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DONALD FORSHA JONES
April 16, 1890-June 19, 1963
BY PAUL C. NIANGELSDORF
I E; THERE were a Nobel prize in agriculture as there is in medi-
cine, it would undoubtedly have been awarded many years
ago to Donald F. Jones for his part in the development of hy-
brid corn. This is an achievement in applied genetics that has
sparked an agricultural revolution in the United States. It has
affected agricultural production in many other parts of the
world as well, including the countries of Latin America, where
corn is the principal source of food, and those of southern
Europe, where corn is an important crop. To this monumental
achievement Dr. Jones made four distinct contributions.
{ones's most practical contribution was the invention in
1917 of the double-cross method of hybrid seed production.
George H. Shull had shown previously that self-pollination in
corn, a naturally cross-pollinated plant, resulted in the isolation
of inbred strains that were uniform and true breeding. Follow-
ing Johanssen, he called these "pure lines." They were much
less vigorous than the open-pollinated varieties from which they
had been derived. However, when two such lines were crossed,
the Fat hybrids were uniform, like their inbred parents, but
much more vigorous and in some cases more productive than
~ Since this was written, a notable contribution to agriculture has been
recognized by the award in 1970 of the Nobel Peace Prize to Dr. Norman E.
Borlaug for his work on the breeding of highly productive new dwarf varieties
of wheat for the underdeveloped regions of the globe.
135
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136
BIOGRAPHICAL MEMOIRS
the original open-pollinated varieties. Shull recognized that
he had discovered a revolutionary new method of corn breed-
ing, and he made determined efforts to persuade agronomists
to adopt and practice it, but with little success. Shull's method
of crossing two inbred strains had one serious shortcoming: The
hybrid seed was borne on the weak, unproductive plants of the
inbred strains used as the female parents. Since production
was low, the hybrid seed was costly—too expensive for the aver-
age farmer to afford. Edward M. East, who independently of
Shull, had observed the effects of inbreeding and crossbreeding
in corn, doubted that the crossing of inbred strains to produce
hybrid seed would ever become practical.
East, who had participated for several years in the famous
experiments at the University of Illinois on selection for chem-
ical composition of corn, initiated a corn-breeding program
at the Connecticut Agricultural Experiment Station in New
Haven in 1905. When East moved to the Bussey Institution of
Harvard as a professor in 1909 he was succeeded by Herbert K.
Hayes, who had been his assistant in New Haven and later be-
came his graduate student at Harvard. When Hayes accepted a
position at Minnesota, ~ones, then a graduate student at Har-
vard, took charge under East's direction of the Connecticut
corn program. He moved to New Haven early in 1915.
During his first year at New Haven Jones did little more
than maintain the experiments that East and Hayes had initi-
ated. But later, impressed by the weakness of the inbred strains
and their feeble production of grain and by the vigorous pro-
duction of their single crosses, he decided to try crossing two
of the single crosses. Thus in 1917 he crossed the single cross
of two strains of Chester's Leaming with a single cross of two
strains of Burr White. Grown in 1918 this cross, which later
came to be known as a "double cross," yielded more than either
of its single-cross parents and considerably more than the best
open-pollinated varieties.
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DONALD FORSHA JONES
137
Jones recognized that double crosses offered a means of
producing hybrid seed corn at a cost within the means of
progressive farmers. This fact was then quickly recognized by
other corn breeders, including Henry Wallace, H. K. Hayes,
and F. D. Richey. Within a few years corn-breeding programs,
including the isolation of inbred strains and testing of single
and double crosses, had been initiated by the U.S. Department
of Agriculture and many of the state experiment stations. By
1933 hybrid corn was in commercial production on a substantial
scale, and the USDA began to collect statistics on it. By 1949,
78 percent of the total U.S. corn acreage was planted in hybrid
corn. By 1959, more than 95 percent was in hybrid corn, and
the average yield of corn in the United States was double that
of 1929.
Ironically, it turns out that tones was not the first to make
double crosses in corn. At the Heterosis Conference held in
Ames, Iowa, in 1950 Shull reported that he had made a number
of such crosses in 1910 and grown the hybrids in 1911. One
of these crosses yielded considerably more than any of the
single crosses. Shull, however, apparently saw no particular
significance in the fact at that time; in 1950 he asked no credit
for making double crosses before tones and reported it only
as a historical fact.
