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J JACK SCHULTZ May 7,1904April 29,1971 BY THOMAS F. ANDERSON JACK SCHULTZ was one of the last graduate students to get his degree in classical Drosophila genetics under Thomas Hunt Morgan. This was in 1929. For the rest of his life, Jack's goal was to understand, in molecular terms, how a set of genes could direct and control the development of an organism; and he lived to see the results of his pioneering research become the foundation on which such an understanding could be based. Jack Schultz was born in Astoria, Long Island, New York, on May 7, 1904, the eldest of three sons of Morris and Bessie (Krones) Schultz. Both parents had been members of large Russian-Jewish families of rabbis, farmers, and tradesmen ]iv- ing in two small villages in the area of Minsk, near the Polish border. During the political difficulties of the 1880s and early 1 890s in Russia, the young adults of both the Schultz and Krones families tended to be ardent socialists and became in- volved in revolutionary activities. Some perished in the abor- tive revolution; others were forced to flee the country. Thus, around 1896, Morris Schultz cut short his training at trade school and came to New York. Bessie Krones also had attended a socialist worker's school and, in addition, helped her family make ends meet by running a knitting machine at home. About 1897 she, too, left her village in Russia to come to New York. Somewhat later Morris Schultz was able to help several members 393

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394 BIOGRAPHICAL MEMOIRS of their families join them. Other members stayed in Russia until World War II, when a few managed to evade death at the hands of the Nazis by escaping to Argentina. Jack's parents met soon after they arrived in New York and were married seven years later. Morris Schultz was a 'gentle soul and a great reader who had liberal, but not extreme, views. Besides being associated with the group that founded the Jewish daily newspaper, Forward, he soon became engaged in various businesses, mostly in establishing and operating neighborhood grocery stores in various industrial communities in the New York City vicinity: west New York, Astoria, Passaic, Clifton, and finally, Long Beach. He was always closely and sympathet- ically involved with the factory workers to whom he supplied food. Immediately after arriving in this country, Jack's mother found work in a knitwear factory and presently became forelady there. She was an energetic and vivacious, kind, yet demanding woman, with a strong influence over the activities of those around her. With a reputation as an excellent cook of tradi- tional dishes, she was devoted to her home and to a large circle of relatives. Jack had two younger brothers, Mortimer, who was born in 1907, and Charles, born in 1912. Their parents were ambitious for all three children and were determined to give them cultural advantages that they themselves had been denied in Russia. The children attended public schools and received, in addition, some religious trainin;, in the Jewish schools of their communities. Although their father found it difficult, he did manage to send his sons to college and to help with the financing of their grad- uate studies. Mortimer became a high school teacher of science in Long, Beach, New York, and served as chairman of the science department there for many years before he retired. Charles, who has literary interests, has been on the advertising staff of Women's Wear Daily for many years.

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JACK SCHULTZ 395 Jack was a studious child, an omnivorous reader at a very early age, as indeed he was throughout his life. He was rapidly promoted through grade school, and was therefore much younger and smaller than his classmates. This, no doubt, ac- counted at least partially for his not being at all interested in athletic activities and for his not being entirely accepted socially by his fellow students. The partial isolation that he experienced served only to reinforce his intellectual interests. His teachers and older relatives considered him a brilliant and independent, but highly impractical, boy. During these years Jack took violin lessons and frequently accompanied his mother on her Saturday afternoon expeditions to the Metropolitan Opera House. Later he took up the flute, which he played occasionally, with great pleasure, for many years. At Clifton High School he became deeply interested in the humanities; this interest was stimulated in large part by his teacher of French, Miss Mary Smith. She discovered {ack's unusual facility with languages and encour- aged him to explore the best in the literature of all the languages he had the opportunity to learn. Later in life this proficiency Bras to help him cement close friendships with people of many . .. . nationalities. During his first two years at Columbia University, he con- centrated on the humanities. He entered eagerly into the stimu- lating atmosphere of undergraduate life in New York City in the early 1920s, when there was a sudden surge of excitement in the theater, painting, literature, social philosophy, and music. This was an important stage in his development, for Jack re- tained an active interest in the humanities throughout the rest of his life. It was only in his junior year that Jack first became seriously interested in making plans for a definite career. He had decided to take a premedical course when he was suddenly, and happily, diverted. Like most young men of those times, he needed extra money for books, concert tickets, and the like; so he answered a

