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WILLIAM HAY TALIAFERRO February 10, 1895-December 21, 1973 BY DAVID W. TALMAGE WILLIAM HAY TALIAFE~O was born prematurely on February 10, IS95 in Portsmouth, Virginia. His doctor consiclered his survival remarkable and gave the credit to his mother, Mary Watkins Leigh, for her solicitous care. He was a member of the ninth generation of Virginia Taliaferros and was descended from Robert (162~1688~. The name is Eng- lish, and it is pronounced Toliver. Both sides of his family were Virginia aristocrats who had become impoverished by the Civil War. Both grandparents were physicians, but his father was barred from the profes- sion by a hunting accident that resulted in the loss of his right hand. With this background it is not surprising that William's boyhood! ambition was to be a physician. William attended public school in Portsmouth until the age of ten. He then went to Sunnyside Seminary, near CIarks- ville, Virginia, for three years. This finishing school for girls was run by three Carrington great-aunts who permitted Wil- liam and several other boys to attend as clay students. The boys added zest to the education of the girls by bringing snakes, frogs, bugs, and other novelties into the classroom. As far as his studies were concerned, William excelled in arith- metic, but he never couIct spell, although he took to 01d English and enjoyed looking up the derivation of words. 375

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376 BIOGRAPHICAL MEMOIRS Once he confided to Karl Lashiey, his closest friend at Hop- kins and thereafter, that he never knew whether "separate" had 2 e's or 2 a's in its middle, whereupon Karl replied that he never couicI decide whether "which" had a t in it or not. In 190S, at the age of thirteen, William received a schol- arship to Norfolk Academy in Norfolk, Virginia. This private preparatory school had a distinguished history. There he was grilled in the oIcI-fashionecT studies of mathematics, English, and history. He took all the science that was offered, but this was limitect to the most elementary physics and chemistry. He was not interested in sports and spent his free time reading. He received the Class Prize in 1910 and the Ingram Prize in 1911. William was shy and timid as a boy and was protected at times from the school bullies by a stronger older brother. This shyness carried over into adulthood as a remarkable gentleness that en~learect him to his friends and students. Shyness did not suppress an early resourcefulness. To assure funds for his later education, William sold eggs from a pedi- greed flock of hens. He devised an ingenious method of feeding them during classroom hours at Norfolk Academy. He turned an alarm clock on its side and balanced a weight on the clapper. At noon, the weight dropped anti upset a box of feed. He always remembered those Plymouth Rocks with pleasure because of "their upturned heals and alert eyes around noontime." Unfortunately, he later developed an allergy to many animal dancers that kept him out of the animal quarters. From the fall of 1911 through the spring of 1915 William attencled the University of Virginia. In his junior and senior years, he had teaching assistantships under Dr. William A. Kepner, who not only took him on innumerable f~elcI trips ~ W. H. Taliaferro, Annual Review of Microbiology, 1968.

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WILLIAM HAY TALIAFERRO 377 and gave him valuable teaching experience but replaced his ambition to study medicine with a lifelong love of zoology. Zoology must have been a little-known profession in those days. When William announced at home that he had decided to be a zoologist, his brother remarked, "Good Lord, William, are you going to be a zookeeper?"2 While at the University of Virginia, he published three papers with Dr. Kepner on the sensory epithelium of Microstoma caudatum, the organs of special sense in Prorhynchus applanatus, and the reactions of Amoeba proteus to food. William graduated from the Univer- sity in 19 ~ 5 with a bachelor of science degree after election to Phi Beta Kappa. He was invited to join the Raven Society of the University in 1944. While at the University of Virginia, William was im- pressed with a book by Dr. Herbert S. Jennings of the Johns Hopkins University, entitled Behavior of the Lower Organisms. He determined to go to Hopkins and study under Jennings. When he arrived there in the fall of 1915, he found that Jennings had changed his field to genetics and was placed under the guidance of Dr. Samuel Mast. Nevertheless, he attended all the lectures given by Dr. Jennings and those by Dr. Burton Livingston on plant physiology. From the latter he learned to use porous filters that were useful in his re- search on antibodies to trypanosomes. William's training in genetics led to the publication of a paper with John Huck on the inheritance of the sickling of human red cells in several Negro families. He and Huck concluded that this phenomenon, later called the sickle-cell trait, is inherited as a single, Mendelian dominant gene (S) that is not sex-linked. Others later showed, on the one hand, that the homozygote recipients of two S genes suffer from severe sickle-cell anemia, and, on the other, that the heterozy- 2 Ibid.

