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CURT STERN August30, 1902-October23, 1981 BY JAMES V. NEEL TH E R E H A V E B E E N three remarkable periods in the his- tory of modern genetics. The first of these, in the early years of this century, encompasses the rediscovery and confirmation of Mendel's findings and the enunciation of the chromosomal theory of inheritance. The seconct period is characterizes! by a concatenation of discoveries regarding the more precise mechanisms of chromosomal behavior, largely based on the use of Drosophila and emanating from the "fly room" at Columbia University. The third is the period in which we now fins! ourselves, initiated in the 1940s by the seminal observations of Avery and collaborators and the later phage work demonstrating that the essential genetic material was DNA and the demonstration, using Neurospora, that genes have essential anc! specific roles in the synthesis of pro- teins. This was followed by the elucidation of the structure of DNA, leading to a cascade of discoveries concerning DNA fine structure and how it can be manipulated. Each of these flowerings clominatect the conceptual biological thinking of the time. 'NOTE: This memoir is an expanded version of a manuscript published in the An- ,~ual Review of Genetics. I am deeply indebted to the many people who have contrib- uted their remembrances and perspective to the writing of this memoir. 443

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444 BIOGRAPHICAL MEMOIRS THE EARLY YEARS Curt Stern, the subject of this memoir, must be regarclect as one of the principal participants (ancl last survivors) of the fly room period. Born in Hamburg, Germany, on August 30, 1902, the first son of Barned anct Anna Stern, he early displayed a strong interest in natural history, ranging from microscopic studies of pondwater to the zoological collections that enliven parental responsibilities. In these interests he received unusual support from two extraordinary high school teachers, who encouraged him to undertake the study of zoology. His father, who was in the dental supply business, and his mother, a schoolteacher, were also highly supportive as their son's biological interests unfoIclecl. Following the family's move to a suburb of Berlin, he entered the University of Berlin in 1920. He must almost immediately have found his way to the Kaiser Wilhelm Institute, where he conductect his doctoral studies. He received a Ph.D. from the University of Berlin in 1923, at that time the youngest person ever to receive the degree from the university. There is no record of an unclergraduate degree. Attendance at the university re- quirec! one to two hours of commuting each way. To achieve his degree so early under these circumstances was an early, clear signal of the remarkable combination of high intellec- tual ability, photographic memory, and stamina that was to characterize his career. Stern's Ph.D. thesis was a descriptive cytological study of mitosis in a protozoan of the order Heliozoa, uncler the di- rection of one of the truly eminent protozoologists of that era, Max Hartmann. One perceives here the momentum of a high school fascination with microscopic pond life. By the time he finished his thesis, he knew he would not remain a ' The only remaining survivor of the "fly room" known to the author is Helen Redfield (firs. Jack) Schultz.

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CURT STERN 445 protozoologist. Reading widely, he became intrigued by ge- netics. A paper by Richard Goldschmidt, then director of the Institute, on the basis for crossing-over caught his attention. Stern felt Goldschmidt's interpretation was incorrect and wrote a critique, which, after great hesitation, he submitted to Goldschmidt. (He was at that time probably the most jun- ior fellow at the Kaiser Wilhelm Institute.) Some six months later, Goldschmidt returned the paper without comment, but shortly thereafter called Stern into his office. Those of us privileged to see Goldschmidt in action after his immigration to the United States will recall the personification at least superficially of the Geheimrat Professor. One can imagine the trepidation with which Stern approached the meeting. With few preliminaries, Goldschmidt offered Stern a fellow- ship, recently funded by the International Education Board of the Rockefeller Foundation, to study genetics with the Morgan group. There apparently was never any discussion of the critique. DR OS OPHILA Stern arrived at Columbia in 1924. The fly room was near or at its zenith. Morgan in ~ 9 ~ 0 had demonstrated sex-linked inheritance in Drosophila and then the recombination of two sex-linked alleles, one responsible for white eye and the other for rudimentary wing. These exciting discoveries, along with the obvious potentiality of Drosophila as an experimental or- ganism, had attracted to Morgan's laboratory a now historic triumvirate Sturtevant, Bridges, and Muller. The publica- tion by Morgan, ~ ' ~3 biturtevant, Muller, and Bridges of "The Mechanisms of Mendelian Heredity" in 1915, which brought together the early data on autosomal as well as X-chromo- some inheritance and linkage, with linkage maps and the evi- dence for nondisjunction, had made clear how major the de- velopments at Columbia were. One result was a rising