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396 ~ ~ ~ ~ ~ . ~ ~ BIOGRAPHICAL MEMOIRS bulletin board ad for someone to wash bottles and make fly food in the laboratory of Professor Thomas Hunt Morgan. Thus, as lUCK WOUND nave it, he was drawn to the celebrated "Fly Room" in the department of zoology. Morgan soon recognized him as a promising young man, and Jack responded bY ranidlv learning ~ to ~ ~ ~ _ _ 1 _ _ A: ~ 1 _ _ , 1 J 1 J ~ c '-~:vo~u~ary ~e~c principles anal were being established by Morgan and his group. Calvin B. Bridges and Alfred H. Sturtevant were especially kind in introducing him to the new genetic theories and techniques. Their experiments and think- ing, more perhaps than those of anyone else, influenced Schultz's later scientific work. Even as a novice, Jack not only listened to what was going on in the Fly Room but was soon tentatively contributing some small points of his own to the discussions. In the course of a few years, he was named a University Scholar, a Teaching Assistant in Zoology, and a National Research Coun- cil Fellow. More important, he enrolled as a graduate student under the guidance of T. H. Morgan, the world's foremost geneticist, and of Edmund B. Wilson, the most outstanding cytologist in the United States. Jack received his bachelor's degree in 1924, his master's in 1925, and his doctorate in 1929. His Ph.D. thesis showed that the large "Minute" class of mutations in Drosophila all pro- duced nearly identical somatic effects, and yet occurred at many different loci. Evidently a large number of independent mutations could lead to similar phenotypic effects on develop- ment. Although many theories have been advanced to explain them, the molecular mechanism for these effects remains a mystery to this day. In his thesis [act: acknowledged the many kindnesses of Morgan, Bridges, and Sturtevant. He also thanked two visitors to the laboratory: Dr. Curt Stern and Dr. Helen Redfield. Stern was a postdoctoral student from the Kaiser Wilhelm Institute in Berlin, who had come to Columbia as a Fellow of the International Education Board. Stern himself had been

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JACK SCHULTZ 397 concerned with the mosaic, normal-appearing patches in Minute flies that occurred when somatic crossin~-over eliminated the Minute region of a chromosome. ~ Tack and Curt both enjoyed and profited from the close scientific relationship that was estab- lished at Columbia and that lasted throughout the rest of iack's life. Helen Redfield had, as an undergraduate at Rice Institute, been introduced to the new developments in genetics by Her- mann i. Muller and Edgar Altenburg. Although she was at that time acting as Assistant in Mathematics, she was fascinated by the Drosophila work and was encouraged to carry out simple experiments using the sex-linked lethal genes then being ex- tensively studied by Muller and Altenburg. For graduate work she went to the University of California at Berkeley, where in the department of zoology she served as Teaching Fellow and 1 1 ~' ~ ~ . ~ ~ - received ner ran.. uncter the direction of Samuel J. Holmes. She came to Morgan's laboratory at Columbia in 1925 as a Fellow of the National Research Council. Here, in addition to completing her study of the maternal inheritance of a sex- limited lethal effect, she embarked on new studies on crossing- over in triploids. There soon developed among the younger members of the group at the Columbia laboratory an unusually rich and rewarding comradeship; and, since Jack's and Helen's interests were so closely related, they became special allies in the discussions and activities. Presently they began the col- laborative work on interchromosomal effects on crossing-over that was to continue for many years. They were married in 1926. At Columbia Jack was also closely associated with Selig Hecht and his newly formed group studying the biophysics of vision. Like Hecht, Jack was convinced of the then revolution- ary, but now almost axiomatic, idea that biological problems could be understood in chemical or physical terms only if ap- propriate physico-chemical methods were used to study them.