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378 . BIOGRAPHICAL MEMOIRS gote carriers of only one S gene demonstrate the sickle-cell trait in vitro, are free of the disease, and are resistant to the lethal effects of the virulent African malarial parasite, Plasmo- ctiumfalciparum. This disease-host relationship has become a model for the stucly of genetics and evolution because it shows how an unfavorable gene may be advantageous under particular conclitions. While at Hopkins, William joined the Beta Theta Pi Fraternity. The boys christened him "folly," an affectionate name he carried the rest of his life. He was never called Bill. The boys also taught him to play poker and bridge and took him to the Saturday burlesque shows at the old Holiciay Street Theatre. It must have been there that he started to accu- mulate the vast stock of stories for which he was famous. In the summer of 1917, William was invited to assist in a course on invertebrate zoology at Woods Hole, Massachusetts. There he met Lucy Graves, a Yankee of AIay- flower descent, who later became his wife. She had just graduates! from Goucher College with a bachelor of arts degree after election to Phi Beta Kappa. Since William anti LACY were each engaged to someone else at the time, they thought it safe ant! pleasant to date each other. In any case. the Goucher table where Lucy was a student must have re- ceived an unequa "l share of the young instructor's time be- cause Dr. Allee is quoted as remarking one day, "Taliaferro, in previous years, the Goucher bunch have always seemed intelligent, but this year they seem to need all your attention while I have to instruct the rest of the ciass."3 Tolly volunteered for service in the army in the fall of 1917 and was assigned to the Division of Chemical Warfare at Yale. While there he finished his thesis, passed his oral examinations, and received his Ph.D. from Hopkins in the ~ L. G. Taliaferro: personal communication.

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WILLIAM HAY TALIAFERRO 379 spring of HIS, at the age of twenty-three. This accelerated program was macle possible by his exceptional training at the Norfolk Academy (he was excused from freshman English and mathematics when he entered the University of Virginia) and continuous attendance at the University of Virginia summer and winter. His thesis was on reactions to light in Planaria macuitata, a small flatworm, with special reference to the function and structure of the eye. The experiments had been welI-planned ant! carried out at the University of Virginia. The technique anal analysis of the results were far in advance of their time. When Tolly and Lucy left Woods Hole in 1917, they never expecter! to see each other again, but they continued to cor- respond. Then, in HIS, Lucy, who was abstracting German articles on gases in the Department of Chemical Warfare, was transferred to New York City. Tolly, who was assisting in a research program on respiratory gases under Dr. Yande} Henderson at Yale, had to go to New York City frequently to get Pyrex glass blown for intricate assemblies of equipment. There he looked up Lucy to take her to dinner. His private's pay was stretched to capacity! Their friendship, however, blossomed to the extent that they became disengaged from their financees and engaged to each other. They were mar- ried in June ~9~9. In the spring of 1919, when his stint in the army endecT with the rank of second lieutenant, Tolly was awarciect a Johnston Scholarship at Hopkins, but declined it to accept a position under Dr. Robert Hegner in the Department of Protozoology anct Mectical Entomology in the newly estab- lished School of Hygiene anct Public Health at Hopkins. While there, the book by Hegner and Taliaferro on Human Protozoollogy appeared. Also, Lucy passed her oral ex- aminations and received her Sc.D. with a thesis on avian malaria. Thus, she became a full member of the Taliaferro

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380 BIOGRAPHICAL MEMOIRS team, and the names Taliaferro and Taliaferro appear on many of the more than 100 contributions to science that are mentioned here. They were a perfect teameach respected the other for what each did best. Lucy always said that Tolly breezed in on the express with ideas and questions, and she and one or two assistants arrived later on the freight loacled with experimental ciata involving injections, blood smears, bleeclings, autopsies, and tests galore. Then, as a team, they graphed, tabulated, arranged, rearranged, described, and summarized the results for publication. Usually all three pro- cesses were occurring simultaneously on different subjects: a manuscript was being readied for publication on old project one, experiments carried out on intermediate project two, and new ideas were being cireamed up for new project three. Some of the cireaming became sidetracked, delayed, or wrecked by sheer lack of time and energy. William and Lucy also bought a house in Roland Park, a suburb of Baltimore, expecting to be settled there for years. It was not to be. In 1924 William was invited by Dr. Edwin 0. Jordan to join the Department of Hygiene and Bacteriology at the University of Chicago. Three years later he was made a full professor. There he and Lucy remained for thirty-six years and clid the bulk of their life's work. During that time they lived in one apartment for eight years and in a second one for twenty-eight years! Their research activities centered around host-parasite relationships and the mechanism of antibody formation with hemolysin formation as a baseline. Although Tolly had never worked on parasites before, his newly found interest in genetics led him to study variability in Trypanosoma 1lewisi, a bloocl-inhabiting parasite of the rat. Fre- quent blood smears were made throughout infections lasting a month or more. Number counts and camera lucida draw- ings from such smears led him to the discovery that the rat not only develops lytic antibodies the parasite counts fell in