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446 BIOGRAPHICAL MEMOIRS number of students and visiting investigators, who, together with Sturtevant, Muller, and Bridges, were all crowcled into a room 16 by 23 feet in size, which contained eight working desks! (For a forthright statement of the physical "together- ness" of this setting, and especially the space allotted to Stern, see Provine ~ 98 ~ .) There quickly ensucc! several "beginner papers," but the first major result of this fellowship, published in 1926-1927, was the demonstration by a combination of genetic and cy- tological techniques that the anomalous genetic behavior of the "bobbed bristles" trait couIcT be explained by the Y-linkecl inheritance of the responsible allele (plus a homologous locus on the X). Until then, the Y-chromosome of Drosophila, al- though associated with male fertility, had been consiclered as otherwise genetically empty. Although Y-linkocI inheritance had previously been demonstratecI by Schmidt in the fish, Lebistes reticulates, this demonstration that abnormalities of Y- chromosome behavior (that is, the occurrence of XXY fe males) accounted for abnormalities in the inheritance of the bobbed trait was unusually elegant for the times, no doubt benefiting greatly from the cytological clemancis of Stern's Ph.D. thesis. His second major contribution appeared in 1931: the demonstration, a ~ 1 using cytologically abnormal X-chromo- somes, one with an X-Y transIocation, one with an X-IV transIocation, that the genetic phenomenon of crossing-over was accompanies! by a physical exchange between the chro- mosomes. (Simultaneously, Creighton and McClintock quite independently clemonstrated the same phenomenon in corn.) This was shortly followed by an ingenious clemonstra- tion (back to the Y-chromosome) that as he acIdect supernu- merary Y-chromosomes bearing the bobbed allele to Droso- phila, the trait gradually ctisappearect, understandable now that we know the bobber] alleles are characterized by varying

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C U RT S T E RN 447 degrees of unclerproduction of ribosomal RNA. At that time the demonstration that adding enough defective genes wouIcI produce normality was novel. Stern's last major contribution to our understanding of the chromosomal basis of inheritance was published in 1936. He had returned to Germany in 1926, but had come back to the United States in 1932, on a second fellowship from the Rockefeller Foundation, spending the year at the California Institute of Technology in the company of a remarkable collection of geneticists: Morgan, Sturtevant, Dobzhansky, Bridges, Schultz, Emerson, Darlington, Kaufmann, and Lin- degren. He had married an American citizen, Evelyn Som- merfield, in 1931. In 1933, when he was clue to return to Germany, Hitler came to power, a clevelopment whose tragic implications for German Jews has been only too well docu- mentecI. While Stern remained in the United States, Evelyn returned to Germany to seek some cautious acivice from his colleagues. What she Earned convinced them it wouIcI be wise not to return. (He became a U.S. citizen in 1939.) Stern accepted a temporary position at Western Reserve University, but quickly moved to the University of Rochester, where he was to remain until 1947, serving from 1941 to 1947 both as chairman of the Department of Zoology and chairman of the Division of Biological Sciences. Shortly after he arrived at Rochester, he began to inves- tigate what was then a puzzling phenomenon: the occurrence in female flies heterozygous for one or several sex-linkoct re- cessive alleles of epidermal spots manifesting the effects of one or all of these alleles or even, if the alleles were on ctif- ferent but homologous chromosomes, "twin spots," exhibit- ing the phenotypes associated with both alleles. A classical analysis revealed that the only consistent explanation re- quired a previously unrecognized phenomenon, mitotic crossing-over. ~ wouIcI like to suggest that this paper was the

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448 BIOGRAPHICAL MEMOIRS ens! of the "chromosomal era" in the history of Drosophila as an experimental organism. It was to continue to provide in- sights on other problems, but the story of how its chromo- somes behaved (in the classical sense) was now essentially complete. In several autobiographical sketches, Stern has empha- sizect that he never attempted any grand research design, but simply "followocl his nose." The beauty of Drosophila was how quickly one could move from one major issue to the next, providing you knew how to manipulate the fly stocks at which Stern was probably seconc! only to Muller. That nose led him unerringly to basic issues. Space permits mention of only three of the outstanding contributions subsequent to 1936 that depended on experiments with Drosophila. T. Isoalleles. The recessive allele cubitus interruptus (ci), lo- cated in the fourth chromosome, causes a gap of variable length in the fourth wing vein. Working with strains thought to be isogenic except for fourth chromosomes of different origins, as well as with strains in which a deficiency of the region encompassing the ci locus was present, and manipu- lating temperature, Stern in 1943 demonstratecl that normal alleles of ci differed greatly in their potency, as measured by their ability to mollify the expression of the ci trait in hetero- zygotes or hemizygotes for this locus. He termed these dif- ferent normal alleles "isoalleles." This demonstration of a range of genetic variation beyond that easily envisioned pre- saged (and may now find an explanation in) the demonstra- tion years later of extensive inapparent biochemical variation. At that time there was still concern amounting to clisbelief among some biologists and paleobiologists that the kinds of traits arising through mutation in Drosophila couIct possibly serve as the stuff of evolution. Since these alleles of small effect presumably arose from the mutational process, this cli- rection of attention to traits of lesser effects player! a signifi

