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GEORGE STREISINGER December27, 1927-SeptemberS, 1984 BY FRANKLIN W. STAHL GEORGE STREISINGER WAS a leading contributor to the genetics of the T-even bacterial viruses, culminating in his demonstration anct rationalization of the circular link- age map. During the infancy of molecular biology, he pro- vicied crucial links between genetics and biochemistry with his demonstration of the consequences of frameshift muta- tions. He identifier! and cleveloped zebra fish as a system for the genetic analysis of vertebrate development. PERSONAL HISTORY George Streisinger was born in Budapest, Hungary, on December 27, 1927. To escape Nazi persecution, he en c} his parents left Budapest for New York when he was ten years oIcI. He attendecI public schools, graduating from the Bronx High School of Science in 1944. During his high school clays, George filled his home with salamanders, spiders, and snakes and colIaboratec! with Theoclosius Dhobzansky on studies of courtship in Drosophila. George's first three scien- tific papers (1944, 1946, 1948), two of them sole authored, resulted! from that precocious enterprise. George obtainer! a B.S. degree in genetics from Cornell University in 1950 en c] a Ph.D. in genetics from the Univer- 353

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354 BIOGRAPHICAL MEMOIRS sity of Illinois in 1953. His postcloctoral studies were at the California Institute of Technology from 1953 to 1956. In 1949 George married Lotte Sielman. They had two children, Lisa and Cory, born in 1955 ant! 1956, respec- tively. George's career was influenced by Salvadore E. Luria, with whom he took his Ph.D., and, more so, by Max Delbruck and lean Weigle, with whom he stuclied as a postdoctoral fellow. George influenced a number of other collaborators and friends, and it is plausible that they influenced him, too. They inclucle Sidney Brenner, Seymour Benzer, Matt Meselson, Robert Edgar, Jan Drake, and myself. PROFESSIONAL HISTORY George was one of many who usecl the Tow-cost School of Agriculture at Ithaca to gain access to a high-quality Cornell University undergracluate education. As always, he put his - - . . . . . etiorts where his Interests were, For instance, while at Cornell' he met and married Lotte Sielman, a refugee from Munich. Furthermore, he spent an academic year on a farm working off a provisional status earnest by his neglect of required ROTC anchor physical education courses. (This require- ment could have been worked off in the summer, but George spent all his summers at Coicl Spring Harbor.) Following his graduation from Cornell, George uncler- took graduate studies in the genetics of T-even coliphage with S. E. Luria in the Bacteriology Department of the Uni- versity of TIlinois. The phages T2 and T4, while cTistinguish- able, are obviously related and tract been shown by Delbruck to recombine with each other to give hybrid phages of vary- ing degrees of viability. George iclentifiecT single genes re- sponsible for the differences between T2 and T4 in W sensitivity ~ ~ 956, ~ ), host range, and serological specificity ( 1956,2), respectively. These studies revealed phenotypic , 1 red ~

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GEORGE STREISINGER 355 mixing, in which a phage with the host-range genotype of one phage type was found in a particle whose phenotype was that of the other (1956,21. George based his Ph.D. the- sis on these experiments. When they were published in 1956, they had a profounc! impact on viral biology. Upon obtaining his degree, George went off to CalTech to pioneer the study of plant somatic cell genetics. He couldn't make those stucties fly, however, and while still at CalTech he returned to the genetics of phage. With lean Weigle, he undertook further studies on T2 x T4 hybrids, which led to the discovery of DNA moctif~cation (by glucosylation) . With Naomi Franklin, George undertook a fine-structure recombination study of the host-range (hi locus of T2, after the fashion of Benzer's studies on the rIl locus of T4. How- ever, whereas, the rII protein escaped (letection, it was prob- able that the protein product of the h gene, T2's tail fibers, would be tractable. Although the connection between gene and protein was not made, these studies foreshaclowed George's later work (described below) on the T4 lysozyme gene. With Victor Bruce (1960), George showed that the known genetic markers of T4 could be arrayed on a single linkage group. This simplification of the previously held view of three linkage groups was an essential step in the coales- cence of genetics with emerging physical data on T4 DNA. After his postdoc at CalTech, George took a position at Cold Spring Harbor. (His science knew no boundaries and his publications no timetable. Adct to that the free flow of personnel ant} ideas among phage labs in those clays, and the chronology of my recitation occasionally becomes a bit vague. ) George took leave from Cold Spring Harbor to spencl a year at the MRC in Cambridge, EnglancI, with Sidney Brenner.

