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THEOPHILUS SHICKEL PAINTER
August 22, October 5, l 969
BY BENTLEY GLASS
IN T H E S T E ~ ~ A R D A Y S O F Drosophila genetics during the
1920s and 1930s, only two principal centers of such re-
search existed in the United States. The California Institute
of Technology attracted Thomas Hunt Morgan from Colum-
bia University in 1929, and he brought with him his two stu-
dents, Alfred H. Sturtevant and Calvin B. Bridges, who a
decade earlier had contributed to the establishment of the
chromosome theory of heredity. The CalTech group also in-
cluded Theoclosius Dobzhansky, lack Schultz, and a constel-
lation of notable visiting fellows, present for a year or two,
such as George Beadle and Curt Stern.
During the same period a second stellar group formed at
the University of Texas in Austin. H. I. Muller, one of the
original trio of Morgan's graduate students, tract created a
great stir in genetics with his 1927 discovery that X-rays will
induce mutations at frequencies hundreds, even thousands,
of times higher than rates of spontaneous mutation. A gen-
erous grant from the Rockefeller Foundation macle it pos-
sible for Muller, joined by I. T. Patterson ant! T. S. Painter of
the Department of Zoology at Austin, to establish a cytoge-
netical program for exploiting the new discovery. Graduate
students were recruiter! and given fellowships, the earliest of
which went to C. P. Oliver, Wilson S. Stone, and the writer of
309
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310
BIOGRAPHICAL MEMOIRS
this memoir. Muller soon found that X-rays produce chro-
mosomal breaks and rearrangements in addition to gene mu-
tations, Oliver worker! out the relation of point mutations to
radiation dosage, Painter collaborates! with Muller in analyz-
ing chromosomal rearrangements, and Patterson explored
an exciting new field mosaic types of mutation produced
by X-rays. Bursts of exciting new findings made the rivalry
with CalTech as hectic as a close basketball game, ant! Painter
was a central figure in all of it.
EARLY LIFE AND EDUCATION
T. S. Painter was born in Salem, Virginia, the son of
Franklin V. N. Painter ant! Laura T. Shicke! Painter. T. S.'s
father was an esteemed educator, a professor of modern lan-
guages and English literature at Roanoke College. Both par-
ents were very religious, and their son was brought up in an
atmosphere of culture and religious faith that marked him
deeply. His midclle name was that of his mother's family; his
given name reflects his parents' Christian orientation. As a
boy, T. S. was sickly and obtained most of his elementary ant}
secondary education by home tutoring. He entered Roanoke
College in 1904 anc! graduated with a B.A. degree in 1908.
The college was a small one and clid not provide a diversity
of scientific courses. Painter was attracted to chemistry ant!
physics but had no opportunity to acquaint himself with biol-
ogy.
Having received a scholarship in chemistry, he entered
Yale University as a graduate student in 1908. Here he met
Professor L. L. Woodruff of the Biology Department and
asker! to be permitted to sit in a corner of the laboratory anal
look at objects under a microscope, which he hac! never had
an opportunity to use before. Professor L. L. Woodruff as-
signec! Painter a microscope and proviclec! him with a hay
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THEOPHILUS SHICKEL PAINTER 311
infusion full of active bacteria, protozoans, and algae. Painter
was fascinates! and soon decided that he wanted to change
his field from chemistry to biology.
He receiver! an M.A. degree in 1909 and a Ph.D. in 1913,
uncler the direction of the fames! authority on spiders Alex-
ancier Petrunkevitch. Painter learned the techniques of cy-
tology as practicer] at that time ant! for his thesis explored
the process of spermatogenesis in a species of spicier. His first
scientific publication (1913, I) was a paper on dimorphism in
males of the jumping spider, Maevia vittata. His second
(1914,1) was his thesis research.
Painter then went to Europe for a year of postdoctoral
study, partly in the laboratory of Theodor Boveri, in
Wurzburg, ant! partly at the famed Marine Zoological Station
at Naples. At that time Boveri was among the foremost cy-
tologists in the world. More than a clecacle earlier he had
established, in studies of the fertilization and development of
Ascaris eggs, that each chromosome controls development in-
clividually. Chromosomes, furthermore—although they
seem to disappear after the close of each mitotic cell divi-
sion—have a persistent continuity and reappear in the next
mitosis in the same place they occupier! before their disap-
pearance. Most surprisingly, they continue to bear whatever
aberrant distinctions they might previously have acquired by
accident. Boveri was a stout supporter of the chromosome
theory of heredity which he had enunciates! inclependently
of W. S. Sutton, a student of E. B. Wilson at Columbia. Later,
when ~ was taking a graduate course with Painter at Austin,
it was a matter of astonishment to me that ~ never heard him
reminisce about those exciting times or make any reference
to Boveri or to what he learned from him.
