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GEORGE WELLS BEADLE
October 22, 1903-fune 9, l 989
BY NORMAN H. HOROWITZ
GEORGE BEADLE WAS A GIANT in the field of modern
genetics. He initiated the great series of advances made
between 1941 and 1953 that brought the era of classical ge-
netics to a close and launched the molecular age. For this
achievement he received many honors, including the Nobel
Prize. He was elected to the National Academy of Sciences in
1944 and served on its Council from 1969 to 1972.
BeacIle also had a clistinguished career as an academic
administrator. When he retired in 1968, he was President of
The University of Chicago. Long years in administration,
however, dial not dampen his love of experimental genetics,
and after his retirement he resumed experimental work on
a favorite subject—the origin of maize. In 1981, he gave up
research altogether because of increasing disability from the
Alzheimer's disease that eventually ended his life.
EDUCATION AND EARLY ElFE
Beadle his oldest friends usually called him by his boy-
hood nickname, "Beets" was born in Wahoo, Nebraska, to
Hattie Albro and Chauncey Elmer Beadle, and he cried in
Pomona, California, at age eighty-five. He grew up on his
father's forty-acre farm near Wahoo. The farm was a mode]
for farms its size and was so designated by the U. S. Depart-
27
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BIOGRAPHICAL MEMOIRS
ment of Agriculture in 1908. Beets' mother died when he
was four years old, and he and his brother ant! sister were
raised by a succession of housekeepers.
As a boy he worked on the farm, and he retained the skills
he learned as a gardener and beekeeper there, and his hand-
iness with tools, for the rest of his life. Gardening remained
one of his greatest pleasures, and the victory garden he grew
around his home at Stanford;during the War produced
enough for two families. This garden included beehives, but
Beets wouldn't eat the honey, saying he had been stung too
many times as a boy. He loved corn, on the other hand, and
raised several kinds, including a small Mexican variety that
gave his garden the distinction of having the earliest sweet
corn at Stanford. After his retirement to Pomona in 1982,
he derived much pleasure from growing flowers, a hobby he
pursued as long as his health permitted.
Beets did well in school and was inspired to go on to col-
lege by his high school science teacher, Bess MacDonald (the
debt to whom he acknowledged more than once in later
years). Despite his father's opinion that a farmer did not need
all that education, he entered the University of Nebraska Col-
lege of Agriculture in 1922. He graduated in 1926 with a
B.S. degree ant! stayed on for another year to work for a
master's degree with Franklin D. Keim.
His first scientific publication, with Keim, dealt with the
ecology of grasses. At some point along the way under Keim's
beneficent influence, Beets became interested in fundamen-
tal genetics and was persuaded to apply to the graduate
school at Cornell University instead of going back to the
farm. He entered Cornell in 1927 with a graduate assistant-
ship and shortly afterward joined R. A. Emerson's research
group on the cytogenetics of maize.
Corn genetics was new and exciting for Beets, and Emer-
son and his team which inclucled Barbara McClintock and
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GEORGE WELLS BEADLE
29
Marcus Rhoades were inspiring. The result was that in the
following five years, Beets published no fewer than fourteen
papers dealing with investigations on maize, all begun while
he was a graduate student at Cornell.
In 1928 he married Marion Hill, a graduate student in
botany at Cornell, who assisted him with some of his early
corn research. Their son, David, was born in 1931.
Beets received his Ph.D. in 1931 and was awarded a Na-
tional Research Council Fellowship to do postdoctoral work
in T. H. Morgan's Division of Biology at the California Insti-
tute of Technology. At Caltech, while finishing the work on
maize cytogenetics he had started at Cornell on genes for
pollen sterility, sticky chromosomes, failure of cytokinesis,
and chromosome behavior in maize-teosinte hybrids (a sub-
ject he wouIc! return to in his retirement) Beadle also began
doing research on Drosophila. Out of it would come one of
the most interesting investigations of his career.
