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HERBERT McLEAN EVANS September 23, Z8S2-March 6, 1971 BY GEORGE W. CORNER HERBERT MCLEAN EVANS, anatomist, endocrinologist, and bibliophile, was born in Modesto, California, Septem- ber 23, 1882. His father, Clayburn Wayne Evans, a native of Alabama, was the leading physician and surgeon in the then- small town; he is said to have been the first in the upper San Joaquin Valley to do abdominal surgery. Herbert Evans's mother, nee Bessie McLean, came of a Virginia family. Her father practiced medicine in Modesto, and her brother, Robert McLean, was professor of surgery and dean of the San Francisco medical faculty of the University of California. Dr. C. W. Evans was a man of vigorous rather than polished character; Bessie McLean Evans and her brother Robert were persons of refined manners and tastes. Herbert Evans thus began life in a strongly medical family and with a varied store of traits and tem- peraments. He attributed much of his early interest in science, literature, and history to a cultivated high school principal in his home town and the excellent library at the school. This exposure to books in boyhood was fortunate, for with his family background and strong pressure from his father it was inevitable that the young man should enter the medical profession and that his college education (at Berkeley) would be directed to that end, at the expense of cultural interests. Indeed, his father seems to 153

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154 BIOGRAPHICAL MEMOIRS have been dubious about letting him have more than the two years of premedical preparation then required; and young Evans himself concentrated the rest of his college years on science. Among his teachers, one who especially won his admiration was the celebrated paleontologist John C. Merriam, who during Evans's senior year in college took him on a field trip to Idaho. Evans's first published contribution to science, about the time of his graduation from college, was the description of a fossil fish spine from the Triassic of Idaho, which Merriam encouraged him to study. In college, Evans distinguished himself sufficiently to be, at the commencement of 1904, one of the two students then custo- marily chosen to deliver brief graduation addresses. He spoke on the importance of biological research for human welfare. In the autumn he enrolled in the medical school of the uni- versity, in whose Berkeley laboratories he pursued the courses of the first year of professional study. The professor of anatomy was Joseph Marshall Flint, a surgeon who had done some ana- tomical research at Johns Hopkins and a stimulating lecturer; his associates were Irving Hardesty, a productive histologist, and Robert Orton Moody, a competent teacher of gross human anatomy. The next summer Evans boldly took two steps that were quite contrary to his parents' judgment. In the first place he decided not to continue the study of medicine at the University of California, but instead to go to the School of Medicine of Johns Hopkins University where, his father feared, his leaning toward science might turn him away from the practice of medi- cine. The other, equally bold step was to marry his college sweetheart, Anabel Tulloch, like him tall and handsome, and in her way as spirited and impetuous as he. Since her family also strongly disapproved the match, the young people were married privately and departed for Baltimore.

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HERBERT MCLEAN EVANS 155 At that time married medical students were unheard of at Johns Hopkins and most other medical schools. Fearing that he might not be admitted, Herbert concealed his marriage from the admissions office. He and Anabel set up housekeeping at some distance from the school, where in a poor city neighborhood the bride from California's broad and verdant spaces was cooped up and separated from her husband all day long, among unintel- lectual neighbors, seeing nothing of his associates and activities, feeling lonely and neglected. Years afterward each of them separately spoke to the writer of this memoir about that strange interlude, he with an expression of remorse, she with a trace of lingering resentment, but each realizing that their sacrifice had helped to put Evans on the way to professional achievement. After about a year, the birth of their daughter Marian in the Johns Hopkins Hospital ended the deception and Anabel's isola- tion, although as the wife of an impecunious and intensely busy student her'lot was still far from easy. During his medical year at Berkeley, Evans had taken the usual course in human anatomy with dissection. At Johns Hopkins, at that time, second-year students were given con- tinuing instruction in that subject, but the professorFranklin P. Mall, a shrewd and subtle judge of menallowed Evans to spend his time in the laboratory more or less as he wished. Because Mall himself had engaged in research on the pattern of microscopic blood vessels in various organs and in embryos, Evans learned from him methods of injecting blood vessels with colored fluids to make them readily visible. Working with finer and finer glass cannulas, under the microscope, he became expert. His special status in the laboratory gave him much closer association with the departmental staff than he would otherwise have had. Mall, the most influential anatomist in the country, had gathered about him probably the strongest ana- tomical research group in the English-speaking world, including