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CURT STERN 449 cant role in what Mayr has termec! "the evolutionary synthe- sis" (Mayr and Provine ~9801. 2. Genetic effects of low-level racliation. During and after World War Il. Stern, in collaboration with Spencer, Caspari, and Uphoff, was drawn into studies of low-level radiation effects, studies sponsored by the Rochester branch of the U.S. Army's Manhattan Engineering District. The question, in the context of the advent of the atomic bomb, was obvious: Was there a threshold in the genetic effects of radiation? The fincI- ing now a cornerstone of racliation genetics was clear: "Viewing all experiments together, it appears that radiation at low doses, administered at low intensity, induces mutation in Drosophila sperm. There is no threshold below which ra- cliation fails to induce mutations" (Uphoff and Stern 19491. It was un(loubtedly this background that led to a term ~ ~ 950- 1953) on the Advisory Committee to the Division of Biology anc! Medicine of the Atomic Energy Commission, a critical period in the (development of the AEC's policy of broadly basect research into radiation effects. The work on the effects of low-level radiation hac! used sex-linkocI lethals as indicator traits. Since it was felt that ra- cliation produces disproportionate numbers of lethals (as the normal spectrum of mutation is unclerstood), an important question was the effect of these "recessives" when heterozy- gous. Given the ratio of heterozygotes to homozygotes pre- clictect by the Harcly-Weinberg formulation, for a rare allele even a small heterozygote effect for an autosomally inherited recessive lethal coup outweigh the impact of the occasional homozygote. Stern-with Carson, Kinst, Novitski, and Up- hoff in 1952 established that under their conditions, the average viability of heterozygotes for sex-linkecl lethals was 96.5 percent normal, a figure still standard. Their ciata clict not permit any distinction between the effects on viability of spontaneous anc! inclucecI mutation.

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450 BIOGRAPHICAL MEMOIRS 3. The extent of cell autonomy in gene expression and the genetic control of patterns. In 1947, Stern left Rochester to become professor of zoology an(l, in 1958, professor of genetics as well, at the University of California, Berkeley; he retirect from these positions in 1970. At Berkeley he returned to an oIc] problem; the basis of the attack had been laid with the 1936 paper on somatic crossing-over and segregation. These . studies had demonstrated a high degree of cell autonomy In the expression of genetic constitution in Drosophila. On the other hanct, transplantation experiments (with Hadorn) hac! demonstrated that the color of the vase efferentia was depen- clent on that of the testis attacher! to them and not on the genetic constitution of the ducts themselves, and that the shape of the testis (spiral or oval) was dependent on the ge- netic constitution of the sperm ducts to which it was attached ~ ~ 939-l 94 ~ ). In the ~ 950s ant! much of the ~ 960s, most of Stern's research efforts, often in collaboration with his stu- clents, were directed toward the difficult problem of the ge- netic control of differentiation, and especially of patterns, still using Drosophila as an experimental organism. The various types of regularly arranged chaetae, so ob- v~ous when one inspects a fly, proved most useful in these interests. Studies with Hannah-Alava, employing genetic mo- saics of various derivations, demonstrated a new level of com- plexity in embryological determination: Differentiation of the male sex-comb (a specialized row of chaetae) clependec! on a field effect within which the development of the sex . comb teeth was cletermined down to very small patches of cells by the sex (maleness) of the cells. These and other studies led to what Stern in 1954 termed the "prepattern hypothesis." Prepattern was a descriptive term for any kind of spatial differentiation in development, development being regarded as a succession of prepatterns. Within the prepat

