Perspectives

Copyright Ó 2006 by the Genetics Society of America Perspectives Anecdotal, Historical and Critical Commentaries on Genetics Edited by James F. Crow and William F. Dove Edward Novitski: Drosophila Virtuoso James F. Crow,*,1 Dan Lindsley† and John Lucchesi‡ *Genetics Laboratory, University of Wisconsin, Madison, Wisconsin 53706, †Section of Cell and Developmental Biology, University of California, San Diego, California 92093, and ‡Department of Biology, Emory University, Atlanta, Georgia 30322 DWARD Novitski, 1918–2006, was the acknowledged attempts to extricate himself make an amusing story. E master of that special art of manipulating chro- The second part of Ed’s book is the story of his own life. mosomes during what Lucchesi (1994) called ‘‘the age The third is his account of the rift between Alfred of Drosophila chromosome mechanics.’’ Following the Sturtevant and Theodosius Dobzhansky. Novitski had Sturtevant tradition, his guiding principle was to derive the unique vantage point of having been successively a as much information as possible from breeding experi- student of each of them, and he is clearly in Sturtevant’s ments with minimum use of direct cytological examina- corner. The fourth part is titled ‘‘The Pleasure of Find- tion. Nobody could perform this kind of chromosome ing Things Out.’’ It recounts some of his most interest- manipulation as well as Ed and he relished new chal- ing intellectual challenges. The book is as idiosyncratic lenges. (Novitski’s closest friends and relatives, especially as Ed himself, a mixture of deep science, anecdotes, in later years, called him Eddie, a name he seems to have intellectual depth, and whimsy. preferred. Those, such as ourselves, whose acquaintance extends over many years, knew him as Ed, which we shall A BRIEF BIOGRAPHY adopt here.) He continued this kind of work long after the development of microbial and molecular genetics Edward Novitski was born in Wilkes-Barre, Pennsylvania, had carried the field in new directions. on July 24, 1918. He had an early interest in science, Another side of Novitski was a love of pranks and prac- leading him to chemistry experiments and building a tical jokes, which were indeed clever, sometimes di- radio receiver. He also had an interest in collecting ani- abolically so. He reached what must be the pinnacle of mals, especially snakes, a practice he engaged in after achievement in this rarified atmosphere, for his repu- slipping out of highly unpleasant church services. Sig- tation was such that pranks that he never carried out nificantly for both their later careers, he and E. B. Lewis were attributed to him nonetheless. He admired Richard attended the same high school and were the leaders of a Feynman, a physics genius who also enjoyed such things biology club. One day, while perusing the journal Sci- as safe cracking, bongo drums, and practical jokes. Both ence, Ed Lewis noticed an ad for Drosophila cultures, so men took great pleasure in ‘‘finding things out’’ and most of the club’s small treasury was used to get some shared highly creative minds and a love of pranks. stocks. It was the beginning of two illustrious Drosophila Fortunately, Ed completed an autobiographical mem- careers. Novitski carried on a correspondence with the oir not long before his death (Novitski 2005). He was Purdue faculty member S. A. Rifenburgh, who had fur- unusual; so, naturally, the book is unusual. It has four nished the flies for the biological supply house. With main sections. The first is ‘‘Fun and Games,’’ a series of Rifenburgh’s help, he got a scholarship to Purdue, with- anecdotes and practical jokes. The first prank is his most out which college would have been unattainable for him. famous, perpetrated on Herschel Roman. Ed manipu- While in high school Ed discovered a mutation, heldout lated the seminar clock to run slowly so that Herschel, (Novitski and Rifenburgh 1938), shown to be an after finishing his talk, seemed to have an embarrass- allele of a gene that was subsequently named decapenta- ingly long period of time remaining to be filled. His plegic. This is a key regulatory gene in dorsal–ventral patterning of the embryo. He also found the mutation, later called asteroid, which he sent to Ed Lewis; this 1Corresponding author: Department of Genetics, University of Wisconsin, mutation marked the beginning of Lewis’s famed ca- Madison, WI 53706. E-mail: [email protected] reer (Crow and Bender 2004). Novitski carried on an Genetics 174: 549–553 (October 2006) 550 J. F. Crow, D. Lindsley and J. Lucchesi Edward Novitski. Courtesy of Charles Novitski. extensive correspondence with C. B. Bridges, remark- Novitski 1941). This article uncovered an error by able for the interest and professional respect that Bridges Dobzhansky, a failure to identify one chromosome arm. showed for this promising unknown scientist. No fewer During this period and later, Novitski became increas- than 36 letters were exchanged. ingly conscious of numerous Dobzhansky errors. An- In high school, Ed learned Latin over the summer other example is the ‘‘sex ratio’’ phenomenon, which and entered the second-year class in the fall. At Purdue Dobzhansky had interpreted as an extra chromosome he soon discovered that he could get credit for college replication. This was later shown to involve no unusual courses simply by learning the material on his own and processes; rather, it involved the nonfunctioning of some taking examinations. In this way, starting with calculus, meiotic products (Novitski et al. 1965). Novitski’s dis- he completed his undergraduate studies at Purdue in illusionment with Dobzhansky and his admiration of 2 years. He influenced his friend Lewis to adopt the Sturtevant argues that, although the differences were same strategy. always underplayed, he must have received considerable At Bridges’ suggestion, he applied to and was ac- satisfaction from discovering the errors, as was probably cepted by Caltech, entering graduate school in the fall true of Sturtevant as well. of 1938. Alas, Bridges was terminally ill at this time, so Novitski received his Ph.D. in 1942. While in what might have been an important scientific collabo- Dobzhansky’s group, he met Esther Rudkin whom he ration never materialized. Novitski began graduate work later married while he was in military service. His service as a student of Dobzhansky, irradiating Drosophila pseudo- in the Army Air Force lasted 3 years, from 1942 to 1945. obscura to produce inversions and using these to isolate During this time the Air Force received a new, state of lethal mutations from natural populations. These le- the art device, which was installed in bombers flying out thals were to be tested for allelism, a heroic task since of England. It failed to function as described, but char- the number of tests goes up with the square of the num- acteristically Ed was able to identify the problem and ber of mutations. Ed soon became disillusioned, think- restore its function. For this, he received a commenda- ing that the data had minimal value for distinguishing tion from his commanding officer. After his discharge among alternative ideas and that the work could, as he from the Air Force, he spent 2 years in the laboratory said, be carried out as easily by a well-trained chimpan- of Curt Stern at the University of Rochester, the first as zee. He finally gave up, destroyed the stocks, and was a Guggenheim fellow and the second on an Atomic promptly sacked as a teaching assistant. Energy project. Then came a year at the University of After some uncertainty, he was finally awarded a fel- Missouri in the laboratory of A. B. Griffen and 2 years at lowship with Sturtevant. He was now on the path that Caltech, again with Sturtevant. In 1951 he joined the he would follow the rest of his life. Together they pub- faculty at the University of Missouri where he stayed lished an article showing the chromosomal homologies until 1956. He then moved to the Biology Division of of various Drosophila species, as evidenced by the lo- the Oak Ridge National Laboratory as head of the cation of corresponding mutations (Sturtevant and Drosophila genetics group. Finally, in 1958, he became Perspectives 551 Professor of Biology at the University of Oregon where, same gamete. These could be perpetuated in subse- except for sojourns in Zurich, Canberra, and Leiden, he quent generations by selecting nondisjunction prod- remained until his retirement in 1983. He died June 29, ucts. An additional crossover could then reconstitute 2006, in his 88th year. the original long inversion sequence with the mutant allele in it. Finally, a double crossover within the larger inversion transferred the desired allele to the normal RESEARCH uninverted chromosome, and the Greens had their de- A central theme in Novitski’s work was the remaking sired allele in a manageable condition. The procedure of the Drosophila genome by attaching chromosome and Muller’s reaction to it are described in Ed’s book arms in novel ways. Attached X chromosomes were well (Novitski 2005, pp. 135ff). known, but there were five other ways in which two X Novitski’s graduate work involved species other than chromosomes could be attached to a single centromere. Drosophila melanogaster (Novitski 1946), but after that Ed succeeded in constructing all five (Novitski 1954a). time he worked exclusively with that species. He dem- These chromosomes were used to analyze various as- onstrated early that low-temperature shocks produced pects of segregation and recombination. A particularly the desemination of mated females, rendering them neat procedure was employed to join the X and Y. At- ‘‘virgins’’ (Novitski and Rush 1948). He demonstrated taching parts of the X to parts of the Y had already been that recombination between some pairs of structurally done, but Novitski and Dan Lindsley succeeded, by a heterozygous chromosomes produces heteromorphic procedure that was both logical and ingenious, in at- dyads from which the smaller element is preferentially taching the entire X and Y chromosomes (Lindsley included in the functional egg nucleus (Novitski 1951).

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