Ironically, too, once the use of double crosses established
the production of hybrid corn on a highly successful scale,
corn breeders found that, by developing more vigorous inbred
strains than those isolated by Shull, East, and {ones, it was
possible to employ single crosses instead of double crosses in the
production of hybrid seed corn. Today much of the hybrid
corn in the United States is represented either by single crosses
or by three-way crosses, the latter being crosses of single crosses
by inbred strains.
It is obvious that hybrid corn, which has revolutionized the
production of corn in the United States and other parts of the
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BIOGRAPHICAL MEMOIRS
world, is a method of exploiting hybrid vigor or heterosis.
Thus heterosis must be considered, along with replication of
the hereditary material (DNA) and photosynthesis, one of the
three principal biological phenomena underlying the practice
of agriculture. What, then, is the genetic basis of heterosis?
In the same year, 1917, that Jones made his first double
cross, which solved the problem of hybrid seed corn production,
he published a theory explaining the phenomenon of heterosis.
He was at that time twenty-seven years old. Jones's theory
applied the concept of chromosomal linkage of Morgan and
his students to the earlier concrete explanation of hybrid vigor
by Keeble and Pellew. These last two had explained the in-
creased height of a cross of two varieties of peas over their
parents in Mendelian terms. It was linkage that explained why
it was not possible to combine in one race the favorable domi-
nant factors of both parents.
Jones's theory probably gave as much stimulus to hybrid
corn breeding as did his double-cross method of seed produc-
tion. Hybrid corn became a practical reality when his method
of seed production made it feasible and his theory of heterosis
made it plausible. This combination was difficult for even the
most conservative agronomists to resist. Seldom in the history
of agriculture has one man made two such significant contribu-
tions, one in theory and the other in practice.
A third contribution to hybrid corn production was one in
which I had the privilege of participating. It involves a method
of employing cytoplasmic male sterility, of a type first described
in corn by Marcus M. Rhoades, to avoid the operation of
emasculation, commonly known as "detasseling," in the seed-
production fields. Detasseling has been called the "peskiest and
most expensive part of producing hybrid seed corn." Before
cytoplasmic sterility was employed to avoid it, some one
hundred and twenty-five thousand workers were engaged on the
peak day of the season in removing tassels from corn plants.
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DONALD FORSHA JONES
139
By employing cytoplasmic male sterility, it was possible to
eliminate as much as two thirds of the detasseling operation.
To ensure fertility in the farmer's crop, sterile hybrids were
mixed with fertile hybrids, involving the same inbred strains
produced in the conventional way.
tones made still another contribution to hybrid corn pro-
duction when he employed fertility-restoring genes to overcome
the cytoplasmic sterility in the farmer's crop. This method
employs hereditary factors in the cytoplasm to make corn sterile
when sterility is a distinct asset and uses hereditary factors on
the chromosomes to make it fertile when fertility is essential. A
patent on the method of using genetic restorers in hybrid-seed-
corn production was issued to Jones in 1956. It was the first
patent on a genetic technique to be granted in the United
States. The validity of the patent was challenged by the seed
corn industry, but finally, after long and extensive litigation,
an amicable agreement was reached: The patent's validity was
generally accepted, and royalties on its use were paid.
For many years the method of employing cytoplasmic male
sterility to avoid the operation of detasseling and using fertility-
restoring genes to overcome this sterility in the farn~er's crop
was phenomenally successful. It not only drastically reduced
the labor required in producing hybrid seed but also eliminated
the reduction in yield of hybrid seed caused by the removal of
one or more leaves in the detasseling operation. The method
has probably been a factor also in making possible on an ex-
tensive scale the replacement of double crosses by higher-yield-
ing single crosses. There is a danger—susceptibility to plant
diseases—inherent in growing these genetically uniform hybrids,
as Jones pointed out in an article in American Naturalist in
1958. But their use, combined with increased applications of
fertilizer, to which the new hybrids were responsive, has un-
doubtedly contributed to the spectacular increase in average
yields in the decade 1959-1969. This is shown in the following
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140
BIOGRAPHICAL ME M OIRS
table, which also demonstrates the remarkable progress that has
occurred between 1929 and 1969 in increasing yields of corn
in the United States.