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CURT STERN 451 tern there are singularities, to which developing cells respond according to their genetic competences. Much of his later work on this subject, well summarized in 1978 by his col- league and collaborator Dr. C. Tokunaga, was directed to- war(1 defining, largely through the use of genetic mosaics, the nature of prepatterns, singularities, and competences. The familiarity with Drosophila mosaics these inquiries cle- mancled inspired an omnivorous interest in mosaics of all types, reflected in the Prather Lectures at Harvard in 1965 in which he summarized his activities in this fielcI. One of Stern's last papers on Drosophila, in 1969, once again illustrated his ability to take full advantage of Drosophila as an experimental organism. The earlier studies on somatic cell crossing-over hacI not provicled a clear approach to the relative frequency of somatic cell versus meiotic crossing- over. Since then, in the 1950s, compound or complex loci hacI been recognized in Drosophila melanogaster, such as the "white eye" (w) locus, within which meiotic crossing-over could occur. As a manifestation of the remarkable degree of cell autonomy in Drosophila, each incliviclual facet of its com- pound eye expresses its genotype as regards pigmentation independently of the other. Stern scored female flies heter- ozygous for two different w alleles between whose mutational sites the frequency of meiotic crossing-over had been cleter- minect, for red-colored spots in their white eyes. Four such spots were found in a total of 6,137 flies. No such spots were observed in the eyes of 27,557 controls. On the assumption that the rect spots resulted from somatic cell recombination in a cell of the developing eye disc, and that the eyes of the flies scored collectively provided a minimum of 9 x lob mi- toses in which the results of somatic recombination couIct be observecl, these 4 spots suggested a frequency of recombi- nation between these mutational sites of less than ~ or 2 in

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452 BIOGRAPHICAL MEMOIRS 2 x lO`; mitoses (! if the exchange was nonreciprocal, 2 if reciprocal). This was 400 to 800 times less common than . . . melotlc crossing-over in t ne same region. HUMAN GENETICS Technically, Stern's advent into the other fielc! of genetics with which his name is so prominently associated, human genetics, dates from a paper entitlecl "Welche Moglichkeiten die Ergebnisse der experimentellen Vererbungslehre ciafur, class clurch verschieclens Symptome charakterisierte Ner- venkrankheiten auf gleicher erblicher Grundlage beruhen?" published in 1928 in Nervenarzt. It is a very clear statement, directed to physicians, of a principle now commonly ac- ceptecI: Indistinguishable phenotypes may have very (liffer- ent genetic bases. His serious entry into the fielct, however, can be ciated more precisely to 1939. That year he supervised his first graduate student seminar in the field of human ge- netics. As one of the half-clozen students who met weekly for a semester, T still have the list of papers he chose for review. Two impressions stand out. First, he hac! manager! to select what little solid ciata existed; the contrast between the then anct the now of human genetics, developments within a single generation, is simply staggering. Second, given the excesses of American eugenicists, and especially the Nazi racism from which he personally had suffered, one might have expected some bitterness and an occasional diatribe on these mon- strous perversions; Stern kept the discussion all science. This occasion must mark the beginning of the most suc- cessful textbook on human genetics ever written. In the course of three English editions ~ 1949, ~ 960,1973),Principles of Human Genetics sold 62,337 copies; there is no way to esti- mate accurately the number of copies soIct of the German, Japanese, Spanish, Portuguese, Hincti, Polish, and Russian

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CURT STERN 463 The determination of color in the vase efferentia of Drosophila me- lanogaster. Science, 86:408. Interaction between cell nucleus and cytoplasm. Nature, 140:770- 71. Methylene blue-staining of peripheral nervous system in Droso- phila melanogaster. DIS, 8:90. 1938 With E. Hadorn. The determination of sterility in Drosophila males without a complete Y-chromosome. Am. Nat., 72:42-52. The innervation of setae in Drosophila. Genetics, 23: 172-73. During which stage in the nuclear cycle do the genes produce their effects in the cytoplasm? Am. Nat., 72:350-57. Control of a species-difference by means of a difference in an in- ductor. Nature, 142: 158. Biology. In: An Orientation in Science, ed. C. W. Watkeys et al., pp. 256-332. New York: McGraw-Hill. 1939 Somatic crossing-over and somatic translocations. Am. Nat., 73:95 -96. With E. Hadorn. The relation between the color of testes and vase efferentia in Drosophila. Genetics, 24: 162-79. 1940 Recent work on the relation between genes and developmental processes. Growth, Suppl. 1:19-36. The prospective significance of imaginal discs in Drosophila. J. Mor- phol., 67: 107-22. On dependent growth and form of the testes in various species of Drosophila. Collect. Net., 15: 1-4. Growth in vitro of the testis of Drosophila. Growth, 4:337-82. 1941 The growth of testes in Drosophila. I. The relation between vas deferens and testis within various species. i. Exp. Zool., 87: 113- 58. The growth of testes in Drosophila. II. The nature of interspecific differences. i. Exp. Zool., 87:159-80.

