirable daily allowances, as well as on the dietary usefulness of vegetable proteins. As chairman of its Committee on Protein Foods, Rose dealt with problems of supply and nutritional quality of protein foods. The re- sults were issued in two comprehensive publications, titled The Evaluation of Protein Nutrition with Emphasis on Amino Acid Proportionalities and The Evaluation of Protein Nutrition. PERSONAL TRAITS Despite his many professional duties and dedication to his research and his students, Rose found time for other interests. In 1913, upon his return from Freiburg, he mar- ried Zula Franklin Hedrick, a North Carolinian. She was at his side for many happy years, and the "two Roses" exerted a wonderfully positive influence on all who knew them. They had no children of their own but instead a large "fam- ily" in which they took a personal interestthe ninety gradu- ate students who studied under Will Rose, of whom fifty-six received the Ph.D. degree. In later years he often com- mented on his extraordinarily happy family life until Zula's death in 1965, his exciting professional life, and the thrill of watching his students grow into professional stature. When asked what accounted for his longevity, Rose sim- ply commented that he had been interested in everything all his life. He particularly enjoyed birdwatching, amateur photography, travels by automobile, and the history of sci- ence. He and his wife made numerous tours of the country by car, and until he was ninety-f~ve he drove annually to Davidson, North Carolina, where he spoke to chemistry classes on the campus. Because of his extreme caution as a driver, it was a source of some amusement to his friends at Davidson College that at age ninety-three Rose got a speeding ticket during the drive down from Illinois. On another such visit

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264 BIOGRAPHICAL MEMOIRS he hac! to have a heart pacemaker installed, after which he got in his car and cirove himself home to Illinois. Rose was the class historian at his undergracluate college, and he took a strong interest in the origins of Davidson College as well as of all the other institutions he was associated with over the years and of science in this country in general. After painstaking verification of the historical facts, he wrote fascinating and insightful accounts of the early clays of Ameri- can biochemistry. In "John R. Young, First American Bio- chemist," an introductory essay to a monograph by Young, originally publishect in ~ 803 and titIec3 An Experimental In- quiry into the Principles of Nutrition and the Digestive Pro cess,3 Rose clescribec! a remarkable thesis submitted to the Uni- versity of Pennsylvania for the ctegree of cloctor of mecli- cine. Considerable space is clevoted to the careers of Young's teachers, including the famous Benjamin Rush, ant! how the youthful author "takes issue with the most revered au- thorities of his era SpalIanzini, Cullen, en c] Rush and then proceeds to prove that each was guilty of erroneous conclu- sions." In "Recollections of Personalities Involves! in the Early History of American Biochemistry,"4 Rose describes his association with scientists responsible for the early de- velopment of this field and provides warm insight into their contributions and personal characteristics. The final para- graph of that article is quoted here because of the remark- able insight it provides: Because of the early start of the Yale laboratories, and the superior genius of Samuel W. Johnson, Russell H. Chittenden, and Lafayette B. Mendel, it is not surprising that such a large proportion of the biochemists produced in this country until approximately 1915 had their training at Yale. This would have occurred wherever Johnson, Chittenden, and Mendel happened to be locatedat Harvard, Chicago, here, or anywhere. Intellects of their caliber would have found a way to do what they did regardless of the place in which fate cast their lot. In the development of a university, as in the life

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WILLIAM GUMMING ROSE 265 and growth of an individual, progress and ultimate attainment depend so largely upon the vision, enthusiasm, and determination of the individual participants. Perhaps, this is a truth that each one needs to remember as he carves his future out of the events and experiences of the present. For many years Rose taught the two core biochemistry courses at Illinois and exhibited a rare talent at imparting enthusiasm about biochemistry to the undergraduate and graduate students who attended his meticulously prepared lectures. The subject came alive with his engrossing stories about the early history of the field and the personalities involved. No mention of his remarkable ability as a teacher would be complete without reference to the weekly gradu- ate student seminars and teas at which he presided, impart- ing scientific knowledge and on some occasions entertain- ing his audience as an incomparable raconteur. His students were somewhat in awe of the professor, per- haps wondering whether they could meet his exacting stan- d~rd.s or could horse to emulate the seeming ease with which he succeeded in all of his professional endeavors. They learned in time that behind his somewhat reserved and formal manner was a genuine warmth and an understand- ing that young scientists develop their full potential only by profiting from their mistakes. His faculty colleagues also . . . - . , . ~ . ~- admired his many talents and sterling personal characteris- tics. Herbert E. Carter, who became the second member of the faculty of the Biochemistry Division in 1932, has com- mented that he became interested in biochemistry upon hearing Rose's lectures and states, "I recall with deep grati- tude that following my graduate work in organic chemistry Dr. Rose invited me to join the Biochemistry Division. His only request of me was that ~ undertake the chemical syn- thesis of the newly discovered threonine, a project which was very fruitful in leading to my own areas of research. He enriched my life as mentor, colleague, and friend for fifty