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WILLIAM HAY TALIAFERRO 38 about a week after an initial rise ant] disappeared later but also displays a peculiar property of inhibiting parasite re- production initial variability of the parasites fell near zero in ten days. Later, when he cletermined that this factor was an antibody, he collect it ablastin. He always considered this clis- covery his greatest achievement, and it was a recurring theme for study throughout his life. In fact, the last research he published in ~ 97 ~ with one of his students, Dr. P. A. D'Alesanciro, was on the effect of adenine on innate and acquired immunity in the rat to T. lewisi. Others, too, were intrigued by the subject. A workshop on ablastin was arranged by D'Alesandro and was held at Rockefeller Uni- versity on June 21 through 22, 1973. The resulting papers were published in Experimental Parasitology in 1975 (Vol. 38:303~91. Tolly and Lucy also studied other trypanosomes in mice, guinea pigs, rats, and dogs, but the mouse was the only host that yielded a reproduction-inhibiting ablastin against T. musculi (=T. duttoni). Tolly always had a yen to go to Africa with the hope that there he could study some indig- enous hosts of trypanosomes, but the opportunity never materialized. In any case, these relationships made a signifi- cant impact on Tolly's own research and lecl much later to his concentrated attack on hemolytic antibodies. The unraveling of such host-parasite relationships, com- bined with an orgy of reading, led to the production of The Immunology of ParasiticInfections, published in 1929. When Dr. Carroll Bull of the School of Hygiene at Hopkins, William's immunological consultant of those clays, read the manu- script, he commended it, but wrote that the chapters needed summaries! Such a suggestion, though justified, caused con- sternation anct the burning of much midnight oil. Once the summaries were completed, the Taliaferros took their first trip to Europe. They spent Palm Sunday in London, saw the tulips spread out in bloom like vast oriental

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382 BIOGRAPH I CAL M EMOI RS carpets in Holianct, and spent a month in Paris with daily trips to surrounding areas. The magnificent cathedrals, with their brilliant red and glowing blue stained-glass windows, were an inspiration. It was a long-rememberec! episode in their life, which they were not able to repeat until 1956. The research and the book brought Taliaferro wide recognition. He was invited to give the Harvey Lecture in 1931 and was elected an honorary member of the Harvey Society. He gave the Delamar Lecture at the School of Hygiene and Public Health in Baltimore in 1932. He was elected president of the American Society of Parasitologists in 1933 and received the Chalmers Mecial of the Royal Society of Tropical Medicine of Englanc! in 1935. From 1925 on, Tolly was also a member of the In- nominates, a collection of about thirty basic scientists at the University of Chicago. They hacl a clinger meeting once a month at the University of Chicago faculty Quadrangle Club for a talk by one of the members. Afterwards, the men met their wives at the home where the ladies had dinecI. During the work on the various trypanosome infections, Tolly was fascinated with the immunological problem of ctis- tinguishing between killing and reproduction-inhibiting ef- fects of innate and acquired immune reactions of the host. He and Lucy then began an intensive study of the parasites that cause various malarial. Examination of these blood- inhabiting plasmodia had two distinct advantages. Blooc! smears collie be user! not only to obtain number counts of a given infection, but, if taken often enough, wouIcl reveal the reproductive cycle because the parasites usually grow and segment synchronously. Such cycles are usually completecl every 24, 36, 4S, or 72 hours, clepending upon the species. The well-known periodic fevers of malaria occur at the time the plasmodia segment in the blood. Many were the evenings Tolly and Lucy carried home canaries, chickens, or monkeys

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WILLIAM HAY TALIAFERRO 383 ant! parked them in the bathtub in order to make blooct smears during the night. This exhaustive and time- consuming search was not very successful as far as the main object was concerned. Parasitic lysis occurred, but no reproduction-inhibiting effect was found in the malarial except for transient ones during the dramatic parasiticidal crises. The various malarial, however, proved to be invaluable for studying the cellular phases of immunity because plas- modial pigment remains for months as an indigestible marker in host phagocytic cells. This work was started with Dr. Paul Cannon and continued with Drs. William Bloom, Hugh Mulligan, and James MouIcler; all were at the Univer- sity of Chicago except Dr. Hugh Mulligan, who came from England by way of India. The onset of the enterprise was sparked by Dr. Alexander Maximow's work on inflammation following the subcutaneous injection of dyes or India ink. The malaria papers were beautifully illustrated by micro- scopic camera lucicla drawings, and praise for these should be accorded Esther BohIman Patterson, Tolly's artist from 1935 on. Examinations of various tissues from infected animals in- cTicatect that the plasmodia inhabiting blood cells are phago- cytize(1 by macrophages in strategically placed organs, such as the spleen, liver, and bone marrow, and that the macro- phages conspicuously increase in number, especially as acquired immunity develops, largely because of the hetero- plastic division and development of lymphocytes and mono- cytes into macrophages. Tolly became convinced that the baffling anal, up to this time, largely ignored lymphocytes were important in defense. In 1937 he and Hugh Mulligan coin`~1 the ~nuronriate term, lymphoid-macrophage system (LMS), to embrace all cells involves! in defense, including the lymphocytes. The term, reticulo-endothelial system (RES), Err- -r-