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156 BIOGRAPHICAL MEMOIRS Warren H. Lewis, Ross G. Harrison, and Florence R. Sabin all of them destined, as was Mall, to become members of the National Academy of Sciences. One day the school's professor of surgery, the austerely scientific William H. Halsted, came to Mall for help with a clinical problem. A patient of his, following an operation on the thyroid gland. had developed acute tetany. Knowing of other surgical mishaps of the same kind, Halsted rightly con- jectured that he had inadvertently either removed the incon- spicuous but indispensable parathyroid glands along with the thyroid, or had tied off their blood supply. He felt that with more exact knowledge of the small arteries supplying the para- thyroids, operations could be planned to conserve the glandules. Mall arranged for Evans to take on the problem under Halsted's supervision. By injection and careful dissection of the branches of the thyroid arteries in a few cadavers, Evans solved the prob- lem, and thus his name appeared with Halsted's at the head of an authoritative little article that appeared in Annals of Surgery in 1907 while the junior author was still a medical student. In the same year Evans published in the American Journal of Anatomy a very creditable article on the blood circulation in the walls of large lymphatic vessels, illustrated with his own drawings made in the style of the great Johns Hopkins medical artist Max Broedel. About the same time he had begun research on embryonic blood vessels. Before taking his medical degree he completed a study of the earliest vessels in the limb buds of chick embryos, which he published in full in 1909. Another significant accomplishment while a medical student was his demonstration of the growth of lymphatic vessels into a malignant tumor, pub- lished in 1908. While Evans was thus exhibiting his remarkable talent for anatomical investigation, Franklin Mall missed no occasion to foster his training for a professional career. He not only en- couraged Evans's research, but had him write book reviews for

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HERBERT MCLEAN EVANS 157 the Anatomical Record, then being published from Mall's labo- ratory, and finally gave him a quite extraordinary opportunity for so young a man. In 1907, Mall was organizing, jointly with Franz Keibel of Freiburg, Baden, Germany, their great Manual of Human Embryology (1910, Philadelphia and Leipzig), to be written by leading investigators of Germany and America. When a European embryologist who was to contribute a section on the blood-vascular system was unable to do so, Mall entrusted the task to Evans. Too ambitious merely to compile what was already known, Evans prepared himself for the assignment not only by studying the literature and then examining well-under- stood human embryos in Mall's collection but also by making original observations on the little-known earliest development of the aorta and the other great vessels. By extremely skillful injection of chick embryos, working under the microscope, he provedagainst the supposition of Hochstetter and othersthat these ultimately large channels begin, as do the peripheral arteries and veins, as a network of capillaries. This fundamental observation was the basis of Evans's authoritative contribution to the Keibel-Mall Manual, Chapter X\7III, Section III. From the first, Evans felt little interest in the clinical courses at Johns Hopkins, particularly when they took his time from such exciting activities as injecting embryonic blood vessels. He cut classes without regard to consequences. A preposterous story got abroad that he was granted his medical diploma only on his promise that he would never practice medicine. He him- self contributed to the persistence of such legends by his own, equally apocryphal statements that he was expelled from the medical school at the end of his third year for incompetence in surgical bandaging, obstetrical manikin exercises, and prescrip- tion-writing and was only restored to academic status with the help and advice of Dr. William H. Howell, then dean of the school. No doubt he did neglect such practical routines, but in ~ Bulletin of the Johns Hopkins Hospital, 6S, 300 (~947) .

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158 BIOGRAPHICAL MEMOIRS sober fact the records of the medical school do not mention any disciplinary action ever taken with respect to H. M. Evans, and he was graduated in medicine in 1908. Immediately afterward he joined Mall's department as assistant in anatomy and by this step confirmed, as his father had been fearing for some years, that he would not return to Modesto to assist with and later take over the older man's practice. To the senior Dr. Evans this was a heavy blow, for he thought practice far more important than research. It was undoubtedly the feeling that his father undervalued his choice of career that instilled in Herbert Evans an urgent desire to impress his parents by success in his chosen work and in later life to win the highest academic honors. While Evans remained at Johns Hopkins, Mall arranged several times for him to go to Germany in the summer vacations. During a stay at Freiburg he was fascinated by the novel experi- ments of the surgeon Erwin Goldman on intravitam staining of animal tissues by acid azo dyes (e.g., trypan blue). With a young Freiburg chemist, Werner Schulemann, he began experiments to find out whether the spectacular coloration, inside and out, of living rats, mice, and rabbits by injection of such dyes is a physical or chemical phenomenon. Continuing the work for some years, he found that these dyes do not truly dissolve in the body fluids, but are dispersed as extremely fine particles, form- ing a fluid suspension. When injected under a rat's skin, the dyestuff gets into the bloodstream as submicroscopic aggregates that the macrophage cells of connective tissues take up and then store in their cytoplasm. The recognition of this important class of cells and their role in the storage of particulate matter was much forwarded by Evans's work. Among the numerous dyes he and Schulemann studied, one now called Evans blue proved ~ Information kindly supplied by Mary E. Foy, Registrar, School of Medicine, Johns Hopkins University.

