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PAUL RUFUS BURKHOLDER
February 1, 1903-August 11, 1972
BY JAMES G. HORSFALL
DISTINGUISHED SCIENTISTS don t just happen. Paul Rufus
Burkholder wasn't just accidentally elected (1949) to the
National Academy of Sciences. He didn't have his greatness
thrust upon him; he earned it.
In him was a concatenation of factors that characterize many
members of the Academy: (a) He was born into and raised in
an intellectual family. His father had a "library" that young
Burkholder devoured. (b) He was a bright boy. (c) He was a
prodigious worker as witness the extensive bibliography at-
tached. (d) He was an introvert at heart, a loner if you please.
This allowed him the time to work. (e) He was stimulated by
his early teachers to pursue science. (f) He had a devoted wife
who shared his hobbies, his life, and his lab. She worked as
hard in biology as he did and raised three sons besides.
I knew Burkholder in the graduate school at Cornell. I
knew him again when I was an adjunct professor in his depart-
ment at Yale. I always admired "Burkie," even though he was
difficult to "know" well.
When he was in graduate school, he compensated for some
of his introvert tendencies by joining Gamma Alpha, a gradu-
ate student fraternity. There he found another "Burkie" (no
relative), whom I also knew. This may not have helped Paul
too much, for the second Burkie was a very tall man and Paul
3
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4
BIOGRAPHICAL MEMOIRS
was a little short. Promptly the two were distinguished as
"big Burkie" and "little Burkie." It must be discouraging to
be called a "little Burkie." On the other hand, it drives the
little Burkies to excell the big Burkies.
At any rate, Paul fought the little-man complex so hard
that he was able to put on giant shoes and take giant strides
· .
in science.
HIS INTELLECTUAL BASE
The Burkholder forebears, religious refugees from the Ger-
man section of Switzerland, came to Pennsylvania during its
early colonial period. William Rankin Burkholder, Paul's
father, was born in 1857, just before the Civil War, on a farm
in central Pennsylvania. As a young man, Paul's father and
an older brother operated a successful general store in Middle-
burg. William was small, quick, and very strong, taking delight
in wrestling, hunting, and driving fast horses. To play cards
and drink hard liquor was not to be scorned either. Money
was plentiful and life was full, strenuous, and satisfying.
In some way, William's older brother became convinced
that he should forsake this way of life and become a minister
in the church. He brought influence to bear on William to do
likewise. This was a difficult problem. Could he give up all
those things that he enjoyed? The diary he kept at the time
bears witness to his struggle. His brother—positive, eloquent,
and persuasive showed him his duty. Once the decision was
made, he went off to college,~as very few did a century ago.
In time he was ordained as a minister in the United Brethren
Church by Bishop Wright, the father of the famous Wright
brothers. As his son was to do later, he pursued a solitary,
dedicated life of service to the small communities in south-
central Pennsylvania. There, when he was thirty-nine years old,
he met and married Mary Ellen Schubert, a young girl of
, . . .
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PAUL RUFUS BURKHOLDER
5
twenty-one. Their son, Paul, an only child, was born seven
years later, on February 1, 1903, at Orrstown, Pennsylvania.
Mary's father, George Schubert, came from Germany as a
young boy. Because the children made fun of his accent, he
left school to become a farmer, married a girl of French
descent, and raised a family of seven children. Mary, the oldest,
like her son later, had an alert mind and a desire to become a
teacher. Her mother pressured her to stay safely at home,
however, to help with the family and the aged grandparents.
In spite of her bitter disappointment, her sense of responsi-
bility won out and she did not rebel. Thus, her formal school-
ing came to an end and she turned to church work for an
outlet. Here, William found her. Typical of ministers' families,
they had very little money, but a house was always provided,
along with space for a garden and a farm to keep a cow and a
few chickens. Here a small boy could grow in an atmosphere
of freedom and with no real sense of hardship. The environ-
ment, if somewhat spartan, was intellectual and enriched by
visitors to the home and church. Paul read and reread his
father's library, consisting mostly of books on religious subjects.
Sundays he spent in church, where three times a day he listened
to his father's sermons or found the time well spent in medi-
tation.
He understood what his father was trying to do and admired
him for his tireless, if often frustrating, efforts to help the
people in their individual, everyday problems and to keep the
churches growing and active. Paul knew that the basic princi-
ples were right, but there must be more than this. Questions
were left unanswered, and Paul went out to find the answers.