The experience at Naples, with its marvels of marine life
for a cytologist to explore, seemed to affect Painter more. His
next publications (lealt with problems of the forces involved
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BIOGRAPHICAL MEMOIRS
in the cleavage of the fertilized egg into a multiplicity of cells
by means of repeated mitotic cell divisions.
Back in the Uniter} States from a war-torn Europe, Painter
received an appointment as an instructor in zoology at Yale
for two years. He was also asked to teach marine invertebrate
zoology at the Woods Hole Laboratory in the summers of
1914 and 1915. There he met two persons who were to be
exceedingly important in his life. The first, Mary Anna
Thomas, was a young student in his course who woulc! later
become his devoted wife. The second, John Thomas Patter-
son, was the young head of the Zoology Department at the
University of Texas in Austin. Patterson offered Painter the
academic post that brought him to the institution where he
would spend the remainder of his life. In his Biographical
Memoir of I. T. Patterson (1965,1), Painter told of the warm
and friendly way in which the two first met while playing
baseball with other teachers and researchers at Woods Hole.
Painter's research at this period greatly resembled the
type of experimentation on developing invertebrate embryos
favored by E. B. Wilson and E. G. Conklin. He first studied
the effects of carbon dioxicle on the developing eggs of As-
caris, the material for which hac} been obtained at Wurzburg.
His next study also took its origin from work begun in Eu-
rope, this time at Naples, where Painter had discoverer! spiral
asters in developing eggs of sea urchins and become curious
about their participation in the process of embryonic cleav-
age. He investigates! the occurrence of monaster eggs, the
light they threw on cell mechanics during division, and the
influence of narcotics on cell division. Painter demonstrates!
that eggs may divide in the absence of asters, that a factor
clerived from the nucleus is requires! for division, and that
the asters presumably play a regulatory role in the distribu-
tion of the nuclear factor.
In May 1916, Painter enlisted in the National Guarc! at
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THEOPHIEUS SHICKEE PAINTER 313
New Haven ant! became a sergeant of the Headquarters
Company of the Tenth Regiment of Fielc! Artillery. Dis-
chargec! in September 1916, he married Anna Thomas on
December 19, 1917. Their children two boys and two
girls and, eventually, their grandchilclren made a warm,
closely knit family.
With the advent of World War ~ in 19 ~ 7, Painter was com-
missioned a first lieutenant of the U.S. Army Signal Corps
and was sent to Toronto's Imperial Flying School to find out
what measures were needed! to establish a ground! school of
aviation in Austin. After the school was established, he served
as a member of its academic board and was promoted in 1918
to captain in the U.S. Army Air Service. In April 1919 he
retiree] as a captain of the Reserve Corps.
Though Painter went to Austin in 1916 as an adjunct pro-
fessor of zoology, military service interrupted his research for
several years, and he was not promoter! to associate professor
until 1921. Four years later, in 1925, he was appointed full
professor with membership in the graduate faculty.
Painter was a man of broad interests and cheerful dispo-
sition. He often visited his students in the laboratory to
exchange ideas, giving them encouragement as well as di-
rection. He taught undergraduate courses in aciclition to
graduate cytology, and for many years- a popular
premedical course in comparative anatomy. He played tennis
and golf ant! lover! swimming, fishing, and crabbing. He was
also an inveterate hunter, liking nothing more than to take
down his rifle to hunt deer or antelope. He was a fine gar-
clener, ant! his flower displays were a marvel to all visitors.
He particularly enjoyed hybridizing irises to produce new
patterns of remarkable color. He was an expert with tools
and macle furniture for his home. In later years he turner!
to jeweIry-making and again cleveloped great skill at produc-
ing objects that reflected his fine taste. He took a strong part
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BIOGRAPHICAL MEMOIRS
in his church's activities and in various clubs. In many ways
the antithesis of the stereotypical Texan, he was both re-
served and self-controlled.
CHROMOSOME CYTOLOGY AND SEX CHROMOSOMES
Back at the University of Texas after his military service,
Painter resumed his cytological studies of spermatogenesis in
a common small lizard, Anolis caroZinensas. But he quickly
turned to a new problem: the number of mammalian chro-
mosomes and their morphology, with particular emphasis on
the nature of sex determination.