DROSOPHILA STUDIES: CROSSING OVER
VERMILION AND CINNABAR
BeadIe's Drosophila studies at Caltech were concerned
with the results of crossing over within various chromosomal
rearrangements. The important study of crossing over in
attached-X chromosomes he conducted with Sterling Emer-
son showed that exchanges occur at random between any two
non-sister chromatics. Another, reported jointly with A. H.
Sturtevant (in a paper called. "monumental" by E. B. Lewis),
was the first systematic investigation of crossing over and dis-
junction in chromosomes bearing inversions.
In 1934, Boris Ephrussi arrived at Caltech from Paris to
study Drosophila genetics with Morgan and Sturtevant. He
was just two years oIcler than Beadle and they became close
friends. Ephrussi soon communicated to Beadle his own in-
terest in the problem of gene action, and the two planned a
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BIOGRAPHICAL MEMOIRS
collaborative study on Drosophila that would use Ephrussi's
skill in the techniques of tissue culture and transplantation.
In mid-1935, the two men went to Paris to carry out ex-
periments in Ephrussi's laboratory at the Institut de Biologie.
Though their attempts to grow imaginal discs in tissue cul-
ture failed, they succeeded in devising a method for trans-
planting discs from one larva to another that allowed! the
discs to continue to develop.
Before year's end, they had gone as far as they could with
this methodology and had worked out a hypothesis to ac-
count for the interaction they observed between the vermi-
lion and cinnabar genes in transplanted flies. The results,
they showecl, could be explainecl by the following assump-
tions: (~) the normal alleles of the two genes control the pro-
duction of two specific substances, called the v+- and cn+-
substances, both necessary for brown eye-pigment forma-
tion; (2) the v+-substance is a precursor of the cn+-substance;
and (3) gene mutation blocks formation of the corresponding
substance. It was not clear until much later that the two sub-
stances are actually precursors of the pigment, and Ephrussi
and Beadle frequently referred to them as "hormones."
At the time, this small step was a great advance in the
science of genetics, for it suggested that development could
be broken clown into series of gene-controlled chemical re-
actions an idea that cried out for further investigation. It
implantec! in Beadle the germ of the one gene-one enzyme
idea that he later brought to full flower. But first, the two eye-
color substances hac! to be identifiecI, a process that took five
years. By that time, Beets was hunting bigger game.
Following his return from Paris, Beadle moved to Har-
vard University as an assistant professor. There, on a few
brief occasions, he met a young woman who wouIc! later be-
come my wife and who remembered him fondly afterwards
as the only member of the Harvard faculty who spoke to
Radcliffe undergraduates at Biology Departmental teas.
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GEORGE WELLS BEADLE
BEADLE AND TATUM
31
Beadle left Harvard the following year (1937) for Stan-
ford University, where he had accepted an appointment as
professor of biology. He was joined by biochemist Edwarc! L.
Tatum (1909-1975) as a research associate.'
Over the next three years, Tatum contributed his skills to
the work of isolating and identifying the two eye-color sub-
stances. With others, they established that the two substances
were derivatives of tryptophan. By 1940, Tatum had ob-
tained a crystalline preparation of the v+-substance, but he
and Beadle were beaten to the identification by Butenancit,
Weidel, and Becker, who hac! adopted the simple procedure
of testing known metabolites of tryptophan for their biolog-
ical activity. These researchers found that kynurenine is ac-
tive as the v+-substance and that OH-kynurenine is active as
the cn+-substance. Much later it was shown that condensation
of two molecules of OH-kynurenine forms brown pigment.
Despite this setback in the laboratory, the years from 1937
to 1939 were not wasted for Beadle. During this period, he
joinec} A. H. Sturtevant in writing a superlative textbook, An
Introa~uction to Genetics (1939,5), praised by]. A. Moore as "the
complete statement of classical genetics."