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HERBERT MCLEAN EVANS 159 useful in a method of measuring the blood volume of living ani- mals and human surgical patients. As a member of Mall's staff, Evans was busy with teaching, with his work on vital staining, and with studying the blood vessels of pig and chick embryos as well as of human embryos on the rare occasion when one was received in a sufficiently fresh state to be injected with indict ink. In 1913, on obtaining funds from the Carnegie Institution of Washington for the support of his large collection of human embryos and to develop research in that field, Mall created the department of embryology of the Carnegie Institution. This was first housed in the Johns Hop- kins anatomical laboratory and later in part of an adjacent new building. Evans, who was given a Carnegie appointment con- currently with his Johns Hopkins post, then devoted a good deal of his time to the sectioning of early human embryos and to reconstructing them in wax from the sections"A wearisome thing to do," he said, "compared with making the living embryo pump indict ink as though it were blood to show the multi- tudinous vascular channels." This strong inclination to experimental rather than purely morphological research was a partial cause of Evans's dropping a major project that Mall had suggested, a descriptive study of the human embryo during the period of somite formation. Another reason was that when Evans finally deft Baltimore, Mall was unwilling to let him take along, even for his temporary use, the rare and precious serially sectioned embryos necessary for the studyone or two of which, at least, Evans had himself collected and laboriously sectioned during Mall's summer absences. Evans was disappointed and hurt by what he regarded as his chief's ungenerosity. However, a few years later when Mall died in the prime of life with the rift between them still unhealed, Evans was deeply grieved. As a kind of penance for his part in the disagreement, he proposed to write a biography of Mall, hinting

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160 BIOGRAPHICAL MEMOIRS that this would be a profound analysis of a distinguished scien- tific mind (as indeed it might well have been), but in time the plan was forgotten. Evans's work on the embryos, however, was not lost. Several years later G. W. Bartelmez of the University of Chicago took up the study of the somite stage of human develop- ment, first studying the Carnegie embryos in Baltimore, then going to Berkeley to secure Evans's collaboration and the use of his notes and drawings. The result was an important mono- graph in the Carnegie Contributions to Embryology (~1926' under their joint authorship. In 1915, when Evans was in his thirty-third year, President Benjamin Ide Wheeler of the University of California offered him the chair of anatomy at Berkeley vacated by the departure of Flint in 1907 and of his acting successor, Irving Hardesty, in 1909. Since then, the direction of the department had reverted to the worthy pedagogue Robert O. Moody, under whom Evans a dozen years before had studied gross human anatomy. Research had practically ceased. Even routine teaching had suffered because of Moody's frail health and the illness of another mem- ber of the small staff. Philip E. Smith was the only man left in the department who was in good health and had sufficient experience to teach gross and microscopic anatomy. During the year before Evans's advent, Smith and his wife, who had done some postgraduate work in biology, had carried almost the entire teaching load, to the detriment of the research program in experimental embryology that Smith had brought with him from Cornell. To build up the department, Evans took with him from Johns Hopkins two young people who had shown competence for anatomical researchKatherine l. Scott (now Katherine Scott Bishop), a medical graduate of 1915, and George W. Corner, who was just completing an internship in the women's clinic. "Gynecologists ought to know more about the female re- productive cycle," said Dr. Evans to Corner. "Come to Berkeley

