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LEMUEL ROSCOE CLEVELAND November 14, Z892kehr7mry 12, l 969 BY WILLIAM TRAGER PROFESSOR CLEVELAND'S scientific work was his life, or so it seemed to his students and colleagues. Always reacly to talk science, he rarely revealed anything of his private life. This is unfortunate, since he had an unusual background. Born November 12, 1892 in Newton County, Mississippi, he grew up in this rural area with Indian children as his play- mates. He worked on the farm and once remarked that in three years with the same mule he had gotten to know it better than most of his human friends. After two years of high school in Union, Mississippi, he entered the University of Mississippi where he receiver! a B.S. in 1917 ant] then spent a year as a graduate student and Instructor in zoology. After a brief period of military service he taught at Emory University for two years and at Kansas State College for one year. He then entered Johns Hopkins University where he began the career of highly productive scientific research that terminated only shortly before his death in 1969. After re- ceiving his Ph.D. in 1923 he stayed on at the Hopkins School of Hygiene and Public Health as a National Research Council Fellow until 1925 when he went to the Department of Trop- ical Medicine at the Harvard Medical School. It was at Harvard that ~ hac! the goof! fortune to meet L. R. Cleveland and to become his first graduate student. In 49

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50 BIOGRAPHICAL MEMOIRS the fall of 1930 Cleveland's laboratory was an exciting place. He was just finishing his studies on cultivation of the human dysentery amoeba, Entamoeba h~stolytica. He and his associates had cleveloped a much improved medium in which the entire life cycle of this important parasite could be propagatecl. They had also macie initial attempts at bacteria-free cultiva- tion of amoebae. With characteristic insight, Cleveland rec- ognizecI the importance of what we now call axenic cultiva- tion, but his efforts toward this end with E. htstolytica were not successful. (Success in this was not achiever! until many years later, with L. S. Diamond's work in 1961.) Most exciting, however, was the new material Cleveland hac! brought back from the mountains of Virginia. He haci discovered that the large wood-dwelling roach Cryptocercus punctulatus contains a seething mass of protozoa in an en- larged portion of its hindgut. All were new species, many representing new genera ant] new families. But what was especially significant and particularly fascinating to Cleve- lanct was the fact that these protozoa were obviously closely related to the symbiotic intestinal flagellates of termites. The protozoa of termites hac! been known for many years, but it was Cleveland who first cliscoverec! their symbiotic nature. In a series of elegant experiments, done while he was a fellow of the National Research Council at the Johns Hopkins Univer- sity School of Hygiene, Cleveland showocl that the ability of termites to live on a diet of wood or cellulose depends on the digestive capacities of their intestinal flagellates. Termites deprived of these protozoa, but still infected with intestinal bacteria and spirochetes, wouIc! die of starvation if fed only wood or cellulose, but they couIcl be saved if reinfected with the protozoa. This was the first instance in which a mutualis- tic relationship between internal microorganisms and their metazoan host was clearly proved. It was pioneering work

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LEMUEL ROSCOE CLEVELAND 51 (published in 1923-1928) that paved the way for many later .. . . . . studies on sym Hotly microorganisms. One can easily imagine Cleveland's delight at finding in a different kind of insect, the wood-feeding roach, the same types of protozoa with which he had already been so success- ful in termites. It did not take him long to establish that the roach Cryptocercus, like termites, depends on its intestinal fla- gelIates for its ability to utilize cellulose as its principal food. He then embarked on a detailed study of all the new species of protozoa living in the hindgut of Cryptocercus. With the aid of a devoted research assistant, Miss lane Collier, and of two postdoctoral fellows, Dr. Elizabeth Sanders Hobbs and Dr. S. R. Hall, he soon Published a classic monograph (see bibliography, 1934~. Cleveland worked for the rest of his life mainly on taxo- nomic and experimental studies with the protozoa of Cr~ypto- cerc~s. Early in these studies he was the first to see and photo- graph in a living cell the fibers of the mitotic apparatus. In part because of their large size, certain of the flagellates of Cryptocercus provided exceptionally favorable material, but it was Cleveland's exacting microscopy and his application of the then newly available phase contrast methods that led to his beautiful results. He also gave much thought to the role of the centriole and its attendant organelles in cellular divi- sion. He produced two monographs (Trans. Am. Philos. Soc., 1949 and 1953) on behavior and structure of chromosomes. Far exceeding all these observations in general biological importance was Cleveland's discovery of the effect of molting of the host insect on sexual reproduction in its intestinal protozoa. Soon after he began working with Cryptocercus he noted cyst formation and various anomalous reproductive stages among the protozoa. There was a period of over five years during which Cleveland published hardly any papers