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266 BIOGRAPHICAL MEMOIRS years." Carl S. VestTing, who became the thirc! faculty mem- ber of the division, macle the following remarks when he served in 1981 as moderator of "Conversations with Will- iam C. Rose," a tape-recorclec3 group discussion: Absolutely uncompromising in all matters involving integrity and sincerity, he has personified many of those qualities of loyalty, unselfishness, and friendliness which mark the unusual individual. He has shown a unique blend of decisiveness and unpretentiousness in his relationships to his asso- ciates. RECOGNITION AND AWARDS Rose's research achievements en c! reputation as a stimu- lating ant! inspiring teacher brought him wide recognition en c! many honors. These incluclecI numerous invitations to lecture anc! serve as a consultant on biochemical aspects of nutrition. In acictition to his work on behalf of the Nutri- tion Founclation ant! the National Research Council, as al- ready clescribecI, he served on the Council on Pharmacy and Chemistry of the American Medical Association, the Advisory Board of the Wistar Institute, anc! the National Advisory Health Council of the U.S. Public Health Service. He received honorary cloctor of science degrees from Davidson College, Yale University, the University of Chi- cago, and the University of TIlinois and was electecl to mem- bership in the National Academy of Sciences. As an indica- tion of his leaclership qualities and the respect of his colleagues nationally, he was electecl to serve as president of the American Society of Biological Chemists from 1939 to 1941 ant! president of the American Institute of Nutri- tion from 1945 to 1946. Other major honors incluclec] the Osborne and Menclel Award of the Institute of Nutrition, of which he was the first recipient (19491; the Willard Gibbs Medal of the American Chemical Society (19521; the Charles F. Spencer Mecial of the American Chemical Society ~9571;

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! WILLIAM GUMMING ROSE 267 the Twentieth Anniversary Award of the Nutrition Founcia- tion (19611; and the National Meclal of Science for 1966, conferrer! by the President at the White House. On the occasion of his ninetieth birthday, Rose's former students, colleagues, ant! friends assembled in Urbana to join him in the celebration. He was much surpriser! when presented with a handsome bronze plaque announcing the establishment of the William C. Rose Lectureship in Bio- chemistry and Nutrition "on the occasion of his 90th birth- day and presented with love, admiration and gratitude by his family of former students and colleagues." The plaque showed, in aciclition to his likeness and a sketch of the Noyes Laboratory, the structures of the essential amino acids and the stereochemistry and crystal structure of threonine, with a quotation and chart from his classical 1935 paper pub- lishecl in The fournal of Biological Chemistry: The data demonstrate conclusively that the crystalline compound is the new essential we have been endeavoring to isolate for several years. Fur- thermore, the experiments shown in Chart 1 represent the first successful efforts to induce growth in animals upon diets carrying synthetic mixtures of highly purified amino acids in place of proteins. It may be noted that this prestigious national award, now administered by the American Society for Biochemistry and Molecular Biology, has been presenter! annually since 1978, and the awardees have all received a duplicate of the same plaque. The lectures of the recipients were originally given in Urbana to allow Rose to attend but are now presented at the society's national meetings. William J. Haines, a former student, made the following closing remarks at the celebra- tion: Dr. Rose enhanced the quality of life for his students by encouraging and supporting those things which enriched the mind and spirit. Good charac- ter was the essential raw material good taste was the product. His per-

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268 BIOGRAPHICAL MEMOIRS sonal dedication to the highest quality of performance was projected in his wise counsel. For all this, he demanded nothing in return, except excel- lence in performance (and behavior) of his academic children and their children, the latter whom he considers to be his academic grandchildren. No scientist count ask for a finer memorial than having made major discoveries that contributed to the welfare of the human race ant! having received the respect, acimira- tion, and gratitude of a family of former students en c! col- leagues. THE AUTHORS ARE GRATEFUL to Leland M. Park for material from the Davidson College Library Archives, Davidson, North Carolina, that deals with Rose's early days there and his longstanding relationship with the college; to Ellen Handler and Robert T. Chapel for useful information from the University of Illinois Archives; and to the National Academy of Sciences for a brief tribute written by Caroline K. McEuen after Rose's death. NOTES 1. H. E. Carter. Identification and synthesis of threonine. Fed. Proc. 38(1979):2684-86. 2. Fifty years later the first two human subjects recalled the early experiments. J. E. Johnson and W. J. Haines. Role of amino acids in human nutrition. FASEB Journal 6 ~ 1992) :2361-62. 3. Published in 1959 by the University of Illinois Press, Urbana. 4. journal of Chemical Education 46 ~ 1 969~: 759-63.