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HERBERT MCLEAN EVANS 161 with me and do your gynecology for a while on rats and rabbits." At Berkeley, recognizing Philip Smith's great abilities, Evans ar- ranged for him a much-lightened teaching schedule and in every possible way facilitated the research that ultimately won for Smith an international reputation and the chair of anatomy at Columbia University. It was during this early period that Smith perfected his operation of hypophysectomy in the rat, which became an invaluable procedure in research on the pituitary gland. Associate Professor Moody, perhaps a little surprised by the inrush of all this youthful enthusiasm, retained charge of gross anatomy, with Smith and one of the newcomers helping him. With the staff thus fully manned, the reorganized depart- ment resumed its work in the autumn of 1919 in a small frame building, once the university's printing shop, that had been adapted for the teaching of human anatomy when the San Francisco earthquake and fire of 1906 forced the transfer of pre- clinical classes to Berkeley. Evans took teaching quite seriously, in his own way, which was tinctured with the pride of intellect he never concealed. His course in histology was radically new in its extensive use of fresh and experimentally prepared tissues along with the traditional fixed and routinely stained sections. In the class laboratory he was usually to be seen at the micro- scope beside one of the better students. With the general run he was tolerant; with the duller minds, barely so and occasionally sarcastic. He did not believe in lecturing on gross anatomy, a finished science, but his lectures on microscopic anatomy were superb from the standpoint of his staff, for whom they consti- tuted a postgraduate course. As for the medical students, he was heard to say that he aimed his lectures at only the four or five best students in the class (of forty), tacitly implying that the assistant professors and instructors could take care of the rest. For the best ten percent the instruction (or, it might be more correct to say, the freedom to learn) provided by this brilliant

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162 BIOGRAPHICAL MEMOIRS professor opened new vistas in medical science. Years later Elmer Belt, a member of the first class Evans taught at Berkeley and now California's most distinguished urologist, wrote of Evans and his young associates that "The effect of their scholar- ship and idealism upon the freshman class in medicine was electric. Each of us realized how great an opportunity it was to enter the study of medicine under their guidance and for us the study of medicine became an obsession. The routine work of gross dissection and histology was time-consuming but most of us, in addition, were stimulated to take up a separate problem in research. We were thus led to seek out and read recent con- tributions to the literature concerned with our special subjects. This pursuit inevitably led us to doubt didactic textbook state- ments unless verified by our own personal observations. This atmosphere of doubt and verification prevailed through the department and led to intense application. For most of us this was our first taste of scholarly research." ~ Departmental administration was for Herbert Evans a duty reluctantly borne. His compulsive urge to work intensively at research led him to put off administrative routine, the writing of articles against deadlines, and other less congenial tasks until the last minute. Thus the course of departmental affairs was in- terrupted from time to time by minor or even major crises. One of these in the early Berkeley days, somewhat mysterious to Evans's associates, evidently caused him deep concern. He and his secretary for several days were intently busy, occupied with account books and the adding machine. Evans's brother, a busi- nessman familiar with accounting, was called in; there were urgent messages to and from the university bursar's office. Prob- ably the professor had overrun his budget. On another occasion he was overtaken by the deadline for an article long promised ~ Elmer Belt in There Was Light: Autobiography of a University, Berkeley, 1868-1969, ed: Irving Stone, Doubleday & Co., Garden City New York, 1970, p. 354.

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HERBERT MCLEAN EVANS 183 With K. Meyer, M. E. Simpson and F. L. Reichert. Disturbance of carbohydrate metabolism in normal dogs injected with the hypophyseal growth hormone. Proc. Soc. Exp. Biol. Med., 29:85- 88. 1932 Vitamin E. l. Am. Med. Assoc., 99:469-75. With Samuel Lepkovsky. Vital need of the body for certain un- saturated fatty acids. I. Experiments with fat-free diets in which sucrose furnishes the sole source of energy. l. Biol. Chem., 96: 143-56. 1933 With M. E. Simpson and P. R. Austin. The hypophyseal substance giving increased gonadotropin effects when combined with prolan. journal of Experimental Medicine, 57:897-906. With K. Meyer, M. E. Simpson, A. l. Szarka, R. I. Pencharz, R. E. Cornish and F. L. Reichert. The growth and gonad-stimulating hormones of the anterior hypophysis. Mem. Univ. Calif., 11:446 PP 1934 First editions in the history of science. In: Exhibition of First Edi- tions of Epochal Achievements in the History of Science. Spon- sored by the History of Science Club, University of California, Berkeley. (Pamphlet, privately printed) With S. Lepkovsky and Elizabeth A. Murphy. Vital need of the body for certain unsaturated fatty acids. IV. Reproduction and lactation upon fat-free diets. i. Biol. Chem., 106:431-40. With S. Lepkovsky and Elizabeth A. Murphy. Vital need of the body for certain unsaturated fatty acids. VI. Male sterility on fat-free diets. l. Biol. Chem., 106:445-50. 1935 With M. E. Simpson and R. I. Pencharz. "Deficiency" changes in the testicular Leydig cells after hypophysectomy. Anat. Record, 61 (abstr. supp.) :44.