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398 all 1 BIOGRAPHICAL MEMOIRS 1 nls no nouot led to his early analyses of the absorption spectra of the eye pigments of various mutant stocks of Drosophila. This work suggested that the pigments were metabolically related and anticipated the hypothesis by Boris Ephrussi and George W. Beadle (1935), and by Beadle and Edward L. Tatum (1941), that each enzyme in a metabolic pathway is the product of a specific gene acting on a specific substrate. Thus, if a mutant gene fails to make an effective enzyme, the substrate of that enzyme (an intermediate pigment in the Drosophila case) might ~ ~ ~ _ _ ~ _ _ 1 ~ , 1 . ~ ~ . 1 ~ Cat acc-uI~lula~e In Ine tissue Involved tto give the eye its mutant color, for example). In 1928 Jack and Helen moved to Pasadena where Morgan and his group were setting up a new laboratory at the California Institute of Technology, under the auspices of the Carnegie Institution of Washington. The two Schultz children, Peter and Jill, were born in Pasadena and spent most of their early childhood there. At this stage Helen preferred to devote only part of her time working at the laboratory on interchromosomal effects. The results of her experiments may be briefly sum marized: In the first place, there were demonstrated, in struc- tural homozygotes with normal sequence, hitherto unsuspected positive and negative correlations of crossing-over in given regions of nonhomologous chromosomes. Some of the positive correlations were believed to be the result of response to com- mon environmental factors, such as the persistence of the polarized pattern of pairing of chromosomes seen at meiosis. However, other correlations, and the negative correlations in particular, gave evidence of a real interchromosomal influence. Extensive data were gathered on the facilitation of crossing-over by heterologous inversions (a phenomenon previously noted by C. B. Bridges and others) faith special attention to the effects of combined inversions on interference and to differential re- sponses of different reunions to different inversions. The hetero- chromatic regions were shown to be important in these inter-

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JACK SCHULTZ 399 chromosomal effects. Also, the regional differences found between triploids and diploids, both in structurally homozygous and in structurally heterozygous types, indicated that the effects of triploidy on crossing-over are to be considered an example of the phenomenon of interchromosomal influence. . At Cal Tech Jack attacked many problems in Drosophila genetics, collaborating with Sturtevant, Bridges, Theodosius Dobzhansky, David G. Catcheside, and others in showing how genes control development and how their effects are modified by suppressor genes. ~ I__ I _ _ ~ ~ ~ _ ~] ~ . ~ ~ T ~ ~ ~ ~ With Morgan and Bridges (and after DIlQ~S Learn, warn viola curry; ne co-authored eleven of the group's annual reports to the Carnegie Institution under the title, "Constitution of the germinal material in relation to hered- ity." Today such a title would imply that the group was reporting results in molecular biology, but the reports actually concerned the classical genetics of Drosophila: descriptions of new mutants, dose effects in sex determination, position effects, the elaboration of salivary gland chromosome maps, effects of X rays on crossing- over, etc. Jack contributed his share to the papers that resulted from this work and even participated in a study with Albert Tyler on the reversibility of fertilization of Urechis eggs. There is a legend that it took only two weeks for lack to teach I. B. S. Haldane the basic genetics of Drosophila. But Jack was primarily interested in the molecular biology implied by the impressive title of the Carnegie reports; he wanted to knots the chemical constitution of the genetic ma- terial and hole it functioned not only in heredity, but how it functioned to produce the phenotype. It had become obvious that chromosomes contained the genes, but their chemical com- position and chemical role in directing development of the phenotype were not amenable to study by the classical staining methods available in Morgan's laboratory. Fortunately, at that time Torbjorn 0. Caspersson in Stocl;- holm alas developing methods for the microspectrophotometry ~ 1 . ~ . . . ~ ~ ~ ~ _ _ _ ~ . . .

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400 BIOGRAPHICAL MEMOIRS of cells in the ultraviolet end of the spectrum; Jack, with his training in both genetics and biophysics, was the ideal person to use these methods in following the metabolism of the nucleic acids in cells. So in 1937, under the auspices of the Rockefeller Foundation, he went to work with Caspersson. Jack and his family spent two very pleasant and stimulating years with their new Swedish friends in Stockholm; those with whom they formed especially warm and long-lasting friendships included the Casperssons, the Gert Bonniers, and the John Runnstroms. In the laboratory, Jack and Caspersson soon showed that there is indeed a relation between the metabolism of the two kinds of nucleic acids: The nucleolus was found to contain large amounts of pentose nucleic acid, whereas the chromosomes themselves largely contain deoxypentose nucleic acid. More- over, the observation that the cytoplasm of rapidly dividing cells is rich in pentose nucleic acid as compared to resting cells gave them a glimmer of current thought as to the mechanism of gene action: mRNA synthesis, ribosome synthesis, tRNA syn- thesis, and protein synthesis. [ack's review of this and of other work, published as early as 1941, "The evidence of the nucleo- protein nature of the gene," concludes, "At the present time the properties of the genes and of nucleoprotein metabolism are evidently parallel: specificity, self reproduction, relations to synthesis and distribution of nucleoproteins in the cell, all are what they should be were the genes nucleoproteins. It would seem therefore, that our present task is to develop the physiology of the nucleoproteins into an effective physiology of the genes." When one also recalls that in 1932 lack had been involved with Bridges in the discovery of specific suppressor genes, one can appreciate his reaction to current concepts of how gene action is controlled by interactions among operons, repressors, . and inducers: It was one of deja flu. As an explorer, Jack had long before sketched that intellectual territory, but its scientific