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WILLIAM GUMMING ROSE SELECTED BIBLIOGRAPHY 1926 269 With G. I. Cox. Further experiments on the alleged interchange- ability of arginine and histidine in metabolism. [. Biol. Chem. 68:217- 23. 1929 With C. P. Berg. Tryptophane and growth. I. Growth.upon a nyptophane- deficient basal diet supplemented at varying intervals by the separate feeding of tryptophane. J. Biol. Chem. 82:479-84. 1931 Feeding experiments with mixtures of highly purified amino acids. I. The inadequacy of diets containing nineteen amino acids. [. Biol. Chem. 94:155-65. With W. Windus and F. L. Catherwood. Feeding experiments with mixtures of highly purified amino acids. III. The supplementing effect of casein fractions. /. Biol. Chem. 94:173-84. 1934 With C. T. Caldwell. Feeding experiments with mixtures of highly purified amino acids. V. Additional properties of the unknown growth essential present in proteins. {. Biol. Chem. 107:57-73. 1935 The metabolism of creatine and creatinine. Annul Rev. Biochem. 4:243- 62. With M. C. Womack. Feeding experiments with mixtures of highly purified amino acids. VII. The dual nature of the "unknown growth essential." {. Biol. Chem. 112:275-82. With R. H. McCoy and C. E. Meyer. Feeding experiments with mix- tures of highly purified amino acids. VIII. Isolation and identifi- cation of a new essential amino acid. [. Biol. Chem. 112:283-302. 1936 With C. E. Meyer. The spatial configuration of oc-amino-B-hydroxy- n-butyric acid. J. Biol. Chem. 115:721 -29.

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270 BIOGRAPHICAL MEMOIRS 1939 With E. E. Rice. The significance of amino acids in canine nutri tion. Science 90:186-87. 1942 With W. I. Haines and i. E. Johnson. The role of the amino acids in human nutrition. i. Biol. Chem. 146: 683-84. 1946 With M. Womack. The partial replacement of dietary phenylalanine by tyrosine for purposes of growth. [. Biol. Chem. 166:429-34. 1948 With M. l. Oesterling and M. Womack. Comparative growth on di- ets containing ten and nineteen amino acids, with further obser- vations upon the role of glutamic and aspartic acids. [. Biol. Chem. 176:753-62. 1949 Amino acid requirements of man. Fed. Proc. 8:546-52. With L. C. Smith, M. Womack, and M. Shane. The utilization of the nitrogen of ammonium salts, urea, and certain other compounds in the synthesis of non-essential amino acids in vivo. J. Biol. Chem. 181 :307-16. 1954 With G. F. Lambert and M. l. Coon. The amino acid requirements of man. VII. General procedures; the tryptophan requirement. J. Biol. Chem. 211 :815-27. 1955 With B. E. Leach, M. I. Coon, and G. F. Lambert. The amino acid requirements of man. IX. The phenylalanine requirement. [. Biol. Chem. 213:913-22. With M. J. Coon, H. B. Lockhart, and G. F. Lambert. The amino acid requirements of man. XI. The threonine and methionine requirements. [. Biol. Chem. 215:101-10. With R. L. Wixom. The amino acid requirements of man. XIII. The

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WILLIAM GUMMING ROSE 271 sparing effect of cystine on the methionine requirement. [. Biol. Chem. 216:763-73. With R. L. Wixom. The amino acid requirements of man. XIV. The sparing effect of tyrosine on the phenylalanine requirement. J. Biol. Chem. 217:95-101. With R. L. Wixom, H. B. Lockhart, and G. F. Lambert. The amino acid requirements of man. XV. The valine requirement; sum- mary and final observations. [. Biol. Chem. 217:987-95. With R. L. Wixom. The amino acid requirements of man. XVI. The role of the nitrogen intake. [. Biol. Chem. 217:997-1004. 1956 With E. E. Dekker. Urea as a source of nitrogen for the biosynthesis of amino acids. {. Biol. Chem. 223:107-21. 1968 The sequence of events leading to the establishment of the amino acid needs of man. Am. f. Publ. Health 58:2020-27. 1979 How did it happen? Ann. N.Y. Acad. Sci. 325:229-34.

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