School and learning were important. Rightly or wrongly,
he felt that he was not so bright and must work extra hard to
compensate. He took full advantage of the knowledge exhib-
ited by the schoolmasters of the little village schools. Summers,
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6
BIOGRAPHICAL MEMOIRS
while still very young, he started working for farmers, weeding
and picking vegetables and fruit and helping to harvest hay
and grain. He came to be known as swift and dependable, so
there was no problem in procuring work. Always he saved the
money he earned. His spare time was spent roaming in the
mountains with his gun, shooting small game for food, and
· .
enjoying nature.
Ready for high school, he walked three miles to a train that
took him to Chambersburg and three miles home every day.
This schedule kept him busy from early until late, but he
enjoyed the work and did well in his classes.
COLLEGE DAYS
In 1920 he enrolled in Dickinson College in Carlisle, Penn-
sylvania. His classes ranged over a great variety of subjects as
available in a small liberal arts college.
1 ~ ~ ~ T
He worked in the
Bleary tor expense money. He appreciated this easy access to
the books and he used them to good advantage. Like many
boys, he worked summers as a laborer.
During his four years at Dickinson, he decided to become
a botanist. This decision was influenced no doubt by his early
and continuing interest in plants. In 1949 Dickinson College
recognized his work with an honorary D.Sc.
GRADUATE SCHOOL
At Cornell, he came under the enthusiastic tutelage of O. F.
Curtis and Louis Knudson and almost automatically became
a plant physiologist. Of his contemporaries who took plant
physiology with him at Cornell, at least four, including himself,
were later elected to the National Academy of Sciences.
Cornell was surely the happy hunting ground for biologists.
There was the science in the laboratory, but the campus and
surrounding hills were laced with gorges through which streams
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PAUL RUFUS BURKHOLDER
cascaded down to the lake from "far above Cayuga's waters"—
to quote the old Cornell song.
What vegetation to study! It was different on the sunny
north side from the shady south side of the gorge. Paul's
Gamma Alpha house was on the south edge of Cascadilla gorge.
He could collect specimens for his classes almost at his doorstep,
or he could go across campus to the Fall Creek gorge to collect
plants on the north edge or swim in Beebe Lake and be eyed
by the Tri Delta girls from their house on the rim above. And
besides he helped the brothers make wine during prohibition
days. He was surely a chip off his father's block.
Like all graduate students in the twenties, before National
Science Foundation student fellowships, he needed money. A
job offer introduced him to the excitement of aquatic biology.
The New York State Conservation Department offered him
a summer job in the limnological survey of the state water-
sheds, beginning with the Cayuga Lake basin, with which he
was already familiar. He did the phytoplankton studies. He and
his colleagues worked hard, had fun, earned money, and pub-
lished their papers. This work was to influence him throughout
the rest of his life.
THE DEPRESSION
He received his Ph.D. in 1929, the year of Black Friday and
the collapse of the stock market, but it did not discourage him.
He was able to continue with the limnological survey. It was
extended to Lake Erie in cooperation with the Buffalo Museum
of Natural Sciences, where he became a curator of biology
and continued plankton studies on the lake. The government
boat, Sheerwater, manned by a discrete crew and carefree
young scientists, came to be known to the "rum runners" on
Lake Erie as a friend in time of need. There were adventures
aplenty on the waters and opportunities for firsthand observa-
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8
BIOGRAPHICAL MEMOIRS
tion of an epic in our history. Buffalo society added to the fun
and frolic.
While at the Museum, he planned and installed exhibits
and participated in the teaching program. Continuing his plank-
ton studies, he spent one summer working in northern New
York waters and another working in some lakes of the upper
Hudson watershed. The last summer he did a study of the
phytoplankton of Frenchman's and Penobscot bays, in Maine,
near Bar Harbor.
In 1930 he married Lillian Miller, a lady who shared his
biological enthusiasms for the rest of his life and who survives
him. With the acquisition of a wife and a little later a son,
Franz, life took on a more serious aspect. He wished to return
to a more academic environment. In the meantime the depres-
sion had deepened and no jobs were available. A National Re-
search Council Fellowship saved him, however, and he spent
two rewarding years, one at Harvard, where a second son, Peter,
was born in 1935, and one at Columbia, where he came under
the influence of E. W. Sinnott, a member of the Academy.
While at Columbia, he took a course in bacteriology. This
became another influence on the rest of his career.
TO CONNECTICUT COLLEGE
Even after two fellowship years, jobs were still scarce, but
one appeared in the nick of time. A young Ph.D., George
Avery, had recently begun teaching at the Connecticut College
for Women, at New London. (This college has now been
"liberated" to Connecticut College and men go there.) Being
alone among all those women, Avery shouted for help to Sinnott
at Columbia. Sinnott sent him Burkholder.