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184 BIOGRAPHICAL MEMOIRS With Miriam E. Simpson. Production of superovulation in normal immature rats by injection of the principle in menopause urine. Proc. Soc. Exp. Biol. Med., 32:1946~7. 1936 With Oliver H. Emerson and Gladys A. Emerson. The isolation from wheat germ oil of an alcohol, <~-tocopherol, having the properties of vitamin E. l. Biol. Chem., 113: 319-32. With Karl Korpi, M. E. Simpson, R. I. Pencharz and D. H. Wonder. On the separation of the interstitial cell-stimulating, luteinizing and follicle-stimulating fractions in the anterior pituitary go- nadotropic complex. University of California Publications in Anatomy, 1 :255-73. 1937 With Clara L. Kohls and D. H. Wonder. Gonadotropic hormone in the blood and urine of early pregnancy; the normal occurrence of transient extremely high levels. l. Am. Med. Assoc., 108:287- 89. With Nellie Halliday. On the fractionation of the vitamin B2 com- plex from various source materials. I. Biol. Chem., 118:255-67. With M. E. Simpson and R. I. Pencharz. An anterior pituitary gonadotropic fraction (ICSH) specifically stimulating the inter- stitial tissue of testis and ovary. Cold Spring Harbor Symposium on Quantitative Biology, 5:229~0. lg38 With Dwight J. Ingle and H. D. Moon. Work performance of hypophysectomized rats treated with anterior pituitary extracts. Am. J. Physiol., 123:620-24. The hypophyseal growth hormone- Its separation from the hor- mones stimulating the thyroid, gonads, adrenal cortex and mam- mary glands. Research Publications of the Association for Research in Nervous and Mental Disease (Proceedings, 1936), 17: 175-92. 1939 With O. H. Emerson, G. A. Emerson, Lee Irvin Smith, Herbert E. Ungnade, W. W. Prichard, F. L. Austin, H. H. Hoehn, J. W.

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HERBERT MCLEAN EVANS 185 Opio and S. Wawzonek. The chemistry of vitamin E. XIII. Specificity and relationship between chemical structure and vitamin E activity. Journal of Organic Chemistry, 4:376-88. 1940 With Nobuko Shimotori and G. A. Emerson. The prevention of nutritional muscular dystrophy in guinea pigs with vitamin E. J. Nutr., 19: 547-54. With i. Fraenkel-Conrat, Heinz Fraenkel-Conrat and M. E. Simp- son. Purification of thyrotropic hormone of the anterior pitui- tary. l. Biol. Chem., 135:199-212. With H. Fraenkel-Conrat and M. E. Simpson. Purification of the follicle stimulating hormone of the anterior pituitary. Anales de la Facultad de Medicina de Montevideo, 25: 617-26. Re- printed in: Anales de la Facultad de Medicina de Montevideo: numero especial con motive del homenaje que la "Sociedad de Biologia de Montevideo" ofrece al Prof. Ludwig Fraenkel, en occasion del 70. aniversario de su nacimiento, pp. 159-68. With H. L. Fraenkel-Conrat, Donald L. Moamber and M. E. Simp- son. Further purification of the growth hormone of the anterior pituitary. Endocrinology, 27: 605-13. With H. Fraenkel-Conrat, Choh Hao Li and M. E. Simpson. In- terstitial cell stimulating hormone. I. Biological properties. Endocrinology, 27: 793-802. N\lith Choh Hao Li and M. E. Simpson. Interstitial cell stimulating hormone. II. Method of preparation and some physico-chem- ical studies. Endocrinology, 27: 803-8. With H. Fraenkel-Conrat and M. E. Simpson. Interstitial cell stim- ulating hormone. III. Methods of estimating the hormonal content of pituitaries. Endocrinology, 27: 809-17. With M. E. Simpson and William R. Lyons. Influence of lactogenic preparations on production of traumatic placentoma in the rat. Proc. Soc. Exp. Biol. Med., 46:586-90. 1941 With W. R. Lyons and M. E. Simpson. Influence of lactogenic preparations on mammary glands and time of vaginal opening in young rats. Proc. Soc. Exp. Biol. Med., 48:634-37.