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384 BIOGRAPHICAL MEMOIRS however, had already been proposed ten years earlier by Aschoff to include cells in defense, although he expressly included lymphocytes. This term was so firmly entrenches] in the literature that it is still the preferred term, although it now needs the tacit assumption that lymphocytes are in- clucled since the role of lymphocytes in defense is well recog- nized and their nature is being intensively analyzecl. Taliaferro and Dr. Merrit P. SarIes also found a similar lymphocyte-macrophage relationship in rats experimentally injected subcutaneously with the small nematode worm, Nip- postrongylus brasiliensis. in this case the strategically located organs, as acquired immunity clevelopecl, are the skin, lungs, . . ancl intestine. From all this work and extensive readings Toliv arrived At +1~ ~ ~ ~ ~ _1 ~ 1 , 1 1 ~ - ~c coupon gnat cellular reactions in all hosts are similar and stereotyped, whether the invacler be living or nonliving, and they only cliffer superficially because of the invader's size, mocle of entrance, and subsequent location in the host. From 1926 through 1954, the Taliaferros took ten three- month trips for work at centers of tropical disease in Puerto Rico and Central ant! South America. Three of theirjourneys were to Panama for intensive work on malaria in monkeys. The Taliaferros always combined a great deal of pleasure with their work on these trips. They swam, played tennis, danced, rocle horseback, and crossed the Andes by bus and boat in the justly famous lake region of Chile. Their first airplane trip in 1954 involvecl crossing the Argentine from Bariloche to Buenos Aires. Publications, in addition to those previously mentioned, inclucled such subjects as a precipitin test in malaria ant! various tests on equine trypanosomiasis and helminth infections. The trips were subsidized by Dr. W. E. Deeks and the United Fruit Company, the Rockefeller Foundation, and the University of Chicago.

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WILLIAM HAY TALIAFERRO 397 Infection and immunity in bird malaria. P. R. I. Public Health Trop. Med., 4: 155-68. A note on the amoeba of the cockroach cultivated by Smith and garret. I. Parasitol., 14:274. The results of Schick tests in Tela, Honduras. l. Prev. Med., 2:213-17. With W. A. Hoffman and D. H. Cook. A precipitin test in intestinal schistosomiasis (S. mansoni). at. Prev. Med., 2:395-414. With L. G. Taliaferro. A precipitin test in malaria; Second report. }. Prev. Med., 2:147-67. 1929 With L. G. Taliaferro. Acquired immunity in avian malaria. I. Immunity to superinfection. I. Prev. Med., 3:197-208. With L. G. Taliaferro. Acquired immunity in avian malaria. II. The absence of protective antibodies in immunity to super- infection. {. Prev. Med., 3:209-23. The Immunology of Parasitic Infections. New York: The Century Com- pany. 414 pp. 1930 With W. A. Hoffman. Skin reactions toDirofilaria immitis in persons infected with Wuchereria bancrofti. at. Prev. Med., 4:261-80. 1931 With P. R. Cannon. Acquired immunity in avian malaria. III. Cellu- lar reactions in infection and superinfection. J. Prev. Med., 5:37-64. The mechanism of acquired immunity in avian malaria. South. Med. l., 24:409-15. With P. R. Cannon and S. Goodloe. The resistance of rats to infec- tion with Trypanosoma lewisi as affected by splenectomy. Am. I. Hyg., 14:1-37. With L. G. Taliaferro. Skin reactions in persons infected with Schistosoma mansoni. P. R. ]. Public Health Trop. Med., 7:23-35. 1932 Trypanocidal and reproduction-inhibiting antibodies to Trypano- soma lewisi in rats and rabbits. Am. J. Hyg., 16:32-84.

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398 BIOGRAPHICAL MEMOIRS Infection and resistance in the blood-inhabiting protozoa. Science, 75:619-29. Experimental studies on the malaria of monkeys. Am. J. Hyg., 16:429~9. 1934 With W. Bloom. A note on the granular leucocytes of the New World monkeys. In: Festschrift fur Prof. N. Anitshbow, Leningrad, pp. 27-30. Some cellular bases for immune reactions in parasitic infections. I. Parasitol., 20:149~1. With L. G. Taliaferro. Complement fixation, precipitin, adhesion, mercuric chloride and Wassermann tests in equine trypano- somiasis of Panama (murrina). }. Immunol., 26: 193-2 13. With L. G. Taliaferro. The transmission of Plasmodiumfalciparum to the howler monkey, Alouatta sp. I. General nature of the infec- tions and morphology of the parasites. Am. J. Hyg., 19:318-34. With P. R. Cannon. The transmission of Plasmodium falciparum to the howler monkey, Alouatta sp. II. Cellular reactions. Am. ]. Hyg., 19:335~2. With L. G. Taliaferro. Morphology, periodicity and course of infec- tions of Plasmodium brasilianum in Panamanian monkeys. Am. I. Hyg., 20: 1~9. With L. G. Taliaferro. Alteration in the time of sporulation of Plasmodium brasilianum in monkeys by reversal of light and dark. Am. J. Hyg., 20:50-59. With L. G. Taliaferro. Superinfection and protective experiments with Plasmodium brasilianum in monkeys. Am. I Hyg., 20:60-72. 1936 With Y. Pavlinova. The course of infection of Trypanosoma duttoni in normal and in splenectomized and blockaded mice. I. Parasitol., 22:29-41. With P. R. Cannon. The cellular reactions during primary infec- tions and superinfections of Plasmodium brasilianum in Pana- manian monkeys. I. Infect. Dis., 59:72-125. With M. P. Sarles. The local points of defense and the passive transfer of acquired immunity to Nippostrongylus muris in rats. J. Infect. Dis., 59:207-20.