NEUROSPORA CRASSA AND GENE ACTION
As a result of his Drosophila experience it became clear
to Beadle that an entirely different method was needed to
make headway with the problem of gene action. No other
nonautonomous traits were known in Drosophila, and the
autonomous ones—of which there were many were of such
towering complexity from the biochemical standpoint that it
was hopeless to attempt to reduce them to their individual
chemical steps.
Beets enjoyed telling how the solution to this problem
came to him while he was listening to Tatum lecture in a
' See p. 356 for Joshua Lederberg's memoir of Tatum.
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BIOGRAPHICAL MEMOIRS
course on comparative biochemistry. Microbial species, Beets
learned, differ in their nutritional requirements even though
they share the same basic biochemistry. If these differences
were genetic in origin, he thought to himself, it should be
possible to induce gene mutations that would produce new
nutritional requirements in the test organism. Such an ap-
proach, if successful, would allow the researchers to identify
genes governing known biochemical compounds immedi-
ately, as opposed to the years needed to identify the unknown
substances controlled by the usual kinds of genes, including
most of those then known.
What was needed for such an undertaking was an orga-
nism that was genetically workable that could be grown on a
chemically defined medium. Beadle knew just the organism.
While still a graduate student at Cornell, he had heard about
Neurospora crassa, the red bread mold. B. O. Docige had come
to the campus from the New York Botanical Garden to give
a lecture on Neurospora. Beets remembered clearly that the
lecture dealt with the genetics of the organism, including re-
sults on first- and second-division segregations of the mating-
type and other loci. Even years later Beets was pleased to
recall that he and a few other graduate students had been
able to explain to the skeptical Dodge that his data could be
explained by crossing over or the lack of it between the
gene and its centromere.
Dodge had played an important role in the history of
Neurospora. It was he who discovered that the ascospores
could be germinated by heat, thus closing its life cycle and
making the organism accessible for genetic study. He also did
basic studies on its genetics and was enthusiastic about its
possibilities for genetic research. He convinced T. H. Mor-
gan, a close friend, to take some cultures with him to Pasa-
dena when, in 192S, Morgan went out to found the Division
of Biology at Caltech. Dodge, according to Beadle, told
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GEORGE WELLS BEADLE
33
Morgan that Neurospora would be "more important than
Drosophila some day," and, in Pasadena, Morgan assigned
the cultures to graduate student Car! Linclegren, for his thesis
in genetics. Lindegren studier} the relation between first-
and second-division segregations anct crossing over. He com-
pleted his thesis in 1931, the year Beadle arrived at Caltech.
In 1940 the question of the nutritional requirements of
Neurospora was still an open one. Previous workers had used
nutrient agar as the growth medium, but this would not clo
for the experiment Beadle had in mincI. Related fungi, how-
ever, were known to have simple requirements, and Tatum
soon showed that Neurospora would grow on a synthetic me-
dium containing sugar, salts, and a single growth factor-
biotin—thenceforth referred to as "minimal medium." For-
tunately, purified concentrates of biotin tract recently become
available, ant] nothing now stood in the way of an experi-
mental test of BeacIle's idea.
The final step was to clear the Drosophila cultures out of
the Stanford lab and convert it into a laboratory for Neuro-
spora genetics. The plan was to x-ray one parent of a cross
and collect offspring (haploid ascospores isolatecl by hand)
onto a medium designed to satisfy the maximum number of
possible nutritional requirements (so-called "complete me-
lium"~. The resulting cultures wouIc! next be transferred to
minimal medium. Growth on complete medium, combined
with failure to grow on minimal medium, was to be taken as
presumptive evidence of an induced nutritional require-
ment. The requirement would be identified, if possible, and
the culture would be crossest to wild type to determine its
heritability.