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186 BIOGRAPHICAL MEMOIRS With Hermann Becks, Edwin A. Kibrick and Walter Marx. The early effect of hypophysectomy and of immediate growth hor- mone therapy on endochondral bone formation. Growth 5:449- 56. 1942 With W. Marx and M. E. Simpson. Bioassay of the growth hor- mone of the anterior pituitary. Endocrinology, 30:1 With H. L. Fraenkel-Conrat, V. V. Herring and M. E. Simpson. Effect of purified pituitary preparations on the insulin content of the rat's pancreas. Am. I. Physiol., 135:404. With M. E. Simpson, E. A. Kibrick and H. Becks. Effect of crystal- line estrin implants on the proximal tibia and costochondral junction of young female rats. Endocrinology, 30:286. With W. Marx, M. E. Simpson and W. O. Reinhardt. Response to growth hormone of hypophysectomized rats when restricted to food intake of controls. Am. l. Physiol., 135:614. With W. Marx and M. E. Simpson. Synergism between thyrotropic and growth hormones of pituitary. Body weight increase in hypophysectomized rat. Proc. Soc. Exp. Biol. Med., 49:594. With A. Gorbman. Urinary gonadotrophins in normal men. Proc. Soc. Exp. Biol. Med., 49:674. With M. E. Simpson and G. H. Li. Biological properties of pitui- tary interstitial cell-stimulating hormone (ICSH). Endocrinology, 30:969. With E. A. Kibrick, M. E. Simpson and H. Becks. Effects of crystal- line estrin implants on the tibia of young hypophysectomized female rats. Endocrinology, 31:93. With C. H. Li and M. E. Simpson. Isolation of adrenocortico- tropic hormone from sheep pituitaries. Science, 96:450. 1943 With M. E. Simpson, W. Marx and E. Kibrick. Bioassay of the pituitary growth hormone. Width of the proximal epiphyseal cartilage of the tibia in hypophysectomized rats. Endocrinology, 32:13. With A. Gorbman. Beginning of function in the thyroid of the fetal rat. Endocrinology, 32: 1 13.

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HERBERT MCLEAN EVANS 187 With W. R. Lyons and M. E. Simpson. Hormonal requirements for pregnancy and mamary development in hypophysectomized rats. Proc. Soc. Exp. Biol. Med., 52:134. With W. Marx, M. E. Simpson and C. H. Li. Antagonism of pitui- tary adrenocorticotropic hormone to growth hormone in hypo- physectomized rats. Endocrinology, 33: 102. With C. H. Li and M. E. Simpson. Adrenocorticotropic hormone. I. Biol. Chem., 149:413. With M. E. Simpson and C. H. Li. Inhibiting effect of adrenocor- ticotropic hormone on the growth of male rats. Endocrinology, 33:237. With M. E. Simpson, C. H. Li and W. O. Reinhardt. Similarity of response of thymus and lymph nodes to administration of adre- nocorticotropic hormone in the rat. Proc. Soc. Exp. Biol. Med., 54:135. With M. E. Simpson and C. H. Li. Bioassay of adrenocorticotropic hormone. Endocrinology, 33:261. With G. A. Emerson. The prophylactic requirement of the rat for alpha tocopherol. J. Nutr., 26:555. 1944 With Gladys A. Emerson. The bioassay of vitamin E. l. Nutr., 27:469. With H. Becks, M. E. Simpson, W. Marx and C. H. Li. Antagonism of pituitary adrenocorticotropic hormone (ACTH) to the action of growth hormone on the osseous system of hypophysectomized rats. Endocrinology, 34:311. With M. E. Simpson, W. Marx and H. Becks. Response to acre nalectomized-hypophysectomized rats to the pituitary growth hormone. Endocrinology, 35:234. With Miriam E. Simpson, W. Marx and H. Becks. Effect of testosterone propionate on the body weight and skeletal system of hypophysectomized rats. Synergism with pituitary hormone. Endocrinology, 35:309. 1945 With C. H. Li and M. E. Simpson. Isolation and properties of the anterior hypophyseal growth hormone. i. Biol. Chem. 159:353.