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JACK SCHULTZ 401 relevance became generally accepted only years later after other molecular biologists had settled into it and painstakingly har- vested specific chemical evidence for its reality. When World War II broke out, Helen and the children sailed for the United States, but lack tarried in Edinburgh lone enough to attend the International Genetics Congress and barely managed to catch one of the last civilian boats home. Back in Pasadena, he wanted to continue the work he and Caspersson had begun. But, as George Rudkin tells us, ``The years following t~ack's] return from Sweden were marked by the search for financial support for the acquisition and operation of the complicated, expensive instrumentation of the type de- veloped in Stockholm, this in an era when grants were rare and in a laboratory where simple equipment was stressed. The period (1939-1943) ended with a harried year spent partly as visiting professor with Lewis John Stadler, at Missouri, working on variegation in corn, partly at Cal Tech and partly at Woods Hole, not far from the Marine Laboratory of the Philadelphia- based Lankenau Hospital Research Institute at North Truro, Massachusetts. That same period saw the completion of a new review, this time from a chemical point of view, 'The Gene as a Chemical Unit,' much of which is still illuminating thirty years later. In 1943, he joined the Lankenau organization, then under the directorship of Dr. Stanley P. Reimann. "Reimann was in the process of building up The Institute for Cancer Research (as it was later called), which was dedicated to the proposition that logical solutions to the cancer problem would come from an understanding of the basic mechanisms underlying development and growth. Jack enthusiastically brought his expertise in 'genetics, embryology, and physiology to this new enterprise. He became immersed in trying to under- stand the many facets of cancer and, as one of the few geneticists in the field, wrote a number of classical reviews on the subject." Jack and Helen were warmly welcomed to the Philadelphia

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402 BIOGRAPHICAL MEMOIRS scene by members of the faculty of the University of Pennsyl- vania, particularly by Charles W. Metz and his associates. The Schultz family lived first in the village of Ithan, on the Main Line west of Philadelphia. In 1949, when the Institute moved from its crowded and makeshift quarters at the old Lankenau Hospital on Girard Avenue, to its modern new building in Fox Chase, they moved to Elkins Park and finally settled in Hunting- don Valley, a suburb in the northeast, where they could happily do some gardening. Jack and Helen also spent many delightful summers at Cold Spring Harbor with such friends as Milislav Demerec, Theodosius Dobzhansky, Curt Stern, Ernst W. Caspari, Berwind P. Kaufmann, Alfred E. Mirsky, and Barbara McClintock. In his own laboratory, Jack and his colleagues worked on projects in many areas. With human material easily available from the Lankenau Hospital, Dr. Reimann soon induced Jack to study human chromosomes. As a result, in 1946 he and Patricia St. Lawrence were able to map two of the chromosomes associated faith the nucleolus in preparations of human pachytene chromosomes and to show that, like the pachytene chromosomes of other species, they have distinctive chromomere patterns. This area of research has been continued at The Institute for Cancer Research by Jack's former student, David Hungerford. Another project involved devising minimal media for grow- ing Drosophila cultures as a preparation for studying the one- gene-one-enzyme hypothesis in a higher organism. Although it was found that different stocks indeed had different require- ments for such substances as tryptophan, the project was eventu- ally dropped because of the difficulty in getting clear-cut non- leaky, nutritional mutants. In another, somewhat related project, Jack and Elizabeth K. Patterson followed the activities of such enzymes as the peptidases during the development of specific organs in Drosophila. Measurements of the nucleic acid