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356 BIOGRAPHICAL MEMOIRS In collaboration with other visiting Americans, they initi- ated studies on T4 proteins with an eye to clecoding the relationship between DNA and protein (1959~. One prod- uct of that work was the identification of T4's endolysin (lysozyme) as a promising object for such studies. At Cold Spring Harbor and the University of Oregon's Institute of Molecular Biology, where George took a posi- tion in 1960, he cleveloped the methods for selecting mu- tants, revertants, and recombinants in the T4e gene, which encodes the phage enclolysin. His first applications of this know-how was in a demonstration that phage containing 5- bromouracil have a high mutation rate when they are al- lowed to grow in medium that is free of 5-bromouracil. The second application was the demonstration, at the level of amino acid sequence, that frameshift mutations in the e gene really do shift the translation reacting frame, as pre- clicted by Francis Crick's hypothesis of a commaless triplet code (1966,1,21. The first in viva codon assignments re- sultec! from this work. The third application of George's mastery of the e gene was an analysis of the roles played by amino acid sequence in determining protein stability. In 1992, impelled by the methods of modern genetics, this work remains a major activity of several chemists and physi- cists at Eugene who study protein folding as well as stability. The physical studies on phage T4 seemed to indicate that each T4 particle contained one Watson-Crick duplex DNA molecule. Contemporaneous genetic studies, however, argued that regions of heterozygosity in T4 were 4-strandec! anct that these regions were variable in position. This para- clox was resolved by Meselson and Streisinger's suggestion that the chromosomes of T4 are circularly permuted an(l~ terminally reclunclant. After George moved to Eugene, he tested predictions of that notion, most notably the preclic- tion that the unitary linkage map of T4 be circular. Since E.

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GEORGE STREISINGER . 357 cold was the only creature previously known to have a circu- lar map, George's demonstration with Bob Edgar (1964) was important in establishing the concept of wiclespreacl circularity among microbes. Subsequent papers support George's notions of terminal reclundancy of a permuted chromosome whose length is cleterminecl by the amount of DNA that can be fit into a phage head (1965, 19671. Frameshift mutations are proflavin inducible, and George's Interests extended to the mechanism of that mutation in- cluction. He offered a proposal that has played a central role in our understanding of the origin of duplications and cleletions (19721. Like many phage workers, George eventually set his sights on more complex systems. Working initially without stu- clents (because it wouicl not be fair to risk their careers), George developed the methods for the mutation en cl ge- netic analysis of zebra fish (1981;1983,l,21. It was his aim to make the fish as tractable as phage so that it could be used for a genetic analysis of the vertebrate nervous system. The degree to which he succeedecI can be juciged by his master- piece on the development of the pigmented retina (1989) and by the many laboratories that are now exploiting this little Indian import to unravel other mysteries of vertebrate development. This work, better even than his work on the code, illustrates his imagination and courage. His career was reaching its zenith when he cried of a heart attack dur- ing his final exam in a scuba diving class. George's research contributions ensured his position in worIc! science. His position in Oregon was ensured equally by his extraordinary contributions as a teacher, a politically involved citizen, a chef, and a warmly sociable frienc! and colleague. As a teacher, George was unbelievably dedicate<] to the students. His dedication was backed by energy (he was al-