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464 BIOGRAPHICAL MEMOIRS With A. Brasted. An analysis of the expression of the mutant "en- ~railed" in Drosophila melanogaster. (genetics. 26: 347-73. ~ . 1942 With L. Birmingham. Boundaries of differentiation of cephalic imaginal discs in Drosophila. ]. Exp. Zool., 91:345-63. 1943 The Hardy-Weinberg law. Science, 97: 137-38. Genic action as studied by means of the effects of different doses and combinations of alleles. Genetics, 28:441-75. Some new types of class experiments with Drosophila at the Univer- sity of Rochester. Ward's Nat. Sci. Bull., 16:67. Effect of environmental differences, and a "Lamarckian" experi- ment. Ward's Nat. Sci. Bull., 16:93. Cumulative and competitive action of alleles and their bearing on the position eject. Genetics, 28:92-93. With E. W. Schaeffer. On primary attributes of alleles in Drosophila melanogastex Proc. Natl. Acad. Sci. USA, 29:351-61. With E. SchaeEer. On wild-type iso-alleles in Drosophila melanogaster. Proc. Natl. Acad. Sci. USA, 29:361-67. 1944 The journey not the goal. Sci. Mon., 58 :96 -100. With E. W. Schaeffer and W. P. Spencer. The genetic basis of dif- ferences between two species of Drosophila. Am. Nat., 78: 183- 88. Peace time research in war time. Science, 99:278-80. A study of race. l. Hered., 35:314-16. With G. Heidenthal. Materials for the study of the position effect of normal and mutant genes. Proc. Natl. Acad. Sci. USA, 30: 197-205. 1945 With D. R. Charles. The Rhesus gene and the effect of consan- guinity. Science, 101 :305 - 7. A letter a college president might write. Bull. Am. Assoc. Univ. Prof., 31:1. Review of Genetics, by E. Altenburg. Science, 102:514-15.

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CURT STERN 465 1946 With E. W. Schaeffer and G. Heidenthal. A comparison between the position effects of normal and mutant alleles. Proc. Natl. Acad. Sci. USA, 32:26-33. Review of Heredity and its Variability, by T. D. Lysenko, Am. Nat., 80:241-43. With G. LeClerc. Occurrence of mitotic crossing-over without meiotic crossing-over. Science, 103:553 -54. With R. MacKnight and M. Kodani. The phenotypes of hemizy- gotes of position alleles and of heterozygotes between alleles in normal and translocated position. Genetics, 31:598-619. With M. Kodani. An invisible chromosome. Science, 104:620-21. 1947 The skin color of children from white by near-white marriages. I. Hered., 38:233-34. 1948 With W. Spencer. Experiments to test the validity of the linear R- dose/mutation frequency relation in Drosophila at low dosage. Genetics, 33:43-74. With E. Caspari. The influence of chronic irradiation with gamma rays at low dosages on the mutation rate in Drosophila melano- gaster. Genetics, 33:75-95. Negative heterosis and decreased effectiveness of alleles in heter- ozygotes. Genetics, 33:215 -19. With Trudy Enders. The frequencies of twins, relative to age of mothers, in American populations. Genetics, 33:263-72. With E. Novitski. The viability of individuals heterozygous for re- cessive lethals. Science, 108: 538-39. The effects of changes in quantity, combination and position of genes. Science, 108:615 -21. 1949 Gene and character. In: Genetics, Paleontology and Evolution, ed. G. L. tepsen, pp. 13-22. Princeton: Princeton University Press. With D. E. Uphoff. The genetic effects of low intensity irradiation. Science, 109:609-10. Selection and eugenics. Science, 110: 1-8.

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466 BIOGRAPHICAL MEMOIRS Principles of Human Genetics. San Francisco: W. H. Freeman and Co. 617 pp. 1950 Anomalies of genetic origin. Pediatrics, 5:324-28. Reply to Bernhard Stern. Science, 111:698. Genetic aspects of sterility. Fertil. Steril., 1 :407-14. Boveri and the early days of genetics. Nature, 166:466. With E. R. Sherwood. The migration of testis sheath cells in Dro- sophila virilis. In: Moderne Biology. Festschrift fur Hans Nachtsheim, ed. H. Gruneberg, pp. 236-40. Berlin: F. W. Peters. 1951 Concluding remarks of the chairman. Cold Spring Harbor Symp. Quant. Biol., 15 :409-12. With A. Hannah. The sex combs in gynanders of Drosophila melan- ogaster. Port. Acta Biol., Ser. A:798-812. With S. T. Fung. The seriation of fourth chromosome loci in Dro- sophila melanogaster. Proc. Natl. Acad. Sci. USA, 37:403-4. Probleme der menschliche Erbforschung. Mitt. Naturforsch. Ges. Bern, 9. Problems of radiobiology with emphasis on radiation genetics. In: Proc. Annul Biol. Colloq., pp. 7-16. Corvallis: Oregon State Uni- versity Press. The genetic future of man. In: Proc. Annul Biol. Colloq., pp.43-51. Corvallis: Oregon State University Press. 1952 Man's genetic future. Sci. Am., 186:68-74. Genetics and the world today. In: The Scientists Look at Our World, ed. }. M. Fogg, pp. 61-82. Philadelphia: University of Pennsyl- vania Press. (Also in: Smithson. Year Annul Rep., Smithson. Inst. for 1953, pp. 263-276.) 1953 With G. Carson, M. Kinst, E. Novitski, and D. Uphoff. The viability of heterozygotes for lethals. Genetics, 37:413-49. With G. Belar. Race crossing in paradise? }. Hered., 49: 154-55.