In the zoology laboratories of the Department, embryol-
ogist Car} G. Hartmann was engaged in studying the repro-
cluction of the opossum. "There was 'possum meat all over
the lab," Painter remarked, a fine opportunity for him to
switch from spillers, marine organisms, and lizards to the
enticing field of mammalian cytology.
Almost nothing was known about mammalian chromo-
somes at the time, although it was supposed that mammals
must have sex chromosomes corresponding to those of in-
sects and that an XXffemale)-XY(rnale) distinction would ex-
ist. It proved quite easy, in fact, to find the sex chromosomes
of the opossum, for they were the smallest pair of chromo-
somes in the cell, and during spermatogenesis they always lay
in the center of a ring of the other, larger chromosomes clur-
ing the metaphase of mitosis. In those days all tissues used
for cytological examination were successively fixed, embed-
ded in paraffin, sectioned, and stained. It was of prime im-
portance to get the tissues fresh from dissection into the fix-
ing fluicI. Painter invented a sort of multiblaclec} knife by
mounting a number of safety razor blades in parallel, close
together, which he used to cut up the spermatogenic tubules
of the testis immediately after the organ was excised.
Painter demonstrated that the male opossum's sex is de-
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THEOPHILUS SHICKEL PAINTER 315
terminec! by a tiny Y-chromosome in place of one of the fe-
male's larger X-chromosomes. He shower! that in meiosis of
the male's spermatocytes prior to formation of spermatozoa,
the X and Y chromosomes pair and then segregate, so that
each male reproductive cell carries either an X- or a Y-
chromosome, but not both. As in insects, then, if all egg cells
carry a single X-chromosome anct if fertilization by the two
sorts of spermatozoa is random, the X-bearing sperm would
produce female offspring; the Y-bearing sperm wouIcl pro-
duce males.
Having thus shown that sex determination in a marsupial
mammal corresponds to the process already known from in-
vertebrates, Painter set his sights on placental, or eutherian,
mammals, and- through a fortunate circumstance was
able to obtain fresh human testicular tissue. One of his for-
mer premedical students was practicing medicine in a state
mental institution in Austin where, "for therapeutic reasons,"
Painter wrote. "they occasionalIv castrated male individuals."
Painter's former student made it possible for him to obtain
and preserve, "within thirty seconds or less after the blood
supply was cut oh, a human testis" (197l,1~. We students in
the Austin laboratory speculated widely that such tissue was
also obtained from criminals executed at the nearby Hunts-
ville prison, but this was probably just idle gossip. Painter
himself never confirmed such a source.
Painter's first work on human chromosomes, therefore,
preceded his study of primates, though their order of pub-
lication was reversed. A year before he publisher! his fuller
account of human spermatogenesis and human sex chro-
mosomes (1923,1), a short announcement on the sex chro-
mosomes of"the monkey" appeared in Science.
To solve the enigma of sex determination in humans,
Painter turned to two species of monkey the New WorIc}
Brown Cebus and the OIct WorIc3 Rhesus (Rhesus macacus). As
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316
BIOGRAPHICAL MEMOIRS
he pointed out in this pioneering work (1924,3), it was highly
desirable and perhaps necessary to establish four matters for
each species examined: (~) the morphology of the diploid
chromosome complex and the chromosome number of the
male; (2) the haploid number revealed in the second sper-
matocytes; (3) the morphology and behavior of the sex chro-
mosomes (X and Y) during meiosis; and (4) the morphology
and chromosome number of the female complex. Cross-
checks among these observations should bar all possibility of
error, even though many species of mammals including the
primates Painter was investigating" have many more and
much smaller chromosomes in their karyotypes than do
opossums or the insect species in which the chromosomal
determination of sex was first established. (A "karyotype" is
the term used to designate the entire group of chromosomes
characteristic of a cell of a particular species. This conic! be a
diploicl cell with two complete sets of chromosomes or, more
frequently, the chromosome complement of a haploic! cell
with a single set of chromosomes one of each distinctive
kind characterizing the species.)
Painter's demonstration of the X-Y type of sex determi-
nation in these mammals ant! in the human species was com-
pelling. His drawings of the larger X-chromosome and the
much smaller Y-chromosome, connected to each other by a
thin strand while segregating in the first prophase of meiosis,
left no doubt.
The number of chromosomes was less certain. Some hu-
man cells seemed! to show a count of forty-eight chromo-
somes in the cliploic! primary spermatocyte, others only forty-
six. Previous investigators of human chromosome number
also varied in their counts, though most settled for forty-
eight.