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WILLIAM HAY TALIAFERRO 1937 399 With H. W. Mulligan. The Histopathology of Malaria with Special Refer- ence to the Function and Origin of the Macrophages in Defence, Indian Medical Research Memoir 29. Calcutta: Thacker, Spink Co. 138 pp. 1938 The effects of splenectomy and blockade on the passive transfer of antibodies against Trypanosoma lewisi. !,- Infect. Dis., 62:98-111. With W. Bloom. Regeneration of the malarial spleen in the canary after infarction and after burning. I. Infect. Dis., 63:54-70. Ablastic and trypanocidal antibodies against Trypanosoma duttoni. ]. Immunol., 35:303-28. 1939 With M. P. Sarles. The cellular reactions in the skin, lungs and intestine of normal and immune rats after infection with Nippo- strongylus muris. ]. Infect. Dis., 64: 157-92. 1940 The mechanism of immunity to metazoan parasites. Am. I. Trop. Med., 20:169-82. With L. G. Taliaferro. Active and passive immunity in chickens against Plasmodium lophurae. it. Infect. Dis., 66: 153~5. With C. G. Huff. The genetics of the parasitic protozoa. Am. Assoc. Adv. Sci., 12:57~1. The mechanism of acquired immunity in infections with parasitic worms. Physiol. Rev., 20:469-92. With C. Kluver. The hematology of malaria (Plasmodium brasilianum) in Panamanian monkeys. 1. Numerical changes in leucocytes. 2. Morphology of leucocytes and origin of mono- cytes and macrophages. I . I nfect. Dis ., 6 7: 1 2 1 - 7 6. 1941 The immunology of the parasitic protozoa. In: Protozoa in Biological Research, ed. G. N. Calkins and F. M. Summers, pp.830-89. New York: Hafner Publishing Company. The cellular basis for immunity in malaria. Am. Assoc. Adv. Sci., 15:239~9; 371-98. Populations of blood-dwelling species. Am. Nat., 75:458-72.

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400 BIOGRAPHICAL MEMOIRS 1942 With M. P. Sarles. The histopathology of the skin, lungs and . . ~ Intestine ot rats during passive immunity to Nippostrongylus muris. ]. Infect. Dis., 71 :69-82. 1943 Antigen-antibody reactions in immunity to metazoan parasites. Proc. Inst. Med. Chicago, 14:358~8. With Y. P. Olsen. The protective action of normal sheep serum against infections of Trypanosoma duttoni in mice. T Infect. Dis., 72:213-21. 1944 Malaria. In: Medicine and the War, ed. W. H. Taliaferro, pp. 55-75. Chicago: University of Chicago Press. Medicine and the War teds. Chicago: University of Chicago Press. 193 pp. Immunity in malaria. Am. I. Clin. Pathol., 14:593-97. With L G. Taliaferro. The effect of immunity on the asexual repro- duction of Plasmodium brasilianum. J. Infect. Dis., 75: 1-32. 1945 With W. Bloom. Inflammatory reactions in the skin of normal and immune canaries and monkeys after the local injections of ma- larial blood. I. Infect. Dis., 77:109-38. With L. G. Taliaferro and E. L. Simmons. Increased parasitemia in chicken malaria (Plasmodium gallinaceum and Plasmodium lophurae) following X-irradiation. I. Infect. Dis., 77: 158-76. With L. G. Taliaferro. Immunological relationships of Plasmodium gallinaceum and Plasmodium lophurae. ]. Infect. Dis., 77:224-48. 1947 The role of the lymphocyte in immunity with special reference to malaria. Rev. Kuba Med. Trop. Parasitol., 3: 150-51. With L. G. Taliaferro. Asexual reproduction of Plasmodium cyno- molgi in rhesus monkeys. J. Infect. Dis., 80:78-104. 1948 The inhibition of reproduction of parasites by immune factors. Bacteriol. Rev., 12: 1-17.

