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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, furthermorealthough they seem to disappear after the close of each mitotic cell divi- sionhave 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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312 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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314 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. Yetgiven 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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318 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 microscopyPainter 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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330 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. The chromosome constitution of the Little and Bagg abnormal- eyed mice. Am. Nat., 62:284-86. Cell size and body size in rabbits. l. Exp. Zool., 50:441-53. 1929 With H. J. Muller. Parallel cytology and genetics of induced trans- locations and deletions in Drosophila. J. Hered., 20:287-98. With H. J. Muller. The cytological expression of changes in gene alignment produced by X-rays in Drosophila. Am. Nat., 63: 193-200. 1930 Recent work on human chromosomes. }. Hered., 21:61-64. Translocations, deletions, and breakage in Drosophila melanogaster. Anat. Rec., 47:392. 1931 With I. T. Patterson. A mottled-eyed Drosophila. Science, 73:530- 31. A cytological map of the X-chromosome of Drosophila melanogaster. Science, 73:647-48. (Also in: Anat. Rec., 51: 111.) 1932 With H. J. Muller. A cytological map of the X-chromosome of Dro- sophila. Proc. 6th Int. Congr. Genetics (Ithaca), 2:147-48. With H. I. Muller. The differentiation of the sex chromosomes of Drosophila into genetically active and inert regions. Z. Indukt. Abstamm. Vererbungsl., 62:316 - 65.

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334 BIOGRAPHICAL MEMOIRS 1933 A method for the qualitative analysis of the chromosomes of Dro- sophila melanogaster. Anat. Rec., 57 (Suppl.~:90. A new method for the study of chromosome rearrangements and the plotting of chromosome maps. Science, 78:585-86. 1934 A new method for the study of chromosome aberrations and the plotting of chromosome maps in Drosophila melanogaster. Genet- ics, 19:175-88. The morphology of the X-chromosome in salivary glands of Dro- sophila melanogaster and a new type of chromosome map for this element. Genetics, 19:448-69. A new type of cytological map of the X-chromosome in Drosophila melanogaster. Am. Nat., 68:75-76. (Also in: Genetics Soc. Am., 2:45-46.) Salivary chromosomes and the attack on the gene. I. Hered., 25:464-76. 1935 Salivary gland chromosomes in Drosophila melanogaster. Am. Nat. 69:74. The morphology of the third chromosome in the salivary gland of Drosophila melanogaster and a new cytological map of this ele- ment. Genetics, 20:301-26. With Wilson S. Stone. Chromosome fusion and speciation in Dro- sophilae. Genet., 20:327-41. With I. T. Patterson. Localization of gene loci in the third chro- mosome of Drosophila melanogaster. Rec. Genetics Soc. Am., 4:76. (Also in: Am. Nat., 70:59.) Some recent advances in our knowledge of chromosomes. Advances inModernBiology, 2:216-23. Moscow: State Biological and Med- ical Press. 1937 Eds. T. S. Painter and I. B. Gatenby. Bolles Lee's The Microtomist's Vade-Mecum, 10th ed. Philadelphia: P. Blakiston's Son & Co. With Allen B. Griffen. The origin and structure of the salivary

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THEOPHILUS SHICKEL PAINTER 335 gland chromosomes of Simulium virgatum. Genet., 22:202-3. (Also in: Rec. Genetics Soc. Am., 5:202-3.) With Allen B. Griffen. The structure and the development of the salivary gland chromosome of Simulium. Genetics, 22:612-33. 1939 The structure of salivary gland chromosomes. Am. Nat., 73:315- 30. An aceto-carmine method for bird and mammalian chromosomes. Science, 90:307-8. With Elizabeth C. Reindorp. Endomitosis in nurse cells of the ovary of Drosophila melanogaster. Chromosoma, 1 :276-83. Chromosomes in relation to heredity. In: Science in Progress. Ed. George Baitsell. New Haven: Yale University Press, 1:210-32. 1940 The chromosomes of the chimpanzee. Science, 91:74-75. On the synthesis of cleavage chromosomes. Proc. Natl. Acad. Sci. USA, 26:95- 100. A review of some recent studies of animal chromosomes. J. R. Mi- crosc. Soc., 60: 161-76. With A. N. Taylor. Nuclear changes associated with the growth of the oocytes in the toad. Anat. Rec., 78(Suppl.~:84. 1941 The effects of an alkaline solution (pH 13) on salivary gland chro- mosomes. Genet., 26: 163-64. (Also in: Rec. Genetics Soc. Am., 9: 163-64.) An experimental study of salivary chromosomes. Cold Spring Har- bor Symp. Quant. Biol., 9:47-53. 1942 With A. N. Taylor. Nucleic acid storage in the toad's egg. Proc. Natl. Acad. Sci. USA, 28:311-17. With }. }. Biesele and H. Poyner. Nuclear phenomena in mouse cancer. Univ. Tex. Publ., 4243: 1-68. 1943 With L. J. Cole. The genetic sex of Pigeon-Ring Dove hybrids as determined by their sex chromosomes. }. Morphol., 72:411-39.

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THEOPHILUS SHICKEL PAINTER 337 1964 Fundamental chromosome structure. Proc. Natl. Acad. Sci. USA, 51: 1282-85. With I. I. Biesele and R. W. Riess. Fine structure of the honey bee's royal jelly gland. Tex. J. Sci., 16:478. 1965 John Thomas Patterson (November 3, 1878-December 4, 19601. In: Biographical Memoirs, vol. 38, pp. 223-62. New York: Co- lumbia University for the National Academy of Sciences. 1966 With I. J. Biesele. The fine structure of the hypopharyngeal gland cell of the honey bee during development and secretion. Proc. Natl. Acad. Sci. USA, 55:1414-19. With J. J. Biesele. A study of the royal jelly gland cells of the honey bee as revealed by electron microscopy. In: Studies in Genetics. III. Morgan Centennial Issue, ed. Marshall R. Wheeler. Univ. Tex. Publ., 6615:475 -99. The role of the E-chromosomes in Cecidomyiidae. Proc. Natl. Acad. Sci. USA, 56:853-55. With J. J. Biesele. Endomitosis and polyribosome formation. Proc. Natl. Acad. Sci. USA, 56: 1920-25. (Also in: Science, 154:426.) 1969 The origin of the nucleic acid bases found in the royal jelly of the honey bee. Proc. Natl. Acad. Sci. USA, 64:64-66. 1971 Chromosomes and genes viewed from a perspective of fifty years of research. L. J. Stadler Memorial Symp., 1:33-42.