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188 With G. S. Gordan, C. H. Li BIOGRAPHICAL MEMOIRS 1946 and L. L. Bennett. Effect of adreno- corticotrophic hormone on urinary nitrogen excretion in the normal rat. Proc. Soc. Exp. Biol. Med., 62:103. With M. E. Simpson. Sensitivity of the reproductive system of hypophysectomized forty-day-old male rats to testosterone pro- pionate. Endocrinology, 39:75. With M. E. Simpson. Comparison of the spermatogenic and andro- genic properties of testosterone propionate with those of pituitary ICSH in hypophysectomized 40-day-old male rats. Endocrinol- gY, 39:281. 1947 With Marjorie M. Nelson. Growth, reproduction and lactation in the rat maintained on purified diets. Archives of Biochemistry, 12:213. Recent progres de nos connaissances sur les hormones du lobe anterieur de l'hypophyse. Journal de Physiologic (Paris), 39: 121. With C. H. Li. The properties of the growth and adrenocortico- tropic hormones. Vitamins and Hormones, 5:197. 1948 With Miriam E. Simpson and C. H. Li. The gigantism produced in normal rats by injection of the pituitary growth hormone. I. Body growth and organ changes. Growth, 12:15. With A. A. Koneff, M. E. Simpson and C. H. Li. The gigantism produced in normal rats by injection of the pituitary growth hormone. II. Histological changes in the pituitary. Growth, 12:33. With C. H. Li and M. E. Simpson. The gigantism produced in normal rats by injection of the pituitary growth hormone. III. Main chemical components of the body. Growth, 12:39. With H. Becks, C. Willet Asling, M. E. Simpson and C. H. Li. The gigantism produced in normal rats by injection of the pituitary growth hormone. IV. Skeletal changes: Tibia, costochondral junction, and caudal vertebrae. Growth, 12:43. With H. Becks, D. A. Collins, C. W. Asling, M. E. Simpson and C. H. Li. The gigantism produced in normal rats by injection

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HERBERT MCLEAN EVANS 189 of the pituitary growth hormone. V. Skeletal changes: Skull and dentition. Growth, 12:55. With C. H. Li. Chemistry of anterior pituitary hormones. In: The Hormones, ed. by G. Pincus and K. V. Thimann, Vol. I, p. 631. New York, Academic Press, Inc. 1949 With F. S. Greenspan, C. H. Li and M. E. Simpson. Bioassay of hypophyseal growth hormone: The tibia test. Endocrinology, 45:455. The search for the diabetogenic principle of the anterior hypophysis. (The Banting Memorial Address y Proceedings of the American Diabetes Association, 9:49. 1950 With H. D. Moon, M. E. Simpson and C. H. Li. Neoplasm in rats treated with pituitary growth hormone. Cancer Res., 10:364. With H. D. Moon, M. E. Simpson and C. H. Li. Neoplasms in rats treated with pituitary growth hormone. III. Reproductive or- gans. Cancer Res., 10:549. With M. E. Simpson. Physiology of the gonadotrophins. In: The Hormones, ed. by G. Pincus and K. V. Thimann, Vol. II, Chap. VI, p. 351. New York, Academic Press, Inc. The hypophysis and diabetes mellitus. In: The Hormones, ed. by G. Pincus and K. V. Thimann, Vol. II, Chap. VIII, p. 405. New York, Academic Press, Inc. With M. E. Simpson and C. Willet Asling. Some endocrine in- fluences on skeletal growth and differentiation. Yale Journal of Biology and Medicine, 23:1. With D. C. Van Dyke, M. E. Simpson and C. H. Li. Survival in the circulation of the growth and adrenocorticotrophic hormones as evidenced by parabiosis. Am. J. Physiol., 163:297. 1951 With A. A. Koneff, H. I). Moon, M. E. Simpson and C. H. Li. Neo- plasms in rats treated with pituitary growth hormone. IV. Pitui- tary gland. Cancer Res., 1 1 : 1 13. With Marjorie M. Nelson. Effect of pyridoxine on reproduction of the rat. J. Nutr., 43: 281.