Painter himself took the evidence of his "best cell" and
reported the number as forty-eight, confirming an error that
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THEOPHIEUS SHICKEE PAINTER 317
would be perpetuated in dozens of textbooks (including one
of my own) until a new set of techniques for counting chro-
mosomes was introducer! in the mid-1950s. In 1956, using
new stains (such as acetocarmine and Feulgen's stain specific
for DNA) and soft somatic tissues (especially embryonic tis-
sues) that could be smeared; using coIchicine to halt clividing
cells in metaphase and hence greatly increase the number of
such cells observable; and using hypotonic salt solutions to
spread the chromosomes of dividing cells apart to eliminate
their clumping into uncountable masses, I. H. Tjio and A.
Levan made a definitive determination that the human dip-
loid chromosome number is forty-six, i.e., twenty-three pairs
of homologous chromosomes in human ctiploid cells.
Painter experienced deep chagrin over this error in what
had long been regarded as a primary discovery for which he
was known ant} universally cited. Yet—given the source of
his material and the procedures available to him in the early
1920s he may not have been entirely wrong. Indivicluals
with mental disorders are not prime material for cletermin-
ing normal chromosome number and morphology, for they
sometimes have forty-seven, forty-eight, or even more chro-
mosomes ant} exhibit more frequently than normal persons
transIocations and deletions of chromosomes that would ap-
pear to alter their number.
Recently T. C. Hsu, a well-known cytogeneticist, reexam-
ined some of the original preparations on which Painter
based his erroneous chromosome count and found that the
chromosomes were so badly clumped and cut into segments
by the microtome knife, it was a marvel Painter was able to
find any cells at all that seemed to give a clear chromosome
count. Given that human chromosomes are exceedingly
small, that the dyes used in the 1920s darkly stained other
matter in addition to chromosomes, ant! that microtome
slices rarely proclucecl whole, undamaged cells for examina-
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BIOGRAPHICAL MEMOIRS
tion, Painter's error was wholly natural and forgivable. In any
case, it in no way diminishes the importance of his discovery
of the XX-XY mechanism for determining sex in mammals
(inclucling humans), a significant contribution to science.
Painter subsequently examined and recorded the chro-
mosome number of the horse (probably 60; XX-XY sex cle-
termination), the bat Nyctinomous mexicanus (2 N = 48), the Eu-
ropean hedgehog (2 N = 48), the armadillo (2 N = 60), the
rabbit (2 N = 44), and the dog (2 N prob. 521. Additional
marsupials examined included besides the opossum
(2 N = 22) Phascolarctus (2 N = 16), Sarcophilus (2 N = 14), Das-
yurus (2 N = 14), and the kangaroo Macropus (2 N = 12~.
Painter identified an XY pair of sex chromosomes in all of
these marsupial and placental mammals except the hedge-
hog, armadillo, ant! clog species he did not investigate ex-
tensively enough to judge- though an XY male type was not
exclucled in them either.
In summary, Painter shower! that marsupial mammals in
general have a lower chromosome number than placental
mammals; that all, or almost all, placentals (including hu-
mans) have a high chromosome number ranging from forty-
four to sixty; ant! that all of them have, or probably have, an
XX-XY type of sex determination depending upon a partic-
ular pair of sex chromosomes in which the Y-chromosome
(carried by the male) is far smaller in size than the X-
chromosome.
If these studies placed Painter in the first rank of cyto-
geneticists, the focus of his next research project established
him firmly in the forefront of classical genetics. One of
Painter's students, E. K. Cox, tract determined that the chro-
mosome number of the common house mouse, Mus musculus,
is forty. Yet W. H. Gates reported that a Japanese waltzing
mouse found in the F! offspring of a cross between normal
(dominant) ant! Japanese waltzer (recessive) parents seemed
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THEOPHILUS SHICKEL PAINTER 327
of science necessarily implies that during a lapse of even two
or three years from the laboratory, fundamental changes in
understanding will have occurred to such an extent that the
returneci scientist's erase of current knowledge and mastery
~ ~ 1
of available techniques are outmocled.
So it was with Painter, but his determination was indom-
itable. His colleagues testify that he spent more time in the
library reacting current periodicals and books than did any
graduate student. He also asked to be reassigned to the teach-
ing of cell biology to untlergracluates anti cytology to grad-
uate students, and thus addled to his burden all the reviewing
and relearning required for teaching. As the Memorial Res-
olution prepared by his fellow faculty members records, he
was successful:
He developed a good knowledge of modern cellular molecular biology.