Burkholder promptly made his late-blooming course in bac-
teriology pay off. He taught the girls about germs, but now his
interest in plant physiology returned. Avery started his work
on plant hormones at Connecticut College and since it was a
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PAUL RUFUS BURKHOLDER
.9
timely subject, they collaborated on work in this field and
produced a book on it. The next three years yielded an im-
pressive return in papers published.
BACK TO AQUATIC BIOLOGY
While at New London, Paul purchased a small farm near
Falmouth, Massachusetts, on Cape Cod. Falmouth harbor was
near, and so Burkholder bought a sail boat and returned to
the water. Wood's Hole was near, too, and here was the oppor-
tunity to return to the study of the creatures that live in the
water.
This lasted only three years, however. In 1938 Dr. Rickett
left the post of plant physiologist at the University of Missouri
and the boy from the hills of Pennsylvania went to the hills of
Missouri. This seemed a logical move. A large well-staffed
agricultural school provided many interested students for the
botany courses. Despite much teaching to be done, he managed
to carry on research work, mostly in plant nutrition. Here the
third son, Karl, was born in 1940. When he left, Dr. Tucker,
chairman of botany, jokingly said, "I will have to hire two
men to take your place, a day man and a night man."
A YALE MAN
In 1940 Sinnott entered his life again. Dr. Sinnott went to
Yale as chairman of the botany department and asked Burk-
holder to go with him as plant physiologist.
Botany at Yale was at an all-time low ebb, partly because of
the depression and partly because of a failure to bring in new
faculty members, but mostly because the university had no
agriculture program to send the students of botany on to jobs.
Zoology was strong at Yale because there was a medical school
to beckon the students from that department.
The next few years were spent in building up the faculty
of botany. Norman Giles, a new Ph.D. in plant genetics, came
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10
BIOGRAPHICAL MEMOIRS
from Harvard; Reader came as a taxonomist in charge of the
herbarium. Ed Tatum brought his Neurospora work and Dave
Bonner followed. Burkholder taught a new course in micro-
biology that became very popular. Galston came to teach plant
physiology. Paul Sears set up a program in conservation.
The department was on the way. Graduate students ap-
peared. Joshua Lederberg, who discovered sex in bacteria; Ed
Adelberg, now head of microbiology at Yale; Guillard at
Wood's Hole; Sy Pomper; Lou Nickell; Ina May Martin from
Jamaica; Joyce and Ralph Lewin; and many others received
degrees from the department.
The years were good at Yale. Burkholder was made Eaton
professor of botany. His research moved apace, especially in
microbiology, production of vitamins by yeasts, and production
of antibiotics by lower fungi.
CHLORAMPHENICOL DISCOVERED
Burkholder soon made a dramatic discovery. Fleming's
penicillin had arrived during the war. Penicillin came from a
fungus. What other antibiotics from microorganisms could be
found? Waksman had discovered streptomycin and received
the Nobel Prize. Burkholder soon discovered chloramphenicol
in an actinomycete, which he isolated from a Venezuelan soil.
The Parke-Davis Company developed, purified, and synthe-
. .
sized it.
Chloramphenicol was discovered just in time to save the
lives of soldiers in the Korean War from scrub typhus, a classical
killer of soldiers in wartime.
Chloramphenicol is a dramatic case in biology for another
reason. Chemists like to chlorinate and nitrate compounds. In
1947, when chloramphenicol was discovered, dogma said that
no organism could do these chemical tricks. They couldn't
attach a chlorine or a NOR group to carbon as chemists could.
Chlorine and nitro were the substituents chemists used to make
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PAUI, RUFUS BURKHOLDER
11
killers for organisms—note dinitrophenol wood preservative or
chlorinated hydrocarbon insecticides.
Burkholder's actinomycete proved the dogma wrong. Not
only could it chlorinate a compound, it could nitrate it, too;
and besides it could put both substituents in the same molecule.
This only proves that chemists were a million years late in
learning these tricks.
A PALACE REVOLUTION
In the meantime, Yale botany was not all serene. Burk-
holder became the unfortunate victim of a palace revolution.
Sinnott had held a brilliant group of thinkers together, but
then he moved up to be dean of the Sheffield Scientific School.
Having always supported Burkholder, Sinnott made him chair-
man of the botany department, but Burkholder had no liking
for scientific politics and had not developed administrative
skills.
Severe unhappiness showed in the microbiological con-
tingent. And Burkholder didn't really hold a union card in
microbiology. He came into it late as a plant physiologist. This
problem was tied to the national picture as well. The bac-
teriologists of the country had arrogated to themselves the term
microbiology. People working with other microorganisms like
protozoa, fungi, and one-celled algae were low in the pecking
order; and Burkholder's specialty was fungi.