This scheme, in its time, was breathtakingly claring. Some
nongeneticists still suspected that genes governed only trivial
biological traits, such as eye color anct bristle pattern, while
important characters were determined in the cytoplasm by
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BIOGRAPHICAL MEMOIRS
an unknown mechanism. Many geneticists believed that gene
action was far too complex to be resolved by any simple ex-
periment. Indeed, the outcome of Beadle and Tatum's trial
run was so uncertain that they agreed at the outset to test
5,000 ascospores before giving up the project and- to avoid
early disappointment isolated and stored over a thousand
spores before testing any of them.
Success came with spore no. 299, which gave rise to a
culture that grew on complete but not on minimal medium
unless this was supplemented! with pyridoxine. This mutant
was followed by others showing requirements for thiamine
and p-aminobenzoic acid, respectively. All three require-
ments were inherited as single-gene defects in crosses to wild
type. These mutants were the subject of the first Neurospora
paper by Beadle and Tatum (1941,21. Before long, mutants
requiring amino acids, purines, and pyrimidines were also
found, and the science of biochemical genetics had been
born.
Beadle recognized that he and Tatum hac! discovered a
new florid of genetics and that more hands would be needed
to explore it. Early in the fall of 1941 he came to Caltech to
give a seminar on the new discoveries and to recruit a couple
of research associates to join the enterprise. Since the first
BeadIe-Tatum paper on Neurospora hac! yet to be published,
no one in the audience hac! an inkling of what was to come.
The seminar was memorable. ~ recorded my recollection
of it in an article written in honor of BeadIe's seventieth
birthday:
"The talk lasted only half an hour, and when it was suddenly over, the
room was silent. The silence was a form of tribute. The audience was
thinking: Nobody with such a discovery could stop talking about it after
just thirty minutes" there must be more. Superimposed on this thought
was the realization that something historic had happened. Each one of us,
I suspect, was mentally surveying, as best he could, the consequences of
the revolution that had just taken place. Finally, when it became clear that
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GEORGE WELLS BEADLE
35
Beadle had actually finished speaking, Prof. Frits Went—whose father had
carried out the first nutritional studies on Neurospora in Java at the turn
of the century got to his feet and, with characteristic enthusiasm, ad-
dressed the graduate students in the room. This lecture proved, said Went,
that biology is not a finished subject- there are still great discoveries to be
made!" Neurospora Newsletter 20~1973~:4-6
BEADLE AS LABORATORY HEAD
David Bonner and ~ accepted appointments with Beadle
and joined his group at Stanford the following year. Later,
H. K. Mitchell and Mary Houlahan (Mitchell) came. There
were also graduate students (including A. H. Doermann and
Adrian Srb) and a steady turnover of visitors in the lab.
The next four years were the most exciting of my life, and
~ imagine the same was true for everyone else in the lab.
Before the Neurospora revolution, the idea of uniting ge-
netics and biochemistry had been only a dream with a few
scattered observations. Now, biochemical genetics was a real
science, and it was all new. Incredibly, we privileged few had
it all to ourselves. Every day brought unexpected new results,
new mutants, new phenomena. It was a time when one went
to work in the morning wondering what new excitement the
clay would bring.
Beadle presided over this scientific paradise with the en-
thusiasm, intelligence, and good humor that characterized
everything he did. He was a popular and much admired boss.
He worker! in the lab with everyone else. He especially en-
joyed working with his hands, and he had plenty of oppor-
tunity to indulge himself in this regard.
The laboratories were located in the basement (the "cat-
~ ~ · ~ ~
.
acombs ) ot Jordan tiall, a location that gave them a certain
remoteness from the campus. There were a bench and lathe
in the lab, and Beets used these to make small equipment
and do minor repairs around the place; he called the campus
shops only for major work and did as much as possible him-
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BIOGRAPHICAL MEMOIRS
self. ~ came to work early one morning and found him paint-
ing one of the rooms.
All this was in addition to his research and his teaching
Buttes as a professor of biology. He always did more than
anyone else. ~ recall going to a lab picnic at the beach one
summer day, over the coast range of hills from the Stanford
campus. We were bicycling to save gas, huffing and wheezing
(we had no gears then); Beets differed from the rest of us
only in that he was carrying a watermelon on his handlebars.