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C U RT S T E RN 467 The geneticist's analysis of the material and the means of evolution. Sci. Mon., 77: 190-97. Model estimates of the frequency of white and near-white segre- gants in the American Negro. Acta Genet. Stat. Med., 4:281- 98. 1954 Listings of unpublished articles. Science, 119:221. George W. Beadle. Science, 119:229 -30. Two or three bristles. Am. Sci., 42:213-47 Needed research. Eugen. Q., 1:161-65. The biology of the Negro. Sci. Am., 191:81-85. The facts of life. Popul. Stud., London, 8: 188-91. Guest editorial: One scientist speaks up. Science, 120:5A. Genes and developmental patterns. In: Caryologia, Proc. 9th Int. Congr. Gen., Part I, pp. 355-69. Firenze: Industria Tipografic . . . . ~ 1orentlua. . 1955 A professor's days. Calif. Mon., 65: 15 -17. Gene action. In: Analysis of Development, ed. B. Willier, P. Weiss, and V. Hamburger, pp. 151-69. Philadelphia: W. B. Saunders Co. With M. Kodani. Studies on the position effect at the cubitus inter- ruptus locus of Drosophila melanogastex Genetics, 40:333-73. Qualitative aspects of the population problem. Science, 121:683- 86. Grundlagen der menschlichen Erblehre. Gottingen: Muster-Schmidt Verlag. 560 pp. 1956 With A. Hannah. Stability of iso-alleles. Nature, 177:42. Die Bedeutung der "Wirbelsaulenmethode nach Kuhne" fur den Vaterschaftsausschluss. Ein Gutachten. Acta Genet. Stat. Med., 6:92-102. Genetics in the atomic age. Eugen. Q., 3:131-38. Hereditary factors affecting adoption. In: A Study of Adoption Prac- tice, vol. 2, by M. Shapiro, pp. 47-58. New York: Child Welfare League of America. The genetic control of developmental competence and morpho

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468 BIOGRAPHICAL MEMOIRS genetic tissue interactions in genetic mosaics. Roux Arch 149: 1-25. Genetic mechanisms in the localized initiation of differentiation. Cold Spring Harbor Symp. Quant. Biol., 21:375-81. The role of genes in differentiation. In: Cytologia, Proc. 9th Int. Congr. Gen., pp. 70-72. 1957 A note on the detection of differential effects of mutagens. I. Ge- net., 55:276 - 79. The scope of genetics. Proc. Natl. Acad. Sci. USA, 43:744-49. Research needed. In: Proceedings World Population Conference, 1954, vol. 6, pp. 665-73. Geneva: World Health Organization. Problems of Y-chromosome inheritance in man. spa. i. Hum. Ge- net., 2:27 - 29. With A. Hannah-Alava. The sexcombs in males and intersexes of Drosophila melanogaster. J. Exp. Zool., 134:533-56. The problem of complete Y-linkage in man. Am. i. Hum. Genet., 9: 147-66. With D. L. Swanson. The control of the ocellar bristle by the scute locus in Drosophila melanogaster. ]. Fac. Sci. Hokkaido Univ., Ser. 6, 13:303-7. O die dum professor. Agros Lisbon, 40:254-59. With G. L. Walls, Jr. The Cunier pedigree of "color blindness." Am. }. Hum. Genet., 9: 249-73. On a case of lethal ichthyosis in Hiroshima. Jpn. T. Hum. Genet., 2:87-88. 1 ~ 1958 With L. S. Penrose. Reconsideration of the Lambert Pedigree (ichthyosis histrix gravior). Ann. Hum. Genet., 22:258-83. The ratio of monozygotic to dizygotic affected twins and the fre- quencies of affected twins in unselected data. Acta Genet. Med. Gemellol., 7:313 -20. Radiation and population genetics. In: Radiation Biology and Medi-- cine, ed. W. D. Claus, pp. 206-28. Reading, Mass.: Addison- Wesley. Selection for subthreshold differences and the origin of pseudo- exogeneous adaptations. Am. Nat., 92:313 -16.