Bacteria are mostly dealt with in two university disciplines—
medicine and agriculture. Yale had no agriculture. Medicine,
therefore, became a magnet for the new microbiology created
by Burkholder at Yale in the botany department. And so it
leas natural for friction to develop within classical botany.
Whereupon, botany lost another child that it had begotten.
The explosion was severe. Tatum left, Galston left, Naylor
left, Burkholder left, and Bonner deserted the botany depart-
ment for the medical school and a new department of micro-
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12
BIOGRAPHICAL MEMOIRS
biology. It seems ironical that Bonner was later elected to the
National Academy of Sciences through the botany section, not
through microbiology.
GEORGIA BECKONS
Georgia beckoned to Burkholder and Burkholder listened.
He labored strenuously there to build up science in general and
microbiology in particular, but it went too slowly for him.
The Sloan-Kettering Cancer Institute in New York thought
so, too. Cornelius Rhoades, its director, suggested to George
Avery, by then director of the Brooklyn Botanic Garden, that
Burkholder could be moved. He could be and was.
The next five years at the Garden, with his old friend George
Avery and with Sloan-Kettering cooperation, saw an intensive
program of screening soil organisms for antibiotics that might
have the potential for chemotherapy of cancer.
There is a fascinating sequel to the Sloan-Ketter~ng phase.
They kept Burkholder's cultures "on deposit" when he left.
Rhoades chanced to mention this to Jasper Kane, then a re-
search executive with the Pfizer Company. Kane procured the
cultures for futher screening. In one of these was the fungus
that produces Terramycin for Pfizer.
BACK TO COLUMBIA
Since the cancer idea did not pay off, Burkholder returned
in 1961 to Columbia and to his old love for the sea and marine
biology. Perhaps the inhabitants of the sea could provide the
hoped-for antibiotics that fungi from the worked-over soil could
no longer supply. He affiliated with the new biology laboratory
in the Lamont Geological Observatory at Columbia, situated
up the Hudson River from New York. He was no longer land-
locked. He had come full circle back to the organisms in the
water.
He spent a season in the Antarctic working with Sieburth
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PAUL RUFUS BURKHOLDER
BIBLIOGRAPHY
KEY TO ABBREVIATIONS
15
Am. I. Bot. = American Journal of Botany
Ann. N.Y. Acad. Sci. -_ Annals of the New York Academy of Sciences
Appl. Microbiol. = Applied Microbiology
Arch. Biochem.—Archives of Biochemistry
Bot. Gaz. _ Botanical Gazette
Bot. Mar. = Botanica Marina
Bull. Buffalo Soc. Nat. Sci. _ Bulletin of the Buffalo Society of Natural
Sciences
Bull. Mar. Sci. Gulf Caribb. _ Bulletin of Marine Science of the Gulf and
Caribbean
Bull. Torrey Bot. Club _ Bulletin of the Torrey Botanical Club
I. Antibiot. = {ournal of Antibiotics
i. Bacterial. = journal of Bacteriology
.T Org. Chem. = Journal of Organic Chemistry
Limnol. Oceanogr. _ Limnology and Oceanography
Mar. Biol. = Marine Biology
Plant Physiol. = Plant Physiology
Proc. Natl. Acad. Sci. = Proceedings of the National Academy of Sciences
Spec. Sci. Rep. Fish. = Special Scientific Report, Fisheries
1930
Microplankton studies of Lake Erie. Bull. Buffalo Soc. Nat. Sci.,
14:73-93.
The heredity and environment exhibit. Hobbies. The Magazine
of the Buffalo Museum of Science.
1931
Studies in the phytoplankton of the Cayuga Lake basin, New York.
Bull. Buffalo Soc. Nat. Sci., 15: 181.
1933
A study of the phytoplankton of Frenchman's Bay and Penobscot
Bay, Maine. Internationale Revue gesamten Hydrobiologie
und Hydrographie, 28:262-84.
Movement in cyanophyceae. The effect of pH upon movement in
Oscillatoria. Journal of General Physiology, 16:875-81.
1934
Movement in the cyanophyceae. The Quarterly Review of Biology,
9:438-59.
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16
BIOGRAPHICAL MEMOIRS
1936
With R. Pratt. Leaf-movement of Mimosa pud ica in relation to
light. Am. I. Bot., 23 :46-52.
The role of light in the life of plants. The Botanical Review,
2:1-52, 97-168.
With G. S. Avery. Polarized growth and cell studies on the Arena
coleoptile, phyto-hormone test object. Bull. Torrey Bot. Club,
63: 1-15.
With Boysen-lensen and G. S. Avery. Growth Hormones in Plants.