Beets knew his responsibilities and took them seriously. It
was wartime, and he concerned himself with all that implied
for the pursuit of fundamental research. He had to find fi-
nancial support for the program while trying to keep his
group together. He succeeded on both scores, obtaining sup-
port from both the Rockefeller and Nutrition foundations—
support that continued throughout the war and even after-
wards. The Committee on Medical Research of the Office of
Scientific Research and Development classified the Neuro-
spora program as essential to the war effort. As ~ recall, no
senior researcher or graduate student was drafted, although
some of us were called up for physical examinations.
Practical applications of Neurospora research were of po-
tential utility to the war effort in developing bioassays for
vitamins and amino acids in preserved foods, and in search-
ing for new vitamins and amino acids. Although the major
thrust of the lab remained basic science, we worked on both
these applications during the war years. Toward the end of
World War Il. Beadle was asked by the War Production
Board to devote part of the effort of the lab to seeking mu-
tants of Penicillium with increased yields of penicillin. He
complied, of course, but we were not successful in this en-
deavor.
The biochemical and genetic studies carried out between
1941 and 1945 on Neurospora mutants in the Stanford lab-
oratory showed that the biosynthesis of any given substance
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GEORGE WELLS BEADLE
HONORS AND DISTINCTIONS
HONORARY DEGREES
Doctor of Science
1947 Yale University
1949 University of Nebraska
1952 Northwestern University
1954 Rutgers University
1955 Kenyon College
1956 Wesleyan University
1959 Birmingham University
1959 Oxford University
1961 Pomona College
1962 Lake Forest College
1963 University of Rochester
1963 University of Illinois
lD64 Brown University
1964 Kansas State University
1964
1966
1967
1970
1971
1972
1973
1975
1976
University of Pennsylvania
Wabash College
Syracuse University
Loyola University, Chicago
Hanover College
Eureka College
Butler University
Gustavus Adolphus College
Indiana State University
I-egum Doctor (LL .D.)
1962 University of California, Los Angeles
1963 University of Miami
1963 Brandeis University
1966 Johns Hopkins University
1966 Beloit College
1969 University of Michigan
43
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BIOGRAPHICAL MEMOIRS
Litterarum Humaniorum Doctor (~.H.D.)
1966 Jewish Theological Seminary of America
1969 DePaul University
1969 University of Chicago
1969 Canisius College
1969 Knox College
1971 Roosevelt University
1971 Carroll College
Doctor of Public Service
1970 Ohio Northern University
AWARDS
1950 Lasker Award
1951 Dyer Award
1953 Emil Christian Hansen Prize (Denmark)
1958 Albert Einstein Commemorative Award in Science
1958 Nobel Prize in Physiology or Medicine (with E. L. Tatum
1959
1960
1967
1967
and J. Lederberg)
National Award, American Cancer Society
Kimber Genetics Award
Priestley Memorial Award
Edison Prize, Best Science Book for Youth (with Muriel
Beadle)
1972 Donald Forsha {ones Medal
1984 Thomas Hunt Morgan Medal
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GEORGE WELLS BEADLE
PROFESSIONAL AND HONORARY SOCIETIES
45
Genetics Society of America (president, 1945)
American Association for the Advancement of Science (president,
1955)
National Academy of Sciences (Council, 1969-1972)
American Philosophical Society
American Academy of Arts and Sciences
Royal Society
Danish Royal Academy of Sciences
Japan Academy
Instituto Lombardo di Scienze e Lettre (Milan)
Genetical Society of Great Britain
Indian Society of Genetics and Plant Breeding
Indian Natural Science Academy
Chicago Horticultural Society (president, 1968-1971)
Phi Beta Kappa
Sigma Xi
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BIOGRAPHICAL MEMOIRS
SELECTED BIBLIOGRAPHY
1927
With F. D. Keim. Relation of time of seeding to root development
and winter survival of fall seeded grasses and legumes. Ecology,
8:251-64.