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CURT STERN 469 In memoriam, Richard Goldschmidt. Experientia, 14:307-8. Richard Goldschmidt, biologist. Science, 128: 1070. Richard Goldschmidt, 12.4.1878 (Frankfurt a.M.) his 25.4.1958. Naturwissenschaften, 45:429-31. The nucleus and somatic cell variation. I. Cell Comp. Physiol., 52:1-34. Acceptance of the honorary degree from McGill University for Kihara, Penrose and Stern. Proc.10th Int. Congr. Genet.,1: 14 15. (Also in: Iden, 12: 27.) Le Daltonisme lie au chromosome X, a-t-il une localisation unique ou double? Expose de deux theories. i. Genet. Hum., 7:302-7. 1959 The chromosomes of man. I. Med. Educ., 34:301-14. (Also Hum. Genet., 12:141.) in: J. Variation and hereditary transmission. Proc. Am. Philos. Soc., 103: 183-89. Colour-blindness in Klinefelter's syndrome. Nature, 183: 1452-53. Use of the term "superfemale." Lancet (Dec. 121:1088. Die Geschlechtsbestimmung. Triangle, 4:131-35. Genetics in the atomic age. In: Protection in Diagnostic Radiology, ed. B. Sonnenblick, pp. 256-65. New Brunswick, N.J.: Rutgers University Press. 1960 The determination of sex. Triangle, 4:131-35. Brain damage in the infant-genetic aspects. Calif. Med., 92:21- 24. O. Vogt and the terms "penetrance" and "expressivity." Am. i. Hum. Genet., 12:141. mosaic of Drosophila consisting of 1X, 2X and 3X tissue and its probable origin by mitotic non-disjunction. Nature, 186:179- 80. Dosage compensation-development of a concept and new facts. (Fifth Huskins Memorial Lecture.) Can. I. Genet. Cytol.,2: 105- 18. Mechanism of meiotic non-disjunction in man. Nature, 187:805.

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470 BIOGRAPHICAL MEMOIRS Principles of Human Genetics, 2d ed. San Francisco: W. H. Freeman and Co. 753 pp. 1961 With E. Sherwood. A search for maternally influenced sex-ratio in Drosophila melanogastex DIS, 35:96. Review of Die philosophischen Grundlagen der Naturwissenschaften, by M. Hartman. Science, 133:697. Letter to the editor: Mongolism. Lancet, 1 (April 8~:775. With S. Sarkar, A. Banerjee, and P. Bhattacharjee. A contribution to the genetics of hypertrichosis of the ear rims. Am. i. Hum. Genet., 13:214-23. 1962 Wilhelm Weinberg. Genetics, 47: 1-5. Wilhelm Weinberg zur hundert jahrigen Wiederkehr seines Ge- burtsjahres. Z. Konstit-Lehre, 36:374-82. In praise of diversity. (Presidential Address, American Society of Zoologists.) Am. Zool., 2:575-79. With E. Sherwood. Can primordial germ cells of the genotype XXY produce functional sperm? DIS, 36:118. The origin of the sternopleural sclerite in Drosophila. Am. Zool., 2:562. With S. Sarkar, A. Banerjee, and P. Bhattacharjee. Letter: Inheri- tance of hairy pinnae. Am. i. Hum. Genet., 14:434-35. 1963 The genetics of sex determination in man. 2d Int. Congr. Hum. Genet., Symp. 3, 1962. Am. J. Med., 34:715 -20. With C. Tokunaga, C. Grisseau, and F. Gottlieb. The cell lineage of the sternopleura in Drosophila melanogaster. Dev. Biol.,7:365- 78. The concentration of rare genes. In: The Genetics of Migrant and Isolate Populations, ed. E. Goldschmidt, pp. 243-49. Baltimore: Williams & Wilkins. Cytogenetica cloveka. Acta Chir. Orthop. Traumatol. Cech., 30: 385-94. 1964 With K. B. DeOme and J. A. ~enkins. William Ernest Castle, 1867 1962. In Memoriam. Univ. of California, pp. 11-13.