New York: McGraw-Hill Book Co. 268 pp.
With G. S. Avery and H. B. Creighton. Plant hormones and mineral
nutrition. Proc. Natl. Acad. Sci., 22:673-78.
1937
With E. S. Johnston. Inactivation of plant growth substance by
light. Smithsonian Miscellaneous Collections, 95~20~: 1-14.
Production and distribution of growth hormone in shoots of Aescu-
lus and Malus and its probable role in stimulating cambial
activity. Am. l. Bot., 24:51-58. \
With G. S. Avery and H. B. Creighton. Avena coleoptile curvature
in relation to different concentrations of certain synthetic
substances. Am. l. Bot., 24:226-32.
With G. S. Avery and H. B. Creighton. Nutrient deficiencies and
growth hormone concentration in Helianthus and Nicotiana.
Am. J. Bot., 24: 553-57.
With G. S. Avery and H. B. Creighton. Growth hormone in termi-
nal shoots of Nicotiana in relation to light. Am. J. Bot.,
24:666-73.
With G. S. Avery and H. B. Creighton. Polarized growth and cell
studies in the first internode and coleoptile of Avena in relation
to light and darkness. Bot. Gaz., 99: 125-43.
1938
With G. S. Avery, H. B. Creighton, and B. A. Scheer. Darwin and
early discoveries in connection with plant hormones. Science,
87:56.
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PAUL RUFUS BURKHOLDER
1939
17
Production of growth substance by bacteria in media containing
specific organic and inorganic nitrogenous compounds. Am. l.
Bot., 26:422-28.
1940
With I. McVeigh. Growth and differentiation of maize in relation
to nitrogen supply. Am. J. Bot., 27:414-24.
With I. McVeigh. Growth of Phycomyces Blakesleeanus in relation
to varied environmental conditions. Am. [. Bot., 27:634~0.
With I. McVeigh. Studies on thiamine in green plants with the
Phycomyces assay method. Am. J. Bot., 27:853-61.
1941
With I. McVeigh. Multinucleate plant cells. Bull. Torrey Bot.
Club, 68: 395-96.
With C. M. Tucker. Calcium deficiency as a factor of abnormal
rooting of Philodendron cuttings. Phytopathology, 31:844~8.
Some experiments with growth curvatures and growth substances.
Am. l. Bot., 28:911-21.
1942
Pyridoxine as a growth factor for Graphium. Science, 95:127-28.
With A. G. Snow, Jr. Thiamine in some common American trees.
Bull. Torrey Bot. Club, 69:421-28.
Synthesis of vitamins by intestinal bacteria. Proc. Natl. Acad. Sci..
28:285-89.
With I. McVeigh. The increase of B vitamins in germinating seeds.
Proc. Natl. Acad. Sci., 28:440-46.
1943
Vitamin deficiencies in yeasts. Am. I. Bot., 30:206-11.
Vitamins in dehydrated seeds and sprouts. Science, 97:562-64.
Synthesis of riboflavin in a yeast. Proc. Natl. Acad. Sci., 29:166-72.
With D. Moyer. Vitamin deficiencies of fifty yeasts and molds.
Bull. Torrey Bot. Club, 70:372-77.
Vitamins in edible soybeans. Science, 98: 188-90.
Influence of some environmental factors upon the production of
riboflavin by a yeast. Arch. Biochem., 3: 121-29.
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18
BIOGRAPHICAL MEMOIRS
With l. Collier and D. Moyer. Synthesis of vitamins by micro-
organisms in relation to vitamin content of fancy cheeses. Food
Research, 8:314-22.
1944
With I. McVeigh and D. Moyer. Niacin in maize. Yale [ournal of
Biology and Medicine, 16~6~: 659-63.
With A. W. Evans, I. McVeigh, and H. K. Thornton. Antibiotic
activity of lichens. Proc. Natl. Acad. Sci., 30:250-55.
Some growth patterns of bacteria in cylinder plate tests for pro-
moting and inhibiting substances. Am. I. Bot., 31:555-58.
With I. McVeigh and D. Moyer. Studies on some growth factors of
yeasts. I. Bacteriol-, 48: 385-91.
1945
With A. W. Evans. Further studies on the antibiotic activity of
lichens. Bull. Torry Bot. Club, 72: 157-64.
With E. W. Sinnott. Morphogenesis of fungus colonies. in sub-
merged shaken cultures. Am. I. Bot., 32:424-31.
With I. McVeigh and K. Wilson. Studies on vitamin "B" produced
by microorganisms. Arch. Biochem., 7:287-303.
With I. McVeigh. The B vitamin content of buds and shoots of
some common trees. Plant Physiol., 20:276-82.