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TACK SCHULTZ 413 chromosome of Drosophila melanogaster. Proc. Natl. Acad. Sci., 17:513-18. With T. H. Morgan and C. B. Bridges. The constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 30:408-15. 1932 The behavior of vermilion-suppressor in mosaics. Proc. Natl. Acad. Sci., 18:485-86. The developmental system affected by the genes for eye color in Drosophila. In: Bloc. 6th Int. Congr. Genet., ed. by Donald F. tones, vol. 2, pp. 178-79. New York: Brooklyn Botanic Garden. (A) With C. B. Bridges. Methods for distinguishing between duplica- tions and specific suppressors. Am. Nat., 65:323-34. With A. Tyler. Inhibition and reversal of fertilization in the eggs of the Echinoid worm, Urech is can pot J. Exp. Zool., 63:509-30. With T. H. Morgan and C. B. Bridges. The constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 31:303-7. 1933 X-ray effects on Droso phila pseud o-o bscura. Genetics, 18: 284-91. With T. Dobzhansky. Triploid hybrids between Drosophila melano- gaster and Drosophila simulans. J. Exp. Zool., 65:73-82. With T. H. Morgan and C. B. Bridges. The constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 32:298-302. 1934 With T. Dobzhansky. The relation of a dominant eye color in Drosophila melanogaster to the associated chromosome rear- rangement. Genetics, 19:344-64. With T. Dobzhansky. The distribution of sex factors in the X- chromosomes of Drosophila melanogaster. J. Genet., 28:349-86. With T. Dobzhansky. Sex in Drosophila and other organisms. Am. Nat., 68:190-91. With T. H. Morgan and C. B. Bridges. The constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 33: 274-80.

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414 BIOGRAPHICAL MEMOIRS 1935 Aspects of the relation between genes and development in Drosoph- ila. Am. Nat., 69:30-54. With T. H. Morgan and C. B. Bridges. The constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 34: 284-91. 1936 Variegation in Drosophila and the inert chromosome regions. Proc. Natl. Acad. Sci., 22:27-33. With T. H. Morgan and C. B. Bridges. The constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 35: 289-97. Radiation and the study of mutation in animals. In: Biological Eects of Radiation, ed. by B. M. Duggar, vol. 2, pp. 1209-61. New York: McGraw-Hill Book Co. 1937 With D. C. Catcheside. The nature of closed X-chromosomes in Droso phila melanogaster. J. Genet., 35: 315-20. With T. H. Morgan and C. B. Bridges. The constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 36:298-305. 1938 With T. Caspersson. Nucleic acid metabolism of the chromosomes in relation to gene reproduction. Nature, 142:294-95. With T. H. Morgan and C. B. Bridges. The constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 37: 304-9. 1939 With T. Caspersson. Heterochromatic regions and the nucleic acid metabolism of the chromosomes. Archiv fur experimentelle Zell- forschung besonders Gewebezuchtung, 22: 650-53. With T. Caspersson. Pentose nucleotides in the cytoplasm of grow- ing tissues. Nature, 143: 602-3. With T. H. Morgan, C. B. Bridges, and V. Curry. Investigations on the constitution of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 38:273-77.

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JACK SCHULTZ 1940 415 With T. Caspersson. Ribonucleic acids in both nucleus and cyto- plasm, and the function of the nucleolus. Proc. Natl. Acad. Sci., 26:507-15. With T. Caspersson and L. Aquilonius. The genetic control of nucleolar composition. Proc. Natl. Acad. Sci., 26:515-23. With T. H. Morgan and V. Curry. Investigations on the constitu- tion of the germinal material in relation to heredity. Carnegie Inst. Wash. Year Book, 40:282-87. 1941 The evidence of the nucleoprotein nature of the gene. Cold Spring Harbor Symp. Quant. Biol., 9:55-65. The function of heterochromatin. In: Proc. 7th Int. Congr. Genet., ed. by R. C. Punnett, pp. 257-62. London: Cambridge Univ. Press. 1943 Physiological aspects of genetics. Annual Review of Physiology, 5:35-62. (Unsigned) Bridges, Calvin Blackman. National Cyclopedia of American Biography, vol. 30, pp. 374-75. New York: James T. White 8c Co., Inc. 1 944 The gene as a chemical unit. In: Colloid Chemistry, Theoretical and Applied, vol. 5, ed. by J. Alexander, pp. 819-50. New York: Reinhold Publishing Corporation. With F. S. lose. Differentiation of chromosomal proteins by stain- ing techniques. Rec. Genet. Soc. Am., 13:34. (A) 1945 Genetic factors in cancer. Clinics, 4:118-27. With F. S. Jose. Differentiation of chromosomal proteins by staining techniques. Genetics, 30:20-21. (A) 1946 The analysis of chromomere patterns in man. Rec. Genet. Soc. Am., 14:58-59. (A) With P. St. Lawrence and D. Newmeyer. A chemically defined