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WILLIAM HAY TALIAFERRO 40 With L. G. Taliaferro. Reduction in immunity in chicken malaria following treatment with nitrogen mustard. I. Infect. Dis., 82:5-30. The role of the spleen and the lymphoid-macrophage system in the quinine treatment of gallinaceum malaria. I. Acquired immunity and phagocytosis. J. Infect. Dis., 83:164-80. Science in the universities. Science, 108:145-48. Acquired immunity in malaria. Proceedings of the Fourth International Congress on Tropical Medicine and Malaria, vol. 1, pp. 776-82. Washington, D.C.: U.S. Government Printing Office. With F. E. Kelsey. The role of the spleen and the lymphoid- macrophage system in the quinine treatment of gallinaceum malaria. II. Quinine blood levels. I. Infect. Dis., 83:181-99. 1949 With L. G. Taliaferro. The role of the spleen and lymphoid- macrophage system in the quinine treatment of gallinaceum malaria. III. The action of quinine and of immunity on the parasite. }. Infect. Dis., 84:187-220. In memoriam for Dr. Charles Morley Wenyon. I Parasitol., 35:322-23. With L. G. Taliaferro. Asexual reproduction of Plasmodium knowlesi in rhesus monkeys. I. Infect. Dis., 85:107-25. The cellular basis of immunity. Annul Rev. Microbiol., 3: 159-94. Immunity to the malaria infections. In: Malariology, ed. M. F. Boyd, vol. 2, pp. 935~5. Philadelphia: W. B. Saunders Co. 1950 With L. G. Taliaferro. Reproduction-inhibiting and parasiticidal effects on Plasmodium gallinaceum and Plasmodium lophurae dur- ing initial infection and homologous superinfection in chickens. }. Infect. Dis., 86:275-94. With L. G. Taliaferro. The dynamics of hemolysin formation in intact and splenectomized rabbits. I. Infect. Dis., 87:37~2. With L. G. Taliaferro. Effect of X-irradiation on hemolysin decline. I. Infect. Dis., 87:201-9. 1951 . With L. G. Taliaferro. Effect of X-rays on immunity: A review. I. Immunol.66:181-212.

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402 BIOGRAPHICAL MEMOIRS With L. G. Taliaferro. The role of the spleen in hemolysin produc- tion in rabbits receiving multiple antigen injections. }. Infect. Dis., 89:143-68. 1952 With L. G. Taliaferro. The role of the spleen and the dynamics of hemolysin production in homologous anamnesis. I. Infect. Dis., 90:205-32. With L. G. Taliaferro and E. F. ianssen. The localization of X-ray injury to the initial phases of antibody response. J. Infect. Dis., 91: 105-24. 1954 With L. G. Taliaferro. Effect of X-rays on hemolysin formation following various immunization and irradiation procedures. }. Infect. Dis., 95:117-33. With L. G. Taliaferro. Further studies on the radio-sensitive stages in hemolysin formation. l. Infect. Dis., 95:134~1. With L. G. Taliaferro. Transfer of antibody-forming capacity in splenic materials. Science, 119: 585-86. 1955 Host-parasite relationships. In: Biological Specificity and Growth, ed. E. G. Butler, pp. 157-76. Princeton, N.T.: Princeton University Press. Studies in antibody formation. In: Some Physiological Aspects and Consequences of Parasitism, ed. W. H. Cole, pp. 50-75. New Brunswick, N.~: Rutgers University Press. With T. Pizzi. Connective tissue reactions in normal and immu- nized mice to a reticulotropic strain of Trypanosoma cruzi. ]. Infect. Dis., 96:199-226. With D. W. Talmage. Absence of amino acid incorporation into antibody during the induction period. J. Infect. Dis., 97:88-98. With L. G. Taliaferro. Reactions of the connective tissue in chickens to Plasmodium gallinaceum and Plasmodium lophurae. I. Histo- pathology during initial infections and superinfections. J. Infect. Dis., 97:99-136. With I. W. Moulder. Reactions of the connective tissue in chickens to Plasmodium gallinaceum and Plasmodium lophurae. II. Glucose