Acre Yields of Corn in the United States
Percent Increase
Percent Bushels over Previous
Year Hybrid Corn per Acre Decade
1929 0 25.7
1939 22.9 29.7 16
1949 78.3 37.8 27
1959 94.S 51.5 36
1969 99+ 80.0 55
Eventually, however, the method became, at least for several
years, the victim of its own success. In 1969 there were reports
that corn hybrids carrying the Texas cytoplasmic male sterility,
the type almost universally employed, were becoming sus-
ceptible to the southern corn blight fungus, Helminthosporium
mayd is. Since susceptibility to a disease determined by the
cytoplasm had never previously been observed in the United
States, these reports were met with skepticism on the part of
some plant pathologists. In 1970, however, the blight, appar-
ently a new mutant strain, spread over the entire eastern half
of the United States. It caused a reduction in corn yields for the
country as a whole of about 13 percent. Many fields in the
southern states, where the infection occurred early in the season,
suffered losses of 50 percent or more. Single and three-way
crosses suffered greater damage from the blight than double
crosses, as Jones some twenty years earlier had warned that they
would when faced with a new hazard. The use of fertility-
restoring genes proved not to be a factor in the susceptibility of
corn hybrids to the blight. As soon as other types of cytoplasmic
male sterility can be introduced into commercial hybrids, it
is probable that the method will again be commonly employed.
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DONALD FORSHA JONES
141
In the meantime the blight, despite its destructive effects
on the 1970 corn crop, has had one beneficial result. It has
focused new attention on the importance of genetic diversity
in the world's major food crops. The new dwarf varieties of
wheat and rice that are revolutionizing the agriculture of
underdeveloped countries have, in the case of each of these
crops, dwarfing genes in common, although they are diverse in
their remaining heredity.
... . .
These common genetic loci may,
like the universally used cytoplasmic male sterility in corn,
become susceptible to new mutant pathogens with disastrous
results in countries and regions already overpopulated. Recog-
nizing this danger, the National Academy of Sciences in 1970
came forward with a proposal to study not only the problem
of the corn blight, but also the genetic vulnerability of major
food crops in general. The study was financed in large part
by Research Corporation, to which Dr. Jones assigned his patent
ort the use of fertility-restoring genes, and with particular ap-
propriateness by the Donald F. Jones Fund, which represents
the corporation's share of the net proceeds from the Jones
patent. The results of this study, published in 1972 in a volume
entitled Genetic Vulnerability of Major Crops, emphasized the
fact that "most major crops are impressively uniform and im-
pressively vulnerable." Thus Donald F. Jones has left a heritage
that continues to affect in tangible ways the course of applied
genetics to which, while living, he made such notable con-
. .
tr1 buttons.
Jones's conviction that selection in self-fertilized lines is one
of the most effective plant breeding methods that can be
practiced in cross-pollinated plants, and his interest in exploit-
ing heterosis was not confined to corn. During the period
from 1921 to 1926, when I served as his assistant, he under-
took breeding programs in numerous field and horticultural
crops, including such diverse crops as alfalfa and asparagus, and
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B I O G R A P H I C A L M E M O I R S
strawberries and squashes. Later, when it became apparent that
the use of cytoplasmic sterility in corn was becoming a re-
sounding success, he encouraged breeders of other field crops to
search for forms of cytoplasmic male sterility in the species to
which they were giving attention. The success of hybrid corn
also led to the exploitation of hybrid vigor in animals, especially
chickens, pigs, and cattle.
Jones's research was by no means confined to practical plant
breeding. His interest in theoretical genetics was as keen as
his concern with crop improvement. I have mentioned his
theory of heterosis. For a number of years he studied and wrote
extensively on somatic segregation, especially in its relation
to various kinds of atypical growths. Another subject to which
he gave attention was sex differentiation in maize and other
plants. In this connection he succeeded in converting maize—
which is normally monoecious, having both sexes on the same
plant, into a dioecious form having the two sexes on separate
plants. From this he concluded that monoecism may be an
intermediate step between perfect flowers and dioecism.