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CURT STERN 471 With E. Stern. Theodor Boveri (translation of an article by F. Baltzer). Science, 144:809-15. With A. S. Mukherjee. Aspects of developmental genetics of the legs of Drosophila. Proc. Natl. Acad. Sci. India, Sect. B., 34: 19- 26. Adventures in dermatological genetics. (Fourth Herman Beerman Lecture.) I. Invest. Dermatol., 43:217-22. Synthesis. In: Genetics Today, Proc. lOth Int. Congr. Genet., Septem- ber 1963, pp. 221-26. Elmsford, N.Y.: Pergamon Press. Review of Kurze Geschichte der Genetik his zur Wiederentdeckung der Vererbungsregeln Gregor Mendels, by Hans Stubbe. Isis, 55:377- 79. 1965 With C. Tokunaga. The developmental autonomy of extra sex combs in Drosophila melanogaster. Dev. Biol., 11 :50 - 81. With A. S. Mukherjee. The effect of sexcombless in genetic mosaics of Drosophila melanogaster. Z. Vererbungsl., 96:36 -48. Thoughts on research. Science, 148: 772-73. Entwicklung und die Genetik von Mustern. Naturwissenschaften, 52:357-65. Mendel and human genetics. Proc. Am. Philos. Soc., 109:216 -26. With C. Tokunaga. Hairy ear rims in Japanese. J. Hered., 56:218- 19. 1966 Pigmentation mosaicism in intersexes of Drosophila. Rev. Suisse Zool., 73:339-55. Population genetics (statement by the moderator). Proc. World Pop. Conf. Belgrade, 1965, 1:114-24. New York: United Na- t~ons. The Origin of Genetics, ed. E. Sherwood and C. Stern. San Francisco: W. H. Freeman and Co. 179 pp. 1967 Richard Benedict Goldschmidt. In: Biographical Memoirs, vol. 39, pp. 141-92. New York: Columbia University for the National Academy of Sciences. With C. Tokunaga. Nonautonomy in differentiation of pattern- determining genes in Drosophila. I. The sex comb of eyeless- dominant. Proc. Natl. Acad. Sci. USA, 57:658-64.

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472 BIOGRAPHICAL MEMOIRS Review of The Cell in Development and Inheritance, by E. B. Wilson. Isis, 58:125 - 27. The genetic resources of man. In: Natural Resources: Quality and Quantity, ed. S. van Ciriacy Wantrup and J. Parsons, pp. 35-51. Berkeley: University of California Press. Mendel's memorabilia. Science, 157: 1119. Genes and people. 3d Int. Congr. Hum. Genet., Chicago, Septem- ber, 1966. Perspect. Biol. Med., 10:500-523. Some general aspects of human genetics. (Fred L. Adair Award Address to the American Gynecological Society, May 1967.) Am. I. Obstet. Gynecol., 99:604-14. 1968 Questions and answers: Color of child of "mixed marriage." I. Am. Med. Assoc., 204:200. With C. Tokunaga. Autonomous pleiotropy in Drosophila. Proc. Natl. Acad. Sci. USA, 60:1252-59. Genetic Mosaics and Other Essays. Cambridge: Harvard University Press. 185 pp. With A. Schaefer and O. I. Ukpe. Hairy pinnae in Nigeria. l. Hered., 59:174-78. An early investigation on the genetics of catalase content in blood. Jpn. I. Hum. Genet., 13: 181-82. 1969 Somatic recombination within the white locus of Drosophila melan- ogaster. Genetics, 62:573 -81. Richard Benedict Goldschmidt. Perspect. Biol. Med., 12: 179-203. Gene expression in genetic mosaics. Genetics, 61: 199-211. A note on the facsimile reproduction of Mendel's manuscript on "Versuche uber Pflanzen-hybriden." Folia Mendeliana, 4:41- 43. With C. Tokunaga. Determination of bristle direction in Drosophila. Dev. Biol., 20:411-25. 1970 Model estimates of the number of gene pairs involved in pigmen- tation variability of the Negro-American. Hum. Hered., 20: 165-68. The continuity of genetics. Dacdalus, 99:882-908.

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CURT STERN 1971 473 With C. Tokunaga. On cell lethals in Drosophila. Proc. Natl. Acad. Sci. USA, 68: 329-31. From crossing-over to developmental genetics. In: Stadler Genetics Symposia 1-2, ed. G. Kimbar and G. Redei, pp. 21-28. Colum- bia: University of Missouri Press. The place of genetics in medicine. Ann. Intern. Med., 75:623-29. 1972 Summary of session. In: Perspectives in Cytogenetics. The Next Decade, ed. S. Wright, B. Crandall, and L. Boyer, pp. 50-52. Spring- field, Ill.: Charles C Thomas. 1973 Principles of Human Genetics, 3d ed. San Francisco: W. H. Freeman and Co. 891 pp. A geneticist's journey. In: Chromosomes and Cancer, ed. I. German, pp. xiii-xxv. New York: John Wiley. High points in human genetics. Am. Biol. Teach., 37:144-49. 1974 Presidential address: "The domain of genetics." In: Proceedings, XIII Congress of Genetics, University of California, Berkeley, 1973. Genetics, 78:21-33.