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416 BI OGRAPHI CAL ME M OIRS medium for the growth of Drosophila melanogaster. Anat. Rec., 96:540. (A) 1 947 The nature of heterochromatin. Cold Spring Harbor Symp. Quant. Biol., 12: 179-91. Nuclear differentiation and the origin of tumors. Cancer Res. 7:41-42. (A) With G. T. Rudkin. Evolution of nutritional requirements in ani- mals: amino acids essential for Drosophila melanogaster. Anat. Rec., suppl., 99:613. (A) 1948 ., With G. T. Rudkin. Absence of a sparing action of tryptophane on the nicotinamide requirements of the fly, Drosophila melano- gaster. Fed. Proc., 7:185. (A) 1949 Chemical genetics in Drosophila. Meeting of American Chemical Society, p. 41C. (A) With G. T. Rudkin. A comparison of the tryptophane require- ments of mutant and wild type Drosophila melanogaster. In: Proc. 8th Int. Congr. Genet., Hereditas suppl., pp. 652-53. (A) With G. T. Rudkin. Nutritional requirements and the chemical genetics of Drosophila melanogaster. In: Proc. 8th Int. Congr. Genet., Hereditas suppl., pp. 657-58. (A) With T. Caspersson. Nucleic acids in Drosophila eggs and Y-chro- mosome effects. Nature, 163: 66-67. With E. K. Patterson and M. E. Dackerman. Peptidase activities of extracts of salivary glands of Drosophila melanogaster. J. Gen. Physiol., 32:607-22. With E. K. Patterson and M. E. Dackerman. Peptidase increase ac- companying growth of the larval salivary gland of Drosophila melanogaster. J. Gen. Physiol., 32:623-45. With R. C. MacDuffee and T. F. Anderson. Smear preparations for the electron microscopy of animal chromosomes. Science, 110: 5-7.

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JACK SCHULTZ 417 With P. St. I,awrence. A cytological basis for a map of the nucleolar chromosome in man. Journal of Heredity, 40:31-38. 1950 Interrelations of factors affecting heterochromatin-induced variega- tion in Drosophila. Genetics, 35:134. (A) The question of plasmagenes. Science, 111:403-7. 1951 \Vith T. Caspersson. Cytochemical measurements in the study of the gene. In: Genetics in the 20th Century, ed. by L. C. Dunn, pp. 155-71. New York: Macmillan Publishing Co., Inc. The effect of ultra-violet radiation on a ring chromosome in Zea Mays. Proc. Natl. Acad. Sci., 37: 590-600. With H. Redfield. Interchromosomal effects on crossing over in Drosophila. Cold Spring Harbor Symp. Quant. Biol., 16:175-97. With Margaret M. Service. Genetic differences in the requirement for ribosenucleic acid and glycine in Drosophila melanogaster. Fed. Proc., 10:245. (A) 1952 Interrelations between nucleus and cytoplasm: problems at the biological level. Exp. Cell Res., suppl. 2. Chemistry and physi- ology of the nucleus), pp. 17~2. 1953 With G. T. Rudkin and H. Temin. Evidence for the equality of chromosomal material in white and wild-type bearing sections of salivary gland chromosomes of Drosophila melanogaster. Genetics, 38:686. (A) With D. A. Hungerford. Characteristics of pairing in the salivary gland chromosomes of Drosophila melanogaster. Genetics, 38: 689. (A) With N. Bischoff. A comparison of the effects of antimetabolites and systemic carcinogens on morphogenesis in Drosophila melano- gaster. Proc. Am. Assoc. Cancer Res., 1:48. (A) With l. F. Aronson, D. A. Hungerford, and G. T. Rudkin. A test system for the microspectrophotometry of chromosomes. l. Histochem. Cytochem., 1:398. (A) With L. Levenbook and E. Travaglini. Nucleic acids and free poly-