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WILLIAM HAY TALIAFERRO 403 metabolism during initial infections. I. Infect. Dis., 97:137-42. With B. N. Jaroslow.Hemolysin production in X-irradiated rabbits. Radiat. Res., 3:237. 1956 Parasitism and parasitology. In: Encyclopaedia Britannica, vol. 17, pp. 271-79. Chicago, London, Toronto: Encyclopaedia Britan- nica. With B. N. Taroslow. The restoration of hemolysin-forming capac- ity in X-irradiated rabbits by tissue and yeast preparations. I. Infect. Dis., 98:75-81. Functions of the spleen in immunity. Am. J. Trop. Med. Hyg., 5:391-410. With D. W. Talmage and G. G. Freter. The effect of repeated injections of sheep red cells on the hemolytic and combining capacities of rabbit antiserums. l. Infect. Dis., 98:293-99. With D. W. Talmage and G. G. Freter. Two antibodies of related specificity but different hemolytic efficiency separated by cen- trifugation. I. Infect. Dis., 98:300-305. With D. W. Talmage and G. G. Freter. The effect of whole body X-radiation on the natural sheep cell hemolysin of the rabbit. J. Infect. Dis., 99:241-45. With D. W. Talmage. Antibodies in the rabbit with different rates of metabolic decay. I. Infect. Dis., 99:21-33. With L. G. Taliaferro. X-ray effects on hemolysin formation in rabbits with the spleen shielded or irradiated. {. Infect. Dis., 99: 109-28. 1957 With L. G. Taliaferro. The effect of repeated doses of X-rays on the hemolysin response in rabbits. I. Infect. Dis., 101:85-99. With L. G. Taliaferro. Amino acid incorporation into precipitin at different stages in the secondary response to bovine serum albu- min. J. Infect. Dis., 101: 252-74. General introduction: synthesis and degradation of antibody. I. Cell Comp. Physiol., 50, Suppl. 1: 1-26. Modification of the immune response by radiation and cortisone. Ann. N.Y. Acad. Sci., 69:745~4.

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404 BIOGRAPHICAL MEMOIRS 1958 The synthesis and activities of antibodies. Rice Inst. Pam., 45: 114~0. With B. N. ~aroslow. Restoration of antibody-forming capacity in X-irradiated rabbits. Second International Conference on the Peace- ful Uses of Atomic Energy, vol. 23, pp. 79-83. Washington, D.C.: U.S. Government Printing Office. With B. N. ~aroslow. Effect of nucleic acid digests in restoration of hemolysin production in irradiated rabbits. Fed. Proc., 17:519. With L. G. Taliaferro and A. Pizzi. Avidity and intercellular trans- fer of hemolysin. Fed. Proc., 17:536. With T. Pizzi and P. D'Alesandro. Antibodies produced in the rat during infection with Trypanosoma lewisi. In: Proceedings of the Sixth International Congresses on Tropical Medicine and Malaria, vol. 3, pp. 259~3. Lisbon: Imprensa Portuese. 1959 With B. N. Jaroslow. Restoration of antibody-forming capacity in X-rayed rabbits. Science, 1 2 9: 1 2 8 9. With P. Stelos. Separation of antibodies by starch zone electro- phoresis. Anal. Chem., 31:845~8. With P. Stelos. Comparative study of rabbit hemolysins to various antigens. 2. Hemolysins to the Forssman antigen of guinea pig kidney, human type A red cells and sheep red cells. J. Infect. Dis., 104:105-18. With L. G. Taliaferro and A. Pizzi. Avidity and intercellular trans- fer of hemolysin. I. Infect. Dis., 105: 197-22 1. 1960 Spleen. In: Encyclopaedia Britannica, vol. 21, pp. 250-51. Chicago, London, and Toronto: Encyclopacdia Britannica. With L. T. Coggeshall. Malaria. In: Encyclopaedia Britannica, vol. 14, pp. 706-8. Chicago, London, and Toronto: Encyclopaedia Bri- tannica. With W. W. Wagener. Parasitism and parasitology. In: Encyclo- haedia Britannica. vol. 17. pp. 272-80. Chin. Lonclon anc1 Toronto: Encyclopacdia Britannica. With T. Pizzi. The inhibition of nucleic acid and protein synthesis in Trypanosoma lewisi by the antibody ablastin. Proc. Natl. Acad. Sci. USA, 46:733-45.

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WILLIAM HAY TALIAFERRO 405 With T. Pizzi. A comparative study of protein and nucleic acid synthesis in different species of trypanosomes. }. Infect. Dis., 107: 100-107. With B. N. iaroslow. The restoration of hemolysin formation in X-rayed rabbits by nucleic acid derivatives and antagonists of nucleic acid synthesis. J. Infect. Dis., 107:341-50. 1961 With L. G. Taliaferro. Intercellular transfer of gamma A-1 and gamma A-2 Forssman hemolysins. Proc. Natl. Acad. Sci. USA, 47:713-24. Whither bound how and why? Cancer Res., 21:1323-24. With J. H. Humphrey, eds. Advances in Immunology, vol. 1 York: Academic Press. 423 pp. 1962 ,. New With }. H. Humphrey, eds. Advances in Immunology, vol. 2. New York: Academic Press. 390 pp. Biochemical aspects of antibody formation. Sci. Rep. Inst. Super. Sanita, 2: 20-36. Remarks on the immunology of leishmaniasis. Sci. Rep. Inst. Super. Sanita, 2: 138~2. With L. G. Taliaferro. Immunologic unresponsiveness during the initial and anamnestic Forssman hemolysin response. I. Re- peated injections of heated sheep red cell stromata into rabbits before and after splenectomy. II. Spleen, bone marrow, lymph node and other tissue transfers. I. Infect. Dis., 110:165-200. With B. N. Jaroslow. Restoration of hemolysin-forming capacity in irradiated rabbits and its relation to induction of antibody syn- thesis. In: The Effects of Ionizing Radiations on Immune Processes, ed. C. A. Leone, pp. 301-14. New York: Gordon and Breach. 1963 The cellular and humoral factors in immunity to protozoa. In: Immunity to Protozoa, ed. P. C. C. Garnham, pp. 22-38. London: Blackwell Press. With L. G. Taliaferro. The effect of antigen dosage on the Forss- man hemolysin response in rabbits. I. Infect. Dis., 113:155~9.