With I. McVeigh. Vitamin content of some mature and germinated
legume seeds. Plant Physiol., 20:301-6.
Studies on the antibiotic activity of actinomycetes. l. Bacteriol.,
52: 503~.
1947
With Norman H. Giles, in Induced biochemical mutation in
Bacillus sub tills. Am. J. Bot., 34: 345~8.
With L. G. Nickell. Inhibition of Azotobacter by soil actinomy-
cetes. Journal of the American Society of Agronomy, 39:771-79.
Chloromycetin, a new antibiotic from a soil actinomycete. Science,
106:2757.
1948
With [. Ehrlich, D. Gottlieb, L. E. Anderson, and T. G. Pridham.
Streptomyces Venezuelae, n. sp., the source of chloromycetin.
I. Bacteriol., 56:467-77.
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PAUL RUFUS BURKHOLDER
19
With L. G. Nickell. Growth responses of Rumex virus wound
tumor in tissue culture. Am. l. Bot., 35:811.
With Mary E. Sanders. Influence of amino acids on growth of
Datura embryos in culture. Proc. Natl. Acad. Sci., 34:516-26.
1949
With L. G. Nickell. Atypical growth of plants. I. Cultivation of
virus tumors of Rumex on nutrient agar. Bot. Gaz., 110:426-37.
With Seymour Pomper. Studies on the biochemical genetics of
yeast. Proc. Natl. Acad. Sci., 35:456-64.
With R. G. H. Sin, R. T. Darby, and E. S. Barghoorn. Specificity
of microbiological attack on cellulose derivatives. Textile Re-
search, 29~8~:484-88.
1950
With L. G. Nickell. Ayptical growth of plants. II. Growth in vitro
of virus tumors of Rumex in relation to temperature, pH and
various sources of nitrogen, carbon, and sulfur. Am. l. Bot.,
37:538~7.
With L. G. Nickell and P. Greenfield. Atypical growth of plants.
III. Growth responses of virus tumors of Rumex to certain
nucleic acid components and related compounds. Bot. Gaz.,
112:42-52.
1951
Determination of vitamin B-12 with a mutant strain of E. coli.
Science, 114:459-60.
1952
Microbial studies on materials which potentiate oral vitamin B-12
therapy in Addisonian anemia. Archives of Biochemistry and
Biophysics, 39:322-32.
Cooperation and conflict among primitive organisms. (Sigma Xi
lecture) American Scientist, 40:601-31.
1954
Speciation and variation in asexual fungi. Ann. N.Y. Acad. Sci.,
60: 1-82.
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BIOGRAPHICAL MEMOIRS
With Sung Huang Sun, l. Ehrlich, and L. E. Anderson. Criteria
of speciation in the genus Streptomyces. Ann. N.Y. Acad. Sci.,
60:102-23.
The spirit of science. Georgia Review, 8~4~:373-82.
1955
With Sung Huang Sun, L. E. Anderson, and I. Ehrlich. The iden-
tity of viomycin-producing cultures of Streptomyces. Bull.
Torrey Bot. Club, 82:108-17.
1956
With L. E. Anderson, l. Ehrlich, and Sung Huang Sun. Strains of
Streptomyces, the sources of azaserine, elaiomycin, griseoviridin,
and viridogrisein. Antibiotics and Chemotherapy, 6~2~: 100-115.
With L. M. Burkholder. Microbiological assay of B-12 in marine
solids. Science, 123: 1071-73.
With L. M. Burkholder. Vitamin B-12 in suspended solids and
marsh muds collected along the coast of Georgia. Limnol.
Oceanogr., 1:202-8.
Studies on the nutritive value of spartina grass growing in the
marsh areas of coastal Georgia. Bull. Torrey Bot. Club, 83:
327-34.
1957
With G. H. Bornside. Decomposition of marsh grass by aerobic
marine bacteria. Bull. Torrey Bot. Club, 84:366-83.
1958
With L. M. Burkholder. Antimicrobial activity of horny corals.
Science, 127:1174-75.
Studies on B vitamins in relation to productivity of the Bahia
Fosforescente, Puerto Rico. Bull. Mar. Sci. Gulf Caribb.,
8:201-23.
1959
Some microbiological aspects of marine productivity in shallow
waters. In: Proceedings of the Salt Marsh Conference, ed. by
R. A. Ragotzkie, pp. 70-72. Sapelo Island, Georgia, Marine
Institute, 1958. University of Georgia: Marine Institute.
OCR for page 21
PAUL RUFUS BURKHOLDER
~1
With L. M. Burkholder and l. A. Rivero. Some chemical constitu-
ents of turtle grass, Thalassia tested inum. Bull. Torrey Bot.
