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OCR for page 353
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.
OCR for page 356
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.
OCR for page 357
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-
OCR for page 358
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-
OCR for page 359
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.
OCR for page 360
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.
OCR for page 361
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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Representative terms from entire chapter:
spring harbor