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190 BIOGRAPHICAL MEMOIRS With H. D. Moon, M. E. Simpson and C. H. Li. Neoplasms in rats treated with pituitary growth hormone. V. Absence of neo- plasms in hypophysectomized rats. Cancer Res., 11:535. With Miriam E. Simpson and C. H. Li. Synergism between pitui- tary follicle stimulating hormone (FSH) and human chorionic gonadotropin (HCG). Endocrinology, 48:370. With C. A. Gemzell, D. C. Van Dyke and C. A. Tobias. Increase in the formation and secretion of ACTH following adre- nalectomy. Endocrinology, 49:325. 1952 With H. D. Moon and M. E. Simpson. Inhibition of methyl- cholanthrene carcinogenesis by hypophysectomy. Science, 116: 331. With M. M. Nelson and C. W. Asling. Production of multiple congenital abnormalities in young by maternal pteroylglutamic acid deficiency during gestation. J. Nutr., 48:61. With D. C. Walker, C. W. Asling, M. E. Simpson and C. H. Li. Structural alterations in rats hypophysectomized at six days of age and their correction with growth hormone. Anat. Record, 1 14:49. With C. W. Asling, D. G. Walker, M. E. Simpson and C. H. Li. Deaths in rats submitted to hypophysectomy at an extremely early age and the survival effected by growth hormone. Anat. Record, 1 14: 19. With A. A. Koneff and D. C. Van Dyke. Increase in the thyrotropic hormone content of blood after thyroidectomy as shown by parabiosis. Endocrinology, 51:249. 1953 With Miriam E. Simpson, D. C. Van Dyke and C. W. Asling. Re- generation of the calvarium in young normal and growth hor- mone-treated hypophysectomized rats. Anat. Record, 115:615. With M. M. Nelson and W. R. Lyons. Comparison of ovarian and pituitary hormones for maintenance of pregnancy in pyri- doxine-deficient rats. Endocrinology, 52: 585. With M. M. Nelson. Relation of dietary protein levels to repro- duction in the rat. J. Nutr., 51:71.

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HERBERT MCLEAN EVANS 191 With R. A. Lyon and M. E. Simpson. Qualitative changes in urinary gonadotrophins in human pregnancy during the period of rapid increase in hormone titer. Endocrinology, 5 3:674. 1954 With R. D. Ray, C. W. Asling, D. G. Walker, M. E. Simpson and C. H. Li. Growth and differentiation of the skeleton in thy- roidectomized-hypophysectomized rats treated with thyroxin, growth hormone, and the combination. journal of Bone and Joint Surgery, 36: A94. With Marjorie M. Nelson. Maintenance of pregnancy in the ab- sence of dietary protein with estrone and progesterone. En- docrinology, b5:543. With Catherine D. C. Baird, M. M. Nelson and I. W. Monie. Con- genital cardiovascular anomalies induced by pteroylglutamic acid deficiency during gestation in the rat. Circulation Research, 2:544. 1955 With Eloise Wooten, M. M. Nelson and M. E. Simpson. Effect of pyridoxine deficiency on the gonadotrophin content of the an- terior pituitary in the rat. Endocrinology, 56:59. With M. M. Nelson. Relation of thiamine to reproduction in the rat. J. Nutr., 55:151. With C. W. Asling, M. E. Simpson, H. D. Moon and C. H. Li. Growth hormone induced bone and joint changes in the adult rat. In: The Hypophyseal Growth Hormone, Nature and Action, ed. by R. W. Smith, ir., et al., Chap. 9, p. 154. New York, McGraw-Hill Book Co., Inc. With M. M. Nelson, H. V. Wright and C. W. Asling. Multiple congenital abnormalities resulting from transitory deficiency of pteroylglutamic acid during gestation in the rat. I. Nutr., 56:349. 1956 With C. W. Asling. Anterior pituitary regulation of skeletal de- velopment. In: The Biochemistry and Physiology of the Bone, ed. by G. H. Bourne, Chap. 21, p. 671. New York, Academic Press, Inc.

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192 BIOGRAPHICAL MEMOIRS 1958 Title E. Mrooten, NI. AI. Nelson and M. E. Simpson. Response of vitamin B6-deficient rats to hypophyseal follicle-stimulating and interstitial-cell-stimulating hormones. Endocrinology, 63:860. 1959 With M. M. Nelson. Dietary requirement for lactation in the rat and other laboratory animals. In: Milk: Its Physiology and Bio- chemistry, ed. by A. T. Cowie and S. K. Kon. New York, Academic Press, Inc. Editor. Men and Moments in the History of Science. Anniversary volume commemorating founding of History of Science Dinner Club, University of California campus, Sept. 12, 1933. Seattle, University of Washington Press. 226 pp.

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