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52 BIOGRAPHICAL MEMOIRS while he was trying to determine what was really going on. Then came a long series of papers on the sexual cycles of the flagellates of Cryptocercus (summarized in a paper in [. Proto- zool., 3~l9561:161~0~. Sex hack been unknown in these fami- lies of protozoa. Cleveland now shower! that the sexual cycle was in all cases related to the molting cycle of the host insect. Some of the protozoa underwent only autogamy, others formed male and female gametes which fused and then un- derwent zygotic meiosis. Though the timing of these events also differed from species to species, it could always be corre- lated with molting in the roach. Furthermore, Cleveland showed that injection of an adult roach with a dose of the molting hormone ec~ysone too small to induce molting in the insect, nevertheless did induce the sexual cycles of the proto- zoa. The effect seems to be a direct one by the host hormone on the symbiotic protozoa. In the later stages of his scientific career Dr. Cleveland prepared several excellent cinemicro- graphs dealing with the sexual cycles of flagellates of Crypto- cercus and with the structure and movement of these protozoa as well as of protozoa in termites. Since Cryptocercus occurs, in the eastern United States, only in the Blue Ridge Mountains, Cleveland regularly spent his summers at the Biological Station at Mountain Lake, Vir- ginia, anti, late in his career, at the Biological Station at High- lancis, North Carolina. Here his rural youth surely server! him well as he swung his ax to break up logs in the search for colonies of Cryptocercus. Cleveland was elected to the National Academy of Sciences in 1952. He was President of the Society of Protozoologists in 1955 and was an honorary member of the Society. At Har- varcI, he mover! in 1936 from the Medical School to the Biol- ogy Department where he was advanced to Professor of Biol- ogy in 1946. He became Emeritus Professor in 1959 and, at the invitation of R. B. McGhee, went to the University of

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LEMUEL ROSCOE CLEVELAND 53 Georgia at Athens. Here he continued his active research. In ~ 965 he was among the protozoologists who were particularly honored at the Seconc! International Congress on Protozool- ogy in London. Unlike his triumphant scientific career, Cleveland's per- sonal life was market with sadness. His first wife, Mabel Bush, whom he marries! in 1925, cried of cancer in 1936, when their daughter, Margaret Elaine, was three years oict. Margaret died when only twenty-five. Clevelan~l's second marriage, to Dorothy Eleanor Colby, was more fortunate. She, and their son, Bruce Taylor Cleve- land, a physicist, survive. Cleve, as we called him, was clearly loved by his friends and family. He was sometimes clifficult, but only because completely honest men are likely to be difficult. He will be remembered mainly, however, for his great body of scientific achievement, and this is surely the way he would want it to be.

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54 BIOGRAPHICAL MEMOIRS BIBLIOGRAPHY 1923 Correlation between the food and morphology of termites and the presence of intestinal protozoa. Am. J. Hyg., 3:444 61. Symbiosis between termites and their intestinal protozoa. Proc. Natl. Acad. Sci. USA, 9:42~28. 1924 The physiological and symbiotic relationships between the in- testinal protozoa of termites and their hosts, with special refer- ence toReticulitennesflavipes Kollar. Biol. Bull., 46:177-225. 1925 Les effete de ['inanition et de ['oxygenation sur la symbiose entre les termites et leurs flagelles intestinaux. Ann. Parasitol. Hum. Comp., 3:35-36. Action toxique de l'oxygene sur les protozoaires in vivo et in vitro son utilization pour debarrasser les animaux de leurs parasites. Ann. Parasitol. Hum. Comp., 3:38~87. The method by which Trachonymph~a campanula, a protozoan in the intestine of termites, ingests solid particles of wood for food. Biol. Bull., 48:282-88. The ability of termites to live perhaps indefinitely on a diet of pure cellulose. Biol. Bull., 48:28~93. The feeding habit of termite castes and its relation to their intestinal flagellates. Biol. Bull., 48:29~306. The effect of oxygenation and starvation on the symbiosis between the termite, Termopsis, and its intestinal flagellates. Biol. Bull., 48:309~26. Toxicity of oxygen for protozoa in vivo and in vitro: animals defau- nated without injury. Biol. Bull., 48:455-68. The social genius of white ants. The Forum, pp. 32~0. 1926 Symbiosis among animals with special reference to termites and their intestinal flagellates. Q. Rev. Biol., 1:51~0.