1928
With B. McClintock. A genie disturbance of meiosis in Zea mays.
Science, 68:433.
1929
Yellow stripe A factor for chlorophyll deficiency in maize located
in the Pr pr chromosome. Am. Nat., 68: 189-192.
A gene for supernumerary mitoses during spore development in
Zea mays. Science, 70:406-7.
1930
Heritable characters in maize. I. Hered., 21:45-48.
Genetical and cytological studies of Mendelian asynapsis in Zea
mays. Cornell Univ. Memoir, 129:3-23.
A fertile tetraploid hybrid between Euchlaena persons and Zea mays.
Am. Nat., 69:190-92.
1931
A gene in Zea mars for failure of cvtokinesis during meiosis. (~lvrol
3: 142-55.
A gene in maize for supernumerary cell divisions following
meiosis. Cornell Univ. Memoir, 135:3 -12.
1932
A possible influence of the spindle fibre on crossing-over in Dro-
sophila. Proc. Natl. Acad. Sci. USA, 18:160-65.
A gene in Zea mays for failure of cytokinesis during meiosis. Cytol.,
3:142-55.
Genes in maize for pollen sterility. Genet., 17:413-31.
The relation of crossing over to chromosome association in Zea-
Euchlaena hybrids. Genet., 17:481-501.
Studies of Euchlaena and its hybrids with Zeal 1. Chromosome be-
, c, , ,
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GEORGE WELLS BEADLE
47
havior in Euchlaena mexicana and its hybrids with Zea mays.
ZIAVA, 62:291-304.
With R. Emerson. Studies of Euchlaena and its hybrids with Zeal
II. Crossing over between the chromosomes of Euchlaena and
those of Zeal ZIAVA, 62:305 -15.
A gene for sticky chromosomes in Zea mays. ZIAVA, 63: 195-217.
1933
Studies of crossing-over in heterozygous translocations in Droso-
phila melanogaster. ZIAVA, 65: 111-28.
With S. Emerson. Crossing-over near the spindle fiber in attached-
X chromosomes of Drosophila melanogaster. ZI AVA, 65: 129-40.
Further studies of asynaptic maize. Cytol., 4:269-287.
Polymitotic maize and the precocity hypothesis of chromosome
conjugation. Cytol., 5: 118-21.
1934
Crossing-over in attached-X triploids of Drosophila melanogaster. ].
Genet., 29:277-309.
1935
Crossing over near the spindle attachment of the X chromosomes
in attachecl-X triploids of Drosophila melanogaster. Genet.,
20: 179-91.
With S. Emerson. Further studies of crossing-over in attached-X
chromosomes of Drosophila melanogaster. Genet., 20: 192-206.
With R. A. Emerson and A. C. Fraser. A summary of linkage stud-
ies in maize. Cornell Univ. Memoir, 180:3-83.
With A. H. Sturtevant. X chromosome inversions and meiosis in
Drosophila melanogaster. Proc. Natl. Acad. Sci. USA, 21 :384-90.
With B. Ephrussi. La transplantation des disques imaginaux chez
la Drosophile. C. R. Acad. Sci., 201:98.
With B. Ephrussi. Differenciation de la couleur de Ceil cinnabar
chez la Drosophile. C. R. Acad. Sci., 201:620.
With B. Ephrussi. La transplantation des ovaires chez la Droso-
phile. Bull. Biol. Belg., 69:492-502.
With B. Ephrussi. Sur les conditions de l'auto-differenciation des
caracteres mendeliens. C. R. Acad. Sci., 201:1148.
With B. Ephrussi. Transplantation in Drosophila. Proc. Natl. Acad.
Sci. USA, 21:642-46.
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~8
BIOGRAPHICAL MEMOIRS
1936
With B. Ephrussi. A technique of transplantation for Drosophila.