Club, 86:88-93.
With L. M. Burkholder and I. A. Rivero. Chlorophyll A in some
corals and marine plants. Nature, 183: 1338-39.
Organic nutrition of some mosses mowing, in nilrP rillt~l'-
gist, 62:6-15.
Antibiotics -the exploitation of microbial antagonisms is having
a challenging impact on medicine and challenging society.
Science, 129:1457-65.
With H. T. Odum and I. Rivero. Measurements of productivity
of turtle grass flats, reefs, and the Bahia Fosforescente of south-
ern Puerto Rico. Publications of the Institute of Marine Science,
University of Texas, Port Aransas, 6: 159-70.
- ~-~ ---a -- ret ace. Bryolo-
1960
Distribution of some chemical values in Lake Erie. In: Limnologi-
cal Survey of Eastern and Central Lake E,^ie, 1928-29, pp. 71-
110. Spec. Sci. Rep. Fish. no. 334, June 1960.
A survey of the microplankton of Lake Erie. In: Limnological Survey '
of Eastern and Central Lake Erie, 1928-29, pp. 123-44. Spec.
Sci. Rep. Fish. no. 334, tune 1960.
With L. M. Burkholder and L. A. Almodovar. Antibiotic activity
of some marine algae of Puerto Rico. Bot. Mar., 2:149-56.
General microbiology of the Antarctic. In: Science in Antarctica.
I. The Life Sciences in A ntarctica, pp. 129-37. Washington,
D.C.: National Academy of Sciences-National Research Council
Publication 839.
With M. D. Tendler. Studies on the thermophilic actinomycetes.
Appl. Microbiol., 9:394-99.
With L. M. Burkholder. Photosynthesis in some alcyonacean corals.
Am. [. Bot., 47: 866-72.
1961
With l. M. Siel~urth. Phytoplankton and chlorophyll in the Ger-
lache and Bransfield straits of Antarctica. Limnol. Oceanogr.,
6~1~:45-52.
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22
BIOGRAPHICAL MEMOIRS
1963
Some nutritional relationships among microbes of sea sediments
and waters. In: Symposium on Marine Microbiology, pp. 133-50.
Springfield, Ill.: Charles C. Thomas.
Radioactivity in some aquatic plants. Nature, 198:601-3.
Drugs from the sea. Armed Forces Chemical Journal, 17:6-16.
1965
With C. W. Dodge and L. M. Burkholder. Estudio de los liquenes
de Tierra del Fuego con especial consideration de suactividad
antibiotica. Centro de Investigacion de Biologia Marina, Buenos
Aires, Argentina. Contribuciones Cientificas no. 21, 1-24.
With R. M. Pfister. Numerical taxonomy of some bacteria isolated
from Antarctica and tropical seawaters. i. Bacteriol., 90: 863-72.
With E. F. Mandelli. Carbon assimilation of marine phytoplankton
in Antarctica. Proc. Natl. Acad. Sci., 54:437-44.
With E. F. Mandelli. Productivity of microalgae in Antarctic sea
ice. Science, 149: 872-74.
With A. Repak and l. Siebert. Studies on some Long Island Sound
littoral communities of microorganisms and their primary pro-
ductivity. Bull. Torrey Bot. Club, 92:378~02.
1966
With R. M. Pfister and F. H. Leitz. Production of a pyrole anti-
biotic by a marine bacterium. Appl. Microbial., 14:649-53.
With L. M. Burkholder and P. Genteno. Nutritive values of shrimp
flour. Nature, 211: 860-61.
With E. F. Mandelli. Primary productivity in the Gerlache and
Bransfield straits of Antarctica. Journal of Marine Research,
24:15-27.
1967
With L. M. Burkholder and L. R. Almodovar. Carbon assimilation
of marine flagellate blooms in neritic waters of southern Puerto
Rico. Bull. Mar. Sci. Gulf Caribb., 17: 1-15.
With E. F. Mandelli and P. Centeno. Some chemical properties of
Munida gregaria and Euphausia superba. journal of Agricul-
tural and Food Chemistry, 1 ~5 :718-20.
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PAUL RUFUS BURKHOLDER
~3
With G. M. Sharma. Studies on antimicrobial substances of sponges.
I. Isolation, purification, and properties of a new bromine-con-
taining antibacterial substance. l. Antibiot. (Japan), 20:200-203.
With G. M. Sharma. Studies on antimicrobial substances of sponges.
II. Structure and synthesis of a bromine-containing antibacterial
compound from a marine sponge. Tetrahedron Letters, 42:
4147-50.