Often he noticed that a researcher's data could be used to answer in part
some classical biological problem, although the author had not mentioned
that possibility. The interpretations were too narrow in coverage. As a
consequence, Dr. Painter decided to teach his students the recent, chemi-
cally-oriented discoveries and to make certain that they had a broader basic
training in biology so that they could understand the biological implica-
tions of the discoveries. To Dr. Painter, a narrow channel of research may
find answers for one small field of interest, but it will not serve the purpose
of biology unless it has some major impact upon a basic biological problem.
One can verify his concern with the broader implications
by glancing at eleven scientific papers written by Painter be-
tween 1955 and 1969. They seem to follow naturally from
the earlier work on the salivary chromosomes of dipterans
ant! the endomitosis in the nurse cells of the ovary. But they
all probe the greater question of how it is that the hereditary
materials passer! down from one generation to another in the
course of reproduction are converted into a multiplicity of
end products in (different tissues.
Working with J. J. Biesele again and with the advantage
of electron microscopy—Painter was able to show how the
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328
BIOGRAPHICAL MEMOIRS
precursors needed for the secretion of royal jelly (the only
food consumed by the queen bee) are produced in the honey-
bee in special gland cells of young worker bees. Producing as
many as 1000 eggs a day, the queen bee requires a consid-
erable supply of both proteins and DNA, which is supplied
by the royal jelly. When workers feed heavily on bee bread,
their gland cells develop and produce the royal jelly.
According to George E. Palade, Keith Porter, and others,
royal jelly gland cells in the young worker bees produce the
proteins by means of an extensively developed endoplasmic
reticulum. Painter and Biesele searched for the origin of this
cellular structure of endoplasmic tubules that apparently de-
rive from outpockets of the nuclear membrane of the cell as
the gland cell undergoes endomitosis. As this process enters
a stage comparable to the prophase of ordinary mitosis, the
numerous nuclei in the gland cell fragment and a myriad of
ribosome-like bodies pass out through nuclear pores to be-
come the polyribosomes attached to the walls of the endo-
plasmic tubules. This process clearly shows how an ovum be-
comes enriched with protein and nucleotides.
In his final paper, Painter advised vouna researchers from
· -
. AS own experience:
J (J
"I get the impression that young people [today] master some sophisticated
technique such as labeling cellular structures with radioactive isotopes fol-
lowed by autoradiography, DNA and RNA hybridization, ultracentrifu-
gation in gradients and all the rest and then look around to see how they
can use their acquired skills! From my experience I think you should first
select and define some broad biological problems, select a suitable material
upon which to work and use any available techniques for the solution of
your problem. The most important thing is for you to have a biological
and not a test tube approach." (1971,1)
How well his own research exemplifies! that ability to identify
the problem, find the right material, and develop the neces-
sary techniques!
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THEOPHIEUS SHICKEE PAINTER 329
Although research always stool! foremost in his heart,
Painter found time and energy for many other activities. He
served on the University of Texas Premedical, Predental, ant!
Library committees. He frequently attencled the meetings of
scientific societies and, in adclition to serving on other com-
mittees of the American Philosophical Society, was a member
of its Council from 1965 to 1967. He server! for six years on
the Council of the National Academy of Sciences ant! six
more on its Finance Committee. He was a member of the
American Society of Zoologists, the Genetics Society of
America, the Association of American Anatomists, the Amer-
ican Society of Naturalists, and the Societa Italiana di Biolo-
gia Sperimentale.
He was a member of the Boy Scouts of America Commit-
tee (1935-40), an advisor to the Dental Research Council
(1949-52), and advisor on research to the American Cancer
Society. He server! on the Commission on Colleges and Uni-
versities of the Southern Association and was its chairman
for three years; the Southern Regional Education Board; the
National Committee on Accreditation; and the Board of the
Institute of Nuclear Studies at Oak Ridge. He was a National
Lecturer for Sigma Xi in 1936-37. Locally, he was a member
of the Rotary Club, Town and Gown, and the English Speak-
ing Union.
He was electect to the Hall of Fame for Famous Ameri-
cans, server! as president of the American Society of Zoolo-
gists in 1940, ant! received the first M. D. Anderson Award
for Scientific Creativity and Teaching from the M. D. Ander-
son Hospital and Tumor Institute in 1969. Perhaps what he
regarded most highly among his honors was his elevation to
the rank of distinguishes! professor of the University of Texas
in 1939.