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BIOGRAPHICAL MEMOIRS 358 ways available), imagination (he conscripted a dance class to illustrate protein synthesis), and, of course, creep uncler- standing. The University of Oregon recognized his teach- ing with a prestigious award. George was politically active both on ant! off campus. He spent a major part of his first two years in Eugene organiz- ing grass-roots resistance to the Vietnam War and legisla- tive opposition to John Kennecly's civil defense program. He played a central role in the successful effort to restrict the use of potentially mutagenic herbicides in Douglas fir reforestation. This extracurricular activity informed his pub- lished work (1983,31. He lee! and won a battle to exclude secret war department research from the University of Or- egon campus. An invitation to dinner at the Streisinger home was never refused, because in the 1960s there were no restaurants in Eugene that collie come close to the cuisine offered there. A barbecue was sometimes a suckling pig, at other times a giant Chinook salmon. Memorable winter meals were tracti- tional Hungarian. Breakfast was for children. It featured crepes poured to resemble animals anc! server! with choco- late syrup. Magic tricks sometimes followed. When George was chairman of our Biology Department, he combatted the paperwork blues by unsuccessfully breed- ing pheasants (the foxes got them) en c] by successfully train- ing to be a goat jucige (he was in great demand at county fairs throughout the West). George's family continues to have its impact on Eugene ant! Oregon. Eugene's well-known Saturday Market was founded in the early 1960s by George's wife, Lotte, a pot- ter, who currently serves the community as an art aciminis- trator. His daughter, Lisa, founded a company in Portland that administers health care systems. Cory, his younger claugh-

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GEORGE STREISINGER 359 ter, served as lawyer for Governor Neil Goldschmidt and is currently lawyer for the Port of Portland. George's impact on the University of Oregon has been symbolized by the naming of a beautiful research building for cell and molecular biology. George's impact on his col- leagues in the Institute of Molecular Biology has been marked by an annual lecture, quickly recognized as both a scientific and a social highlight of our community. LOTTE STRElSINGER AND AARON NOVICK made important contributions to this memorial.

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360 BIOGRAPHICAL MEMOIRS S E L E C T E D BIBLIOGRAPHY 1944 With T. Dobzhansky. Experiments on sexual isolation in Drosophila II. Geographic strains of Drosophila prosaltans. Proc. Natl. Acad. Sci. U.S.A. 30:340-45. 1946 The cardini species group of the genus Drosophila. J. N.Y. Entomol. Soc. 54:105-13. 1948 Experiments on sexual isolation in Drosophila IX. Behavior of males with etherized females. Evolution 2:187-88. 1956 The genetic control of ultraviolet sensitivity levels in bacterio- phages T2 and T4. Virology 2:377-87. Phenotypic mixing of host range and serological specificity in bac- teriophages T2 and T4. Virology 2:388-98. 1959 With S. Brenner et al. Structural components of bacteriophage. [. Mol.Biol.1:281-92. 1960 With V. Bruce. Linkage of genetic markers in phages T2 and T4. Genetics 45:1289-96. 1964 With R. S. Edgar and G. H. Denhardt. Chromosome structure in phage T4, I. Circularity of the linkage map. Proc. Natl. A cad. Sci. U.S.A. 51:775-79. 1965 With T. Sechaud et al. Chromosome structure in phage T4, II. Ter- minal redundancy and heterozygosis. Proc. Natl. Acad. Sci. U.S.A. 54:1333-39.

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GE O RGE STREI SIN GER 1966 361 With E. Terzaghi et al. Change of a sequence of amino acids in phage T4 lysozyme by acridine-induced mutations. Proc. Natl. Acad. Sci. U.S.A. 56:500-507. With others. Frameshift mutations and the genetic code. Cold Spring Harbor Symp. Quant. Biol. 31:77-84. 1967 With I. Emrich and M. M. Stahl. Chromosome structure in phage T4, III. Terminal redundancy and length determination. Proc. Natl. Acad. Sci. U. S.A. 57:292-95. 1972 With Y. Okada et al. Molecular basis of a mutational hot spot in the lysozyme gene of bacteriophage T4. Nature 236:338-41. 1981 With others. Production of clones of homozygous diploid zebra fish (Brachydanio rerio) . Nature 291 :293-96. 1983 With S. Chakrabati et al. Frequency of 7-ray induced specific-locus and recessive lethal mutation in mature germ cells of the zebrafish (Brachydanio rerio). Genetics 103:109-23. With C. Walker. Induction of mutations by 7-rays in pregonial germ cells of zebrafish embryos. Genetics 103:125-36. Extrapolation from species to species and from various cell types in assessing risks from chemical mutagens. Mutat. Res. 114:93-105. 1989 With others. Clonal origins of cells in the pigmented retina of the zebrafish eye. Dev. Biol. 131:60-69. (In the publication, this work, which was performed at the University of Oregon's Institute of Molecular Biology, is unaccountably attributed to the University of Utah School of Medicine.)

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