Am. Nat., 70:218-25.
With B. Ephrussi. The differentiation of eye pigments in Droso-
phila as studied by transplantation. Genet., 21:225-47.
With B. Ephrussi. Development of eye colors in Drosophila: Trans-
plantation experiments with suppressor of vermilion. Proc.
Natl. Acad. Sci. USA, 22:536-40.
With B. Ephrussi and C. W. Clancy. Influence de la lymphe sur la
couleur des yeux vermilion chez la Drosophile. C. R. Acad. Sci.,
203:545.
With A. H. Sturtevant. The relation of inversions in the X chro-
mosome of Drosophila melanogaster to crossing-over and disjunc-
tion. Genet., 21:554-604.
With Th. Dobzhansky. Studies on hybrid sterility IV. Transplanted
testes in Drosophila pseudoobscura. Genet., 21:832-40.
With B. Ephrussi. Development of eye colors in Drosophila: Stud-
ies of the mutant claret. I. Genet., 33:407-10.
1937
With B. Ephrussi. Development of eye colors in Drosophila: Trans-
plantation experiments on the interaction of vermilion with
other eye colors. Genet., 22:65-75.
With B. Ephrussi. Development of eye colors in Drosophila: Dif-
fusible substances and their interrelations. Genet., 22:76-85.
With B. I:phrussi. Developpement des couleurs des yeux chez la
Drosophile: Influence des implants sur la couleur des yeux de
l'hote. Bull. Biol. Belg., 71:75-90.
With B. Ephrussi. Developpement des couleurs des yeux chez la
Drosophile: Revue des experiences de transplantation. Bull.
Biol. Belg., 71 :54-74.
With C. W. Clancy. Ovary transplants in Drosophila melanogaster:
Studies of the characters singed, fused, and female-sterile. B~ol.
Bull, 72:47-56.
With B. Ephrussi. Development of eye colors in Drosophila: The
mutants bright and mahogany. Am. Nat., 71:91-95.
The development of eye colors in Drosophila as studied by trans-
plantation. Am. Nat., 71:120-26.
With K. V. Thimann. Development of eye colors in Drosophila:
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GEORGE WELLS BEADLE
49
Extraction of the diffusible substances concerned. Proc. Natl.
Acad. Sci. USA, 23:143-46.
Development of eye colors in Drosophila: Fat bodies and Malpigh-
ian tubes as sources of diffusible substances. Proc. Natl. Acad.
Sci. USA, 23:146-52.
With C. W. Clancy and B. Ephrussi. Development of eye colours in
Drosophila: Pupal transplants and the influence of body fluid
on vermilion. Proc. R. Soc. London, 122:98-105.
The inheritance of the color of Malpighian tubes in Drosophila me-
lanogaster. Am. Nat., 71:277-79.
With B. Ephrussi. Ovary transplants in Drosophila melanogaster.
Meiosis and crossing-over in superfemales. Proc. Natl. Acad.
Sci. USA, 23:356-60.
With B. Ephrussi. Development of eye colors in Drosophila: Pro-
duction and release of en+ substance by the eyes of different
eye color mutants. Genet., 22:479-83.
Chromosome aberration and gene mutation in sticky chromosome
plants of Zea ways. Cytol. Fujii Jubilee, pp. 43-56.
Development of eye colors in Drosophila: Fat bodies and Malpigh-
ian tubes in relation to diffusible substances. Genet., 22:587-
611.
1938
With L. W. Law. Influence on eye color of feeding diffusible sub-
stances to Drosophila melanogaster. Proc. Soc. Exp. Biol. Med.,
37:621-23.
With R. Anderson and J. Maxwell. A comparison of the diffusible
substances concerned with eye color development in Drosophila,
Ephestia and Habrobracon. Proc. Natl. Acad. Sci. USA, 24:80-
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Representative terms from entire chapter:
eye colors