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418 BIOGRAPHICAL MEMOIRS nucleotide fragments in the egg of Drosophila. Anat. Rec., 117 585. (A) l 9~4 With i. F. Aronson and G. T. Bodkin. A comparison of giant X chromosomes in male and female Drosophila melanogaster by cytophotometry in the ultraviolet. I. Histochem. Cytochem., 2:458-59. (A) With E. K. Patterson, H. Lang, and M. E. Dackerman. Chemical determinations of the effect of the X and Y chromosomes of the nucleic acid content of the larval salivary glands of Drosophila melanogaster. Exp. Cell Res., 6: 181-94. The place of cytogenetics in cancer research. In: Proceedings of the 2d National Cancer Conference, pp. 1152-60. New York: American Cancer Society, Inc. With T. S. Hauschka. Cytologic aspects of immunogenetic spe- cificity. Transplantation Bulletin, 1:203-6. 1955 With N. Rothman and M. M. Aronson. The growth and morpho- genesis of Drosophila melanogaster as criteria for screening tests. Cancer Res., suppl. no. 3, pp. 86-99. With G. T. Rudkin, J. F. Aronson, and D. A. Hungerford. A com- parison of the ultraviolet absorption of haploid and diploid salivary gland chromosomes. Exp. Cell Res., 9:193-211. 1956 With L. Levenbook and E. C. Travaglini. The effect of the Y chromosome on nucleic acids and polynucleotide fragments in the unfertilized Drosophila melanogaster egg. Be Congres Inter- national de Biochimie, Brussels. Resume des Communications 70, ed. by Claude Liebecq. New York: Academic Press, Inc. The shape of the cancer problem. Cancer Res., 16:720-21. With G. T. Rudkin. An attempt to compare the sarcosomes of diploids and triploids in Drosophila melanogaster. Cold Spring Harbor Symp. Quant. Biol., 21:303-6. The relation of the heterochromatic chromosome regions to the nu- cleic acids of the cell. Cold Spring Harbor Symp; Quant. Biol., 21: 307-28.

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JACK SCHULTZ 1958 419 With E. C. Travaglini. Cytoplasmic DNA deoxyribosides and Y chromosome heteroploidy in the egg of Drosophila melanogaster. Proc. Am. Assoc. Cancer Res., 2:352. (A) Malignancy and the genetics of the somatic cell. Annals of the New York Academy of Science, 71: 994-1008. Bridges, Calvin Blackman. Dictionary of American Biography, vol. 22, suppl. 2, pp. 60-62. New York: Charles Scribner's Sons. With L. Levenbook and E. C. Travaglini. Nucleic acids and their components as affected by the Y chromosome of Drosophila melanogaster. I. Constitution and amount of the ribonucleic acids in the unfertilized egg. Exp. Cell Res., 15:43-61. With E. C. Travaglini and L. Levenbook. Nucleic acids and their components as affected by the Y chromosome of Droso phila melanogaster. II. Nucleosides and related compounds in the acid soluble fraction of the unfertilized egg. Exp. Cell Res., 15:62-79. The nature of variegation in Drosophila. In: Proc. 10th Int. Congr. Genet., vol. 2, p. 255. Toronto: Univ. of Toronto Press. (A) 1959 Antigens and antibodies as cell phenotypes. Science, 129:937-43. Integrative mechanisms in biology. Am. Nat., 93:209-11. With l. T. Freed, l. l. Engle, and G. T. Rudkin. Ultraviolet radia- tion effects on Ehrlich ascites tumor cells. Journal of Biophysics and Biochemical Cytology, 5: 205-16. The role of somatic mutation in neoplastic growth. In: Genetics and Cancer, pp. 25~2. Thirteenth Annual Symposium on Fundamental Cancer Research. Austin: Univ. of Texas Press. 1960 With G. T. Rudkin. DNA content in relation to the genetic unit in Drosophila melanogaster. Tenth International Congress of Cell Biology, Paris, p. 71. Paris: L'Expansion scientifique franc~aise. (A) Studies on the effects of ultraviolet radiation on cell structure and

