Below are the first 10 and last 10 pages of uncorrected machine-read text (when available) of this chapter, followed by the top 30 algorithmically extracted key phrases from the chapter as a whole.
Intended to provide our own search engines and external engines with highly rich, chapter-representative searchable text on the opening pages of each chapter.
Because it is UNCORRECTED material, please consider the following text as a useful but insufficient proxy for the authoritative book pages.
Do not use for reproduction, copying, pasting, or reading; exclusively for search engines.
OCR for page 48
OCR for page 49
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
OCR for page 50
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
OCR for page 51
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
OCR for page 52
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
OCR for page 53
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.
OCR for page 54
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.
OCR for page 55
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.
OCR for page 56
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.
OCR for page 57
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-
OCR for page 58
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.
OCR for page 59
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.
OCR for page 60
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.
OCR for page 61
Representative terms from entire chapter:
intestinal flagellates