It was characteristic of him that he died as he had lived-
suddenly, on his return home to Fort Stockton, Texas, from
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BIOGRAPHICAL MEMOIRS
a hunting trip, in his eighty-first year and as active as ever.
Two papers- "The Origin of the Nucleic Acid Bases Found
in the Royal Jelly of the Honeybee" (1969,1) and "Chromo-
somes and Genes Viewed from a Perspective of Fifty Years"
~197 I, I) appeared posthumously.
THE AUTHOR OF THIS MEMOIR iS deeply indebted to the Univer-
sity of Texas Faculty Committee that prepared the Memorial Min-
ute on T. S. Painter that is quoted above. Members of this Com-
mittee were C. P. Oliver, chairman; I. I. Biesele; and R. P. Wagner.
I would also like to acknowledge with deep gratitude the receipt of
various documents, both published and unpublished, from Mrs.
T. S. Painter. Without access to them there would have been seri-
ous gaps in the account, especially in respect to T. S. Painter's ad-
. . .
ministrative career.
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THEOPHILUS SHICKEL PAINTER 331
SELECTED BIBLIOGRAPHY
1913
On the dimorphism of the males of Maevia vittata, a jumping spi-
der. Zool. Jahrb. Abt. Syst. Oekol. Geogr. Tiere, 35:625-35.
1914
Spermatogenesis in spiders. I. Zool. tahrb. Abt. Anat. Ontog.
Tiere, 38:509-76.
The effect of carbon dioxide on the eggs of Ascaris. Proc. Soc. Exp.
Biol. Med., 11 :62-64.
1915
An experimental study in cleavage. I. Exp. Zool., 18:299-323.
The effects of carbon dioxide on the eggs of Ascaris. l. Exp. Zool.,
19:355-85.
1916
Some phases of cell mechanics. Anat. Rec., 10:232-33.
Contributions to the study of cell mechanics. I. Spiral asters. I. Exp.
Zool., 20:509-27.
1917
A wing mutation in Piophila casei. Am. Nat., 51:306-8.
1918
Contributions to the study of cell mechanics. II. Monaster eggs and
narcotized eggs. J. Exp. Zool., 24:445-97.
1919
The spermatogenesis of Anolis carolinensis. Anat. Rec., 17: 328-29.
1921
Studies in reptilian spermatogenesis. I. The spermatogenesis of
lizards. J. Exp. Zool., 34:281-327.
The Y-chromosome in mammals. Science, 503-4.
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332
BIOGRAPHICAL MEMOIRS
1922
Studies in mammalian spermatogenesis. I. The spermatogenesis of
the opossum (Didelphys virginiana). ]. Exp. Zool., 35: 13-38.
The sex chromosomes of the monkey. Science, 56:286-87.
1923
Studies in mammalian spermatogenesis. II. The spermatogenesis
of man. l. Exp. Zool., 37:291-336.
Further observations on the sex chromosomes of mammals. Sci-
ence, 58:247-48.
1924
A technique for the study of mammalian chromosomes. Anat. Rec.,
27:77-86.
Studies in mammalian spermatogenesis. III. The fate of the chro-
matin-nucleolus in the opossum. l. Exp. Zool., 39:197-227.
Studies in mammalian spermatogenesis. IV. The sex chromosomes
of monkeys. J. Exp. Zool., 39:433-62.
Studies in mammalian spermatogenesis. V. The chromosomes of
the horse. J. Exp. Zool., 39:229-47.
The sex chromosomes of man. Am. Nat., 58:506-24.
1925
Chromosome numbers in mammals. Science, 61:423-24.
A comparative study of the chromosomes of mammals. Am. Nat..
59:385-409.
The chromosomes of the rabbit. Anat. Rec., 31:304.
A comparative study of the chromosomes of the largest and the
smallest races of rabbits. Anat. Rec., 31:304.
A comparative study of mammalian chromosomes. Anat. Rec.,
31:305.
1926
The chromosomes of rodents. Science, 64:336.
Studies in mammalian spermatogenesis. VI. The chromosomes of
the rabbit. I. Morphol. Physiol., 43: 1-43.
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THEOPHILUS SHICKEL PAINTER 333
1927
The chromosome constitution of Gates' "non-disjunction" (v-o)
mice. Genetics, 12:379-92.
1928
A comparison of the chromosomes of the rat and mouse with ref-
erence to the question of chromosome homology in mammals.
Genetics, 13: 180-89.
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Representative terms from entire chapter:
sex chromosomes