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420 BIOGRAPHICAL MEMOIRS behavior. IRE Transactions on Medical Electronics, vol. ME7, no. 4, p. 367. (A) With G. T. Rudkin. Direct measurement of deoxyribonucleic acid content of genetic loci in Drosophila. Science, 132:1499-1500. (A) With E. Travaglini. Genetic control of RNA constitution in the unfertilized Drosophila egg. In: Symposium of the International Institution of Embryology, Pallanza, Italy, pp. 20-22. (A) 1961 With G. T. Rudkin. Disproportionate synthesis of DNA in polytene chromosome regions in Drosophila melanogaster. Genetics, 46: 893-94. (A) 1963 The genetics of somatic cells. In: Proceedings of the 16th Interna- tional Congress of Zoology, ed. by John A. Moore, vol. 3, pp. 216-21. Washington: National Research Council. With H. Redfield. Recombination for the X of structurally stand- ard triploids of Drosophila melanogaster. Drosophila Informa- tion Service, 38: 46-48. 1964 With F. T. Ashton. Stereoscopic analysis of the fine structure of chromosomes in diploid Drosophila nuclei. l. Cell. Biol., 23:7. (A) 1965 With E. C. Travaglini. Evidence for homeostatic control of ribo- somal content in Drosophila melanogaster eggs. Genetics, 52:473. (A) Genes, differentiation, and animal development. Brookhaven Sym- posia in Biology, 18:116-47. With E. C. Travaglini. Changes in the base composition of RNA species during the embryonic development of Droso phila melanogaster. J. Cell. Biol., 27:1 08A. (A) 1 966 Perspectives for the conference on the nucleolus. National Cancer Institute Monograph, 23: 1-9.

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JACK SCHULTZ 421 1967 Innovators and controversies. Book review of The Gene: A Critical History, by E. A. Carlson. Science, 157:296-301. 1968 With E. C. Travaglini Andy. Petrovic. Two satellite cytoplasmic DNA's in Drosophila. J. Cell. Biol., 39~334~:136A. (A) 1970 With E. Hadorn and R. Gsell. Stability of a position-effect varie- gation in normal and transdetermined larval blastemas from DYOSOPhila melanogaster. Proc. Natl. Acad. Sci., 65:633-37. With B. S. Fansler, E. C. Travaglini, and L. A. Loeb. Structure of Drosophila melanogaster dAT replicated in an in vitro system. Biochemical and Biophysical Research Communication, 40: 1266-72. Foreword. (In honor of Torbjorn Caspersson on his sixtieth birth- day) Exp. Cell Res., 62:3-4. 1971 With F. T. Ashton. The three-dimensional fine structure of chromo- somes in a prophase Drosophila nucleus. Chromosoma, 35: 383-92. 1972 New ideas. Perspectives in Biology and Medicine, 15:435-42. With E. C. Travaglini and l. Petrovic.~ Characterization of the DNA in Drosophila melanogasteY. Genetics, 72:419-30. With E. C. Travaglini and l. Petrovic. ~ Satellite DNA's in the embryos of various species of the genus Drosophila. Genetics, 72:431-39. With E. C. Travaglini.~ Circular DNA molecules in the genus DYOSOPhila. Genetics, 72:441-50. 1973 Human values and human genetics. Am. Nat., 107:585-97. ~ Written by co-authors after the death of Dr. Schultz.

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422 BIOGRAPHICAL MEMOIRS BIOGRAPHICAL NOTICES Talbot, T. R., Jr., 1971. Jack Schultz, May 7, 1904-April 29, 1971. Sixteenth Scientific Report, The Institute for Cancer Research, Philadelphia, pp. 15-16. Rudkin, G. T., 1971. Jack Schultz, 7 May 1904-29 April 1971. Genetics, 68:s97-98. Rudkin, G. T., 1971. Jack Schultz, May 7, 1904-April 29, 1971. Sixteenth Scientific Report, The Institute for Cancer Research, Philadelphia, pp. 97-101. Rudkin, G. T., 1972. Jack Schultz, May 7, 1909WApril 29, 1971. In: Results and Problems in Cell Differentiation, ed. by W. Beer- mann, vol. 4, pp. 5-6. New York: Springer-Verlag.

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