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LEMUEL ROSCOE CLEVELAND 1927 55 The encystment of Paramoecium in the recta of frogs. Natural and experimental ingestion of Paramoecium by cockroaches. Science, 66:221-22. 1928 Further observations and experiments on the symbiosis between termites and their intestinal protozoa. Biol. Bull., 54:231-37. Tritrichomonasfecalis nov. sp. of man; its ability to grow and multiply indefinitely in faeces diluted with tap water and in frogs and tadpoles. Am. J. Hyg., 8:232-55. The separation of a Tritrichomonas of man from bacteria; its failure to grow in media free of living bacteria; measurement of its growth and division rate in pure cultures of various bacteria. Am. J. Hyg., 8:256-78. The suitability of various bacteria, molds, yeasts, and spirochaetes as food for the flagellate Tratrichomonasfecalis of man as brought out by the measurement of its fission rate, population density, and longevity in pure cultures of these microorganisms. Am. I. Hyg., 8:990~1013. 1930 With Elizabeth P. Sanders. Encystation, multiple fission without encystment, excystation, metacystic development, and variation in a pure line and nine strains of Entamoeba histolytica. Arch. Protistenkd., 70:223-66. With Elizabeth P. Sanders. The morphology and life-cycle of Ent- amoeba terrapinae spec. nov., from the terrapin, Chrysemys elegans. Arch. Protistenkd., 70:267-72. With Elizabeth P. Sanders. The production of bacteria-free amoe- bic abscesses in the liver of cats and observations on the amoebae in various media with and without bacteria. Science, 72: 149-51. With Elizabeth P. Sanders. The virulence of a pure line and several strains of Entamoeba histolytica for the liver of cats and the rela- tion of bacteria, cultivation, and liver passage to virulence. Am. J. Hyg., 12:569~05. With lane Collier. Various improvements in the cultivation of Ent- amoeba histolytica. Am. I. Hyg., 12:606-13.

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56 BIOGRAPHICAL MEMOIRS With lane Collier. The cultivation and differentiation of haemo- flagellates in autoclaved media. Am. }. Hyg., 12:61~23. The symbiosis between the wood-feeding roach, Cryptocercus punc- tulat?`s Scudder, and its intestinal flagellates. Anat. Rec., 47:293-94. 1934 With S. R. Hall, Elizabeth P. Sanders, and Jane Collier. The wood- feeding roach C,yptocercus, its protozoa, and the symbiosis be- tween protozoa and roach. Mem. Am. Acad. Arts Sci., 17~2~: 18~342. 1935 The centriole and its role in mitosis as seen in living cells. Science, 81 :59~600. The centrioles of Pseudotr~chonympha and their role in mitosis. Biol. Bull., 69:4~51. The intranuclear achromatic figure of Oxymonas grandis sp. nov. Biol. Bull., 69:54~63. 1938 Longitudinal and transverse division in two closely related flagel- lates. Biol. Bull., 74:1-24. Origin and development of the achromatic figure. Biol. Bull., 74:41-55. Morphology and mitosis of Teranymph~a. Arch. Protistenkd. 91:442-51. Mitosis in Pyrsonympha. Arch. Protistenkd., 91 :452-55. 1947 Sex produced in the protozoa of Crytocercus by molting. Science, 105: 16-18. The origin and evolution of meiosis. Science, 105:287. 1948 An ideal partnership. Sci. Mon., 67:173-77. 1949 The whole life cycle of chromosomes and their coiling systems. Trans. Am. Philos. Soc., 39:1-100.

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LEMUEL ROSCOE CLEVELAND 57 Hormone-induced sexual cycles of flagellates. I. Gametogenesis, fertilization, and meiosis in Trachonymph~a. ]. Morphol., 85: 197-296. 1950 Hormone-induced sexual cycles of flagellates. II. Gametogenesis, fertilization, and one-division meiosis in Oxymonas. ]. Morphol., 86: 185-214. Hormone-induced sexual cycles of flagellates. III. Gametogenesis, fertilization, and one-division meiosis in Saccinobaculus. ]. Mor- phol., 86:215-28. Hormone-induced sexual cycles of flagellates. IV. Meiosis after syngamy and before nuclear fusion in Notila. ]. Morphof., 87:317~8. Hormone-induced sexual cycles of flagellates. V. Fertilization in Eucomonympha. ]. Morphol., 87:349 68. 1951 Hormone-induced sexual cycles of flagellates. VI. Gametogenesis, fertilization, meiosis, oocysts, and gametocytes in Leptospironym- pha. J. Morphol., 88: 199-244. Hormone-induced sexual cycles of flagellates. VII. One-division meiosis and autogamy without cell division in Ur~nvmtha. I. Mor- phol., 88:385~40. 1952 ~ 1 a Hormone-induced sexual cycles of flagellates. VIII. Meiosis in Rhynchonympha in one cytoplasmic and two nuclear divisions followed by autogamy. J. Morphol., 91:269-324. 1953 Hormone-induced sexual cycles of flagellates. IX. Haploid gameto- genesis and fertilization in Barbulanympha. ]. Morphol., 93: 371-404. With A. M. Winchester. Photographs of living chromosomes. l. Hered.,44:118-27. Studies on chromosomes and nuclear division. I. Fusion of nucleoli independent of chromosomal homology. II. Spontaneous aber- rations, homologous and non-homologous union of fragments. III. Pairing, segregation, and crossing-over. IV. Photomicro-