Donald Forsha Jones was born near Hutchinson, Kansas,
on April 16, 1890. He was the second of four children of Oliver
Winslow Jones and Minnie Wilcox Bush Jones. Both parents
were descendants of New England families. The Winslow in
Jones's father's name traces back to Kenelm Winslow, who came
to Plymouth, Massachusetts, in 1629. His maternal ancestors
were among the early settlers who founded Hartford, Con-
necticut. Jones, in one biographical sketch, has been aptly
called "a Yankee from Kansas."
At the time of their marriage both parents were school-
teachers. The family subsequently moved to Mulvane, Kansas,
and later to Wichita, where the father continued as a teacher
and a school principal. Their home, a small farm on the out-
skirts of Wichita, had ample space for gardening and for a
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DONALD FORSHA JONES
143
horse, a cow, a pig, and a flock of chickens. Donald and his
older brother, Kenneth, later a professor at Northwestern Uni-
versity Medical School, had a newspaper delivery route for
several years that required their getting up before daylight,
hitching the horse to the buggy, and completing their deliveries
in time to have breakfast at home. The family garden became
Donald's special responsibility, and he tried to grow some of
virtually everything. His boyhood interest in gardening was to
persist throughout his life. He was an ardent and talented
gardener. Even after a day's work in the experimental corn field
during the summer months he was not too tired to spend the
evening in his own garden. His gardening included a wide
variety of vegetables, small fruits, and ornamentals, all of which
responded to his gardening skill. Many of the popular articles
on horticulture that he wrote were based on his own firsthand
. .
gardening experience.
After completing his secondary school education, Jones at-
tended Kansas State Agricultural College, where he majored
in horticulture. ~~ ' ' ~ ~
His college years apparently were neither
particularly inspiring nor enjoyable, and he was not regarded
as an outstanding student. However, one of the honors that
he cherished most in his later years was an honorary degree,
Doctor of Science, awarded to him in 1947 by his alma mater.
Graduating from college in 1911, he took a position at the
Arizona Experiment Station, where one of his principal job.s
was pollinating alfalfa flowers in a breeding program. He once
spoke disparagingly of his role in this position as "taking the
place of the bumble bee." While there he read the recently
published bulletin by East and Hayes, "Heterozygosis in Evolu-
tion and Plant Breeding." Having already observed the effects
of inbreeding on alfalfa, he found this publication of unusual
interest, and he wrote to East inquiring about the possibility
of doing graduate work at Harvard under East's direction. In
the meantime, however, he took a position at Syracuse Univer-
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BIOGRAPHICAL MEMOIRS
sity, where he spent the year 1913-1914 teaching and working
toward the master's degree. He enrolled as a graduate student
at Harvard in the fall of 1914. In February 1915 he moved to
New Haven to take charge under East's supervision of the plant-
breeding program, which had been initiated by East in 1905
at the Connecticut station. For the next several years he di-
vided his time between New Haven and Cambridge
Jones remained at the Connecticut Agricultural Experiment
Station for the rest of his professional life, and this continuity
of effort was undoubtedly one factor in his lifelong productive-
ness in research and publication. His technical scientific nuh-
- rib
lications cover a period from 1915 to ,964 and include ap-
proximately one hundred titles. In addition he wrote many
popular and semipopular articles covering a wide variety of
subjects for the farm and garden press. When I joined him as
a graduate assistant in 1921, he advised me also to write for
these media, partly in the interest of informing the general
public on new developments in agricultural science and partly
as practical experience in writing, which for me it proved
indeed to be.
An inveterate reader as well as a prolific writer, Jones,
during the winter months, usually spent his mornings writing
and reading scientific literature. His evenings at home, except
during the summer when he worked in his garden until dark-
ness, were most commonly spent in reading. Among his favorite
subjects were biography and history. Even his luncheon periods
were devoted partly to reading. Alternating in reading aloud
to each other, he and I, in the years between 1921 and 1926,
went through various works, including Darwin's Voyage of the
Beagle.
One of the most unusual aspects of Jones's career was his
close and congenial collaboration for a number of years with
Edward M. East, his graduate-school mentor. The two men
were quite different in temperament.