With L. M. Burkholder. Primary productivity in surface waters of
the South Pacific ocean. Limnol. Oceanogr., 12:606-17.
1968
With L. M. Burkholder, A. Chu, N. Kostyk, and O. A. Roels. Fish
fermentation. Food Technology, 22: 1278-84.
Patterns of B vitamin requirements of neritic
Canadian journal of Microbiology, 14:537-43.
marine bacteria.
With H. G. Udell, I. Zarudsky, and T. E. Doheny. Productivity
and nutrient values of plants growing in the salt marshes of
the town of Hempstead, Long Island. Bull. Torrey Bot. Club,
96:42-51.
With T. E. Doheny. Biology of eelgrass. Contribution no. 3.
Hempstead, L.I.: Department of Conservation and Waterways,
Marine Laboratory.
Antimicrobial substances from the sea. In: Drugs from the Sea,
ed. by Hugo D. Freudenthal, pp. 87-112. Transactions of a
symposium held at the University of Rhode Island, 1967.
Washington, D.C.: Marine Technology Society.
With G. M. Sharma and B. Vig. Studies on antimicrobial sub-
stances of sponges. III. Chemical properties of some antibacterial
compounds from marine sponges. In: Drugs from the Sea, ed.
by Hugo D. Freudenthal, pp. 119-26. Transactions of a sym-
posium held at the University of Rhode Island, 1967. Wash-
ington, D.C.: Marine Technology Society.
With G. M. Sharma and L. Michaels. Coniodomin, a new antibiotic
from a dinoflagellate. l. Antibiot. Japan), 21:659-64.
With O. A. Roels. Biological oceanology at Lamont Geological
Observatory, pp. 72-85. New York State Science and Tech-
nology Foundation.
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24
BIOGRAPHICAL MEMOIRS
1969
With K. Ruetzler. Antimicrobial activity of some marine sponges.
Nature, 222:983-84. (L)
With G. M. Sharma. Antimicrobial agents from the sea. Lloydia,
32:466-88.
1970
With G. M. Sharma and B. Vig. Studies on antimicrobial substances
of sponges. IV. Structure of a bromine-containing compound
from a marine sponge. J. Org. Chem., 35:2823-26.
With P. Hargraves and R. Brody. Phytoplankton of the lesser An-
tilles region. Bull. Mar. Sci. Gulf Caribb., 20:331-49.
Some biomedical aspects of marine microbiology. In: Food-Drugs
from the Sea Proceedings 1969, ed. by H. W. Youngken, Jr.,
pp. 255-79. Held at the University of Rhode Island, 1969.
Washington, D.C.: Marine Technology Society.
With G. M. Sharma and B. Vig. Antimicrobial substances of marine
sponges. IV. In: Food-Drugs from the Sea Proceed ings 1969,
ed. by H. W. Youngken, Jr., pp. 307-10. Held at the University
of Rhode Island, 1969. Washington, D.C.: Marine Technology
Society.
With G. Cintron and W. S. Maddux. Consequences of brine pollu-
tion in the Bahia Fosforescente, Puerto Rico. Limnol. Oceanogr.,
15(2):246~9.
With E. F. Mandelli, T. Doheny, and R. Brady. Primary produc-
tivity in coastal waters of southern Long Island, New York.
Mar. Biol., 7~2~: 153-60.
1971
With L. Almodovar. Species composition and productivity of man-
grove algae communities in Puerto Rico. Florida Scientist,
36:66-74.
With G. M. Sharma. Structure of dibromophakellin, a new bro-
mine-containing alkaloid from the marine sponge Phakellia
flabellata. Chemical Communications (London), pp. 151-52.
With L. Almodovar and L. M. Burkholder. Nutritive constituents
of some Caribbean marine algae. Bot. Mar., 14~2~:132-35.
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PAUL RUFUS BURKHOLDER
25
With T. Doheny. Proximate analysis and amino acid composition
of some marine phytoplankton and bacteria. Contribution
no. 8. Hempstead, L.I.: Department of Conservation and
Waterways, Marine Laboratory.
The ecology of marine antibiotics. In: Coral Reefs of The World,
ed. by R. Endean and 0. A. tones, pp. 117-82. New York:
Academic Press, Inc.
1972
With T. E. Doheny. Use of selected enrichment experiments in
predicting the eutrophication of an estuary. Contribution no.
7. Hempstead, L.I.: Department of Conservation and Waterways,
Marine Laboratory.
With R. Brody and A. E. Dammann. Some phytoplankton blooms
in the Virgin Islands. Caribbean Journal of Science, 12:23-28.
\
Representative terms from entire chapter:
paul rufus