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58 BIOGRAPHICAL MEMOIRS graphs of living cells during meiotic divisions. Trans. Am. Philos. Soc., 43 :809~9. 1954 Hormone-induced sexual cycles of flagellates. X. Autogamy and endomitosis in Barbulanympha resulting from interruption of haploid gametogenesis. J. Morphol., 95: 189-212. Hormone-induced sexual cycles of flagellates. XI. Reorganization in the zygote of Barbulanympha without nuclear or cytoplasmic division. J. Morphol., 95:213-36. Hormone-induced sexual cycles of flagellates. XII. Meiosis in Bar- bulanympha following fertilization, autogamy, and endomitosis. J. Morphol., 95:557-620. 1955 Hormone-induced sexual cycles of flagellates. XIII. Unusual behavior of gametes and centrioles of Barbulanympha. J. Mor- phol., 97:511~2. With W. L. Nutting. Suppression of sexual cycles and death of the protozoa of Cryptocercus resulting from change of hosts during molting period. J. Exp. Zool., 130:485-514. 1956 Hormone-induced sexual cycles of flagellates. XIV. Gametic meio- sis and fertilization in Macrospironympha. Arch. Protistenkd., 101 :99-169. With Arthur W. Burke, Jr. Effects of temperature and tension on oxygen toxicity for the protozoa of Cryptocercus. J. Protozool., 3:74-77. Cell division without chromatin in Trtchonympha and Barbulanym- pha. J. Protozool., 3 :78-83. Brief accounts of the sexual cycles of the flagellates of Cryptocercus. J. Protozool., 3:161-80. 1957 Types and life cycles of centrioles of flagellates. J. Protozool., 4:23~41. Achromatic figure formation of multiple centrioles of Barbulanym- pha. J. Protozool., 4:241-48.

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LEMUEL ROSCOE CLEVELAND 1958 59 A fractural analysis of chromosomal movement in Barbulanympha. I. Protozool., 5:47-62. Movement of chromosomes in Spirotrichonympha to centrioles in- stead of the ends of central spindles. I. Protozool., 5:63~8. Photographs of fertilization in the smaller species of Trtchonympha. I. Protozool., 5: 105-15. Photographs of fertilization in Trichonympha grandis. j. Protozool., 5:115-22. With Max Day. Spirotrichonymphidae of Stolotermes. Arch. Pro- tistenkd., 103: 1-53. 1959 Sex induced with ecdysone. Proc. Natl. Acad. Sci. USA, 45:747-53. 1960 Photographs of living centrioles in resting cells of Trichonympha collards. Arch. Protistenkd., 105: 11 ~ 12. Photographs of fertilization in Eucomonympha. Arch. Protistenkd., 105: 137~8. The centrioles of Trichomonas and their functions in cell reproduc- tion. Arch. Protistenkd., 105: 14~62. Pairing and segregation in haploids and diploids of Holomastigotoi- des. Arch. Protistenkd., 195: 16~72. With A. W. Burke, Jr., and P. Karlson. Ecdysone induced modifica- tions in the sexual cycles of the protozoa of Cryptocercus. ]. Pro- tozool., 7:229-39. With A. W. Burke, fir. Modifications induced in the sexual cycles of the protozoa of Cryptocercus by change of host. l. Protozool. 7:240~5. The centrioles of Trichonympha from termites and their functions in reproduction. I. Protozool., 7:326 41. Induction and acceleration of gametogenesis in flagellates by the insect hormone ecdysone. Science, 131: 1317. Effects of insect hormones on the protozoa of Cryptocercus and termites. In: Host Influence on Parasite Physiology, ed. Leslie A. Stauber, pp. ~10. New Brunswick, N.~.: Rutgers Univ. Press.

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60 B I OGRAPH I CAL MEMOI RS 1961 Induction and acceleration of gametogenesis in flagellates by the insect hormone ecdysone (Cryptocercus punctulatusJ. Frog. Proto- zool., Proc. Int. Cong. Protozool., 1:289-91. 1963 Functions of flagellate and other centrioles in cell reproduction. In: The Cell in Mitosis, ed. Laurence Levine, Proceedings of the First Annual Symposium of the Wayne State Fund Research Rec- ognition Award, pp. 3-53. N.Y.: Academic Press. 1964 With A. V. Grimstone. Fine structure of flagellate Mixotricha para- doxa and its associated micro-organisms. Proc. R. Soc. Lond. Ser. B. 159:668-85. 1965 With A. V. Grimstone. Fine structure and function of contractile axostyles of certain flagellates. I. Cell Biol., 24:387~00. 1966 Nuclear division without cytokinesis followed by fusion of pronu clef in Paranotila lata gen. et sp. nov. l. Protozool., 13:132-36. Reproduction by binary and multiple fission in Gigantomonas. J. Protozool., 13:573-85.

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