Henry Wallace once
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DONALD FORSHA JONES
147
In Joneses later years, in addition to his duties at the Con-
necticut Station, he was a lecturer in genetics at Yale University
and at the University of Connecticut. He was Sprague Me-
morial Lecturer at Michigan State College in 1935; research
fellow at California Institute of Technology in 1935-1936 and
again in 1946-1947; and visiting professor at the University of
Washington in 1953.
{ones was the recipient of many honors. He was elected to
the American Academy of Arts and Sciences in 1934, to the
National Academy of Sciences in 1939, and to honorary mem-
bership in the Societa Italiana Genetica Agraria in 1955. I
have already mentioned the honorary degree that he received
from his alma mater in 1947. The accompanying citation
reads:
"As a major contributor to the development of hybrid
corn . . . he has conferred immense benefits upon a hungry
world. As research scientist, teacher, and leader in scientific
societies, he has inspired his colleagues and students by . . . his
extraordinary achievements . . ."
The New England Council and the Governors of New Eng-
land made him a charter member of the `'Fellowship of Agri-
cultural Adventurers," saying that,
"with an imagination which delved below and soared above
the findings of his predecessors, he translated the learning of the
laboratory into the fruitfulness of the field."
He was awarded gold medals by the Massachusetts Horti-
cultural Society and the American Farm Bureau. In addition
he was given awards by the Connecticut State Confederation of
Women's Clubs, the American Seed Trade Association, the Con-
necticut State Grange, the New York Farmers' Club, and the
Botanical Society of America.
Dr. Jones died at his home in Hamden, Connecticut, on
June 19, 1963. He is survived by his wife, Eleanor March
Jones, a son, Loring M. Jones, and a daughter, Mrs. Margaret
Owen.
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148
KEY TO ABBREVIATIONS
BIOGRAPHICAL MEMOIRS
BIBLIOGRAPHY
Prepared by Carolyn A. Sleekly
Am. .T- Bot.—American journal of Botany
Am. Nat. _ American Naturalist
Biol. Bull. _ Biological Bulletin
Bot. Gaz. Botanical Gazette
Conn. Agric. Exp. Stn. Bull. Connecticut Agricultural Experiment Sta-
tion Bulletin
Conn. Agric. Exp. Stn. Circ. Connecticut Agricultural Experiment Sta-
tion Circular
Conn. Agric. Exp. Stn. Rep. = Connecticut Agricultural Experiment Sta-
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I. Am. Soc. Agron. Journal of the American Society of Agronomy
I. Hered. Journal of Heredity
Proc. Natl. Acad. Sci. Proceedings of the National Academy of Sciences
Proc. Int. Congr. Genet. Proceedings of the International
Congress of Genetics
Sci. Mon. Scientific Monthly
Seventh Int. Manage. Congr. = Seventh International Management Con-
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1915
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DONALD FORSHA JONES
1917
149
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i
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DONALD FORSHA JONES
151
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BIOGRAPHICAL MEMOIRS
1931
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With W. R. Singleton and L. C. Curtis. The correlation between
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With E. Huntington. The adaptation of corn to climate. J. Am.
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DONALD FORSHA JONES
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A multiple mosaic in maize. .T- Hered., 26: 191-92.
With E. Huntington. Further comments on adaptation of corn to
climate. I. Am. Soc. Agron., 27:682-83.
1936
Segregation of color and growth-regulating genes in somatic tissue
of maize. Proc. Natl. Acad. Sci., 22:163-66.
Tumors in Drosophila melanogaster resulting from somatic segre-
gation. Science, 84: 135.
Mutation rate in somatic cells of maize.
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Atypical growth. Am. Nat., 70:86-92.
'with W. R. Singleton. Sweet corn inbreds.
Circ. 112, pp. 49-58.
1937
Somatic segregation in relation to atypical growth.
the American Philosophical Society, 77:411-16.
Somatic segregation and its relation to atypical growth. Genetics,
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The life and work of Luther Burbank. Spragg Memorial Lectures
on Plant Breeding, Michigan State College, East Lansing, Michi-
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Proceedings of
O
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1938
Crops and forests: their production, protection and use. Seventh
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Utilization of plant resources. Seventh Int. Manage. Congr., Wash-
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Translocation in relation to mosaic formation in maize. Proc. Natl.