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406 BIOGRAPHICAL MEMOIRS 1964 With L. G. Taliaferro. The relation of radiation dosage to enhance- ment, depression and recovery of the initial Forssman hemo- lysin response in rabbits. I. Infect. Dis., 114:285-303. With L. G. Taliaferro and B. N. Jaroslow. Radiation and Immune Mechanisms. New York: Academic Press. 152 pp. 1965 With L. G. Taliaferro. Enhancement of natural hemolysin in adult rabbits after radiation. Proc. Natl. Acad. Sci. USA, 53:139-46. 1966 With I. R. Marrack. Immunity and immunization. In: Encyclo- paedia Britannica, vol. 11, pp. 1108-14. Chicago, London, and Toronto: Encyclopaedia Britannica. With B. N. {aroslow. The effect of colchicine on the hemolysin response in unirradiated and irradiated rabbits. I. Infect. Dis., 1 16: 139-50. With L. G. Taliaferro. Persistence of hemolysin anamnestic reac- tivity in rabbits. Proc. Natl. Acad. Sci. USA, 56:1151-54. 1967 With L. G. Taliaferro. Effect of 5-bromodeoxyuridine on the hemolysin response in rabbits. Proc. Soc. Exp. Biol. Med., 124: 67 1-75. A retrospective look at the immunologic aspects of parasitic infec- tions. In: Immunologic Aspects of Parasitic Infections, pp. 3-20, 130-36. W. H. O. Scientific Publications No. 150. A retrospective look at the immunological aspects of parasitic infec- tions. Argonne Natl. Lab. Rev., 4:5146. 1968 With L. G. Taliaferro. The hemolytic immunoglobulins produced by unirradiated and irradiated rabbits immunized with Forss- man antigens. J. Infect. Dis., 1 18:278-88. With L. A. Stauber. Immunology of protozoan infections. In: Re- search in Protozoology, ed. T. T. Chen, vol. 3, pp. 50744. New York: Pergamon Press. Prefatory chapter: The lure of the unknown. Annul Rev. Micro- biol., 22: 1-14.

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WILLIAM HAY TALIAFERRO 407 1969 With L. G. Taliaferro. Effects of radiation on the initial and anam- nestic IgM hemolysin responses in rabbits; antigen injection after X-rays. }. Immunol., 103:55949. 1970 With L. G. Taliaferro. Effects of radiation on the initial and anam- nestic hemolysin responses in rabbits; antigen injection before X-rays. J. I mmunol., 104: 1364-76. With H. J. Shaughnessy. Immunity and immunization. In: Encyclo- paedia Britannica, vol. 12, pp. 2A-2G. Chicago, London, and Toronto: Encyclopaedia Britannica. With L. G. Taliaferro. The cellular reactions in the skin of normal and immune rabbits injected with Trichinella with special refer- ence to the hematogenous origin of macrophages. In: Srivastava Commemorative Volume, ed. K. S. Singh, pp. 517-26. Izatnager, U. P., India: Indian Vet. Res. Inst. With L. G. Taliaferro. Actinomycin D. before and during primary and secondary anti-Forssman immunoglobulin hemolysin re- sponses in rabbits. Proc. Natl. Acad. Sci. USA, 66:1036~3. 1971 With P. A. D'Alesandro. Trypanosoma lewisi infection in the rat: Effect of adenine. Proc. Natl. Acad. Sci. USA, 68:1-5. With M. D. Young. Malaria. In: Encyclopaedia Britannica, vol. 14, pp. 669-72. Chicago, London, Toronto: Encyclopacdia Britannica. With editors. Spleen. In: Encyclopaedia Britannica, vol. 21, pp. 44- 45. Chicago, London, and Toronto: Encyclopacdia Britannica. With W. W. Wagener. Parasitism and Parasitology. In: Encyclo- hacdia Britannica. vol. 17, pp. 323-~(). (ethical. 1 onclon. anc1 Toronto: Encyclopaedia Britannica. 1976 ~ _, With L. G. Taliaferro. Methods and applications of radiation in immunological research. In: Methods in Immunology and Immuno- chemistry, ed. C. A. Williams and M. W. Chase, vol. 5, pp. 239-59. New York: Academic Press.