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Nuclear control of cell activity. Science, 88:400-401.
1939
Sex intergrades in dioecious maize. Am. ]. Bot., 26:412-15.
Continued inbreeding in maize. Genetics, 24:462-73; Russian
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OCR for page 154
154
BIOGRAPHICAL MEMOIRS
With W. R. Singleton. The Shelton and Hebron
Conn. Agric. Exp. Stn. Chic. 137. 4 pp.
With W. R. Singleton. Early sweet corn hybrids—Spancross, Mar-
cross, and Carmelcross. Conn. Agric. Exp. Stn. Circ. 138, pp.
5-11.
1940
Nuclear changes affecting growth. Am. J. Bot., 27:149-55.
With W. R. Singleton. The improvement of naturally cross-polli-
nated plants by selection in self-fertilized lines. III. Investi~a-
tions with vegetatively propagated fruits.
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1941
a
Conn. Agric. Exp.
Somatic segregation. Botanical Review, 7:291-307.
An investigation of growth in plants. Science, 93:40.
Natural and induced changes in chromosome structure in maize
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With W. R. Singleton. Sweet corn hybrids Lexington, Lincoln
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1942
Chromosome degeneration in relation to growth and hybrid vigor.
Proc. Natl. Acad. Sci., 28: 38~4.
1943
Review of Producer of Better Fruits and Fairer Flowers (Luther
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Chromosome degeneration.
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1944
Equilibrium.in genie materials. Proc. Natl. Acad. Sci., 30:82-87.
Growth changes in maize endosperm associated with the relocation
of chromosome parts. Genetics, 29:420-27.
1945
Edward Murray East (1879-1938~. In: National Academy of Sci-
ences, Biographical Memoirs, 23:217~2.
Heterosis resulting from degenerative changes.
Genetics, 30:527~2.
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DONALD FORSHA JONES
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The importance of degenerative changes in living organisms. Sci-
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With W. R. Singleton and O. E. Nelson, fir. The improvement of
naturally cross-pollinated plants by selection in self-fertilized
lines. IV. Combining ability of successive generations of inbred
sweet corn. Conn. Agric. Exp. Stn. Bull. 490, pp. 453-98.
1947
Effect of temperature on the growth and sterility of maize. Science,
105:390-91.
The nature of gene action as shown by cell-limited and cell-dif-
fusible gene products. Proc. Natl. Acad. Sci., 33:363-65.
1948
Induced and naturally occurring mutations. In: Proceedings, Au-
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With W. R. Singleton and H. L. Everett. Sweet-dent silage.
Conn. Agric. Exp. Stn. Circ. 165. 14 pp.
1949
With H. L. Everett. Hybrid field corn.
Bull. 532. 39 pp.
Conn. Agric. Exp. Stn.
1950
The interrelation of plasmagenes and chromogenes in pollen pro-
duction in maize. Genetics, 35:507-12.
1951
The cytoplasmic separation of species. Proc. Natl. Acad. Sci., 37:
408-10.
With P. C. Mangelsdorf. The production of hybrid corn seed with-
out detasseling. Conn. Agric. Exp. Stn. Bull. 550. 21 pp.
The induction of cytoplasmic pollen sterility and the restoration of
fertility in maize. Genetics, 36: 557(A).
1952
Plasmagenes and chromogenes in heterosis. In: Heterosis, ed. by
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156
BIOGRAPHICAL MEMOIRS
1954
Gene and cytoplasm interaction in species separation. Proc. 9th
Int. Congr. Genet., pp. 1225-27. Caryologia. Vol. VI, Suppl.
1956
Genic and cytoplasmic control of pollen abortion in maize. Tn
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1957
Gene action in heterosis. Genetics, 42:93-103.
Edith H. T. Stinson, [r. and U. Khoo. Transmissible variations in
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With H. T. Stinson, [r. and U. Khoo. Pollen restoring genes.
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1958
Heterosis and homeostasis in evolution and in applied genetics.
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1959
Basic research in plant and animal improvement.
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Proc. 10th Int.
1960
The genotype as the sum of plasmatype and chromotype. Am. Nat.,
94: 181-83.
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Representative terms from entire chapter:
inbred strains
