2002 GSA Honors and Awards

The 2002 Thomas Hunt Morgan Medal

The Genetics Society of America annually honors members who have made outstanding contributions to genetics. The Thomas Hunt Morgan Medal recognizes a lifetime contribution to the science of genetics. The Genetics Society of America Medal recognizes particularly outstanding contributions to the science of genetics within the past 15 years. The George W. Beadle Medal recognizes distinguished service to the field of genetics and the community of geneticists. We are pleased to announce the 2002 awards.

IRA Herskowitz is the richly deserving recipient of the 2002 Thomas Hunt Morgan Medal, the highest honor awarded by the Genetics Society of America. This award recognizes 33 years of outstanding achievement since the publication of his first research paper in 1970. We are sad to note that Ira Herskowitz succumbed to pancreatic cancer on April 28, 2003.

Ira's research career started during his undergraduate days at Cal Tech, where he characterizes his experience as "I was 20 minutes late for my first class and I never caught up." Fortunately for genetics, he found his way into a research project on bacteriophage with Robert Edgar and Jonathan King, where he developed a passion for experimental genetics. Perhaps more importantly he gained a deep appreciation of experimental contexts in which intellectual rigor could be matched by experimental care, as this, he believed, was the best way to train scientists.

Imprinted by the gospel of microbial genetics, Ira found his way to graduate school at MIT, where he joined Ethan Signer's lab for graduate work. There he developed an interest in the biology of phage and its interaction with the physiology of its host. Lambda appealed to Ira deeply because of the alternate fates, lysis or lysogeny, that were controlled by genes of and of Escherichia coli. This study of alternative fates was to be a theme that continued throughout his career.

After completing his Ph.D. in 1971, Ira enjoyed a brief stint with yeast and Cold Spring Harbor Laboratory during a postdoctoral year with David Botstein before beginning as an Assistant Professor in the Institute of Molecular Biology at the University of Oregon. The institute at that time was an odd mixture of geneticists and biophysicists. Two of his colleagues who made a deep impression on Ira were George Streisinger and Frank Stahl, top figures in microbial genetics and molecular biology.

Ira's approach to science was a refreshing change of pace for the graduate students first attracted by more biophysical interests. Indeed his first six graduate students were walk-ons who gave up their original interests to follow Ira's research agenda and style. As the seventh member to join the lab, I remember Ira's uncanny ability to take any result and help you see how it could be developed into a much bigger story with great significance. There was more to Ira's appeal than intricate hypotheses on chalkboards, toothpicks, and Petri plates. His passion for both music and ping pong enlivened many occasions, and the track meets in Eugene were always fun and occasionally dry. Joining his lab was the easiest decision that most of us had made.

It's hard to say with certainty what any great scientist's principal strength is. With Ira, certainly the ability to choose compelling research problems was one of them. He described his strategy as one in which he looked for situations in which there are two or more observations that all parties agree upon, yet the observations are mutually incompatible. This philosophy led Ira directly into a central mystery in the biology of the Saccharomyces yeast. Everyone agreed that the MATa and MAT alleles of the mating-type locus were codominant. Yet everyone also agreed that Don Hawthorne had a deletion that converted an cell into an a cell. Clearly, the conversion of one codominant allele into another challenges all normal conventions in genetic thinking.

This insight led to an amazingly beautiful set of experiments, initiated by Jim Hicks and Jeff Strathern, which proposed and proved that yeast switch mating types by way of transposable cassettes. This body of work may well represent the last great problem in biology that was fundamentally resolved purely by classical genetic analysis.

In 1979, Ira spent a sabbatical at UC San Francisco; in 1981 he joined the faculty at UCSF, where he remained. From 1981 to 2001, he headed the Division of Genetics and from 1982 to 1995, served first as Vice Chairman and then Chairman of the Department of Biochemistry and Biophysics. This early period in Ira's career at UCSF was heavily influenced by the pervasive strength in cell biology at UCSF and the experimental facility that yeast lent his lab in tackling these issues with genetics. The leading example from this period was his lab's contribution to understanding the cascade of events that enable two yeast cells of opposite mating types to engage in preconjugal communication, coordinating their cell cycles to facilitate mating. This process is now understood from the receptors, through the G protein, through a cascade of MAP kinases, to the transcription factors that are the targets of the signal. By looking for mutants in the classic way that could still respond to the ligand but did not arrest the cell cycle, Ira and his colleagues worked out the coordination between signaling and cell cycle progression.

Other examples include a detailed dissection of how the expression of the HO gene is regulated. His lab has been one of the principle contributors to our understanding of how the HO endonuclease is expressed only in G1, and then only in mother cells and not in their daughters. Understanding the genetic basis of asymmetry at HO expression was particularly gratifying to Ira, both as a capstone to his lab's discovery of the asymmetry in mating-type switching, but also as the first well understood example of a problem fundamental to developmental biology: how new cell types can be programmed to appear at the right place at the right time.

More recent studies include an analysis of why the budding pattern of haploid and diploid cells is different, which has blossomed into other aspects of cell polarization, and studies of the role of chromatin components and chromatin remodeling factors on gene expression. As in all cases, Ira came to these interests not by reading the papers of others. Instead he simply allowed the mutants isolated in his own lab to reveal the stories they had to tell.

His most recent work continued an interest in signaling of other types, including pathogenesis in Ustilago, and moved into more central medical problems. He had begun a study of prion autocatalysis in yeast and became interested in the significance of polymorphisms in human genes that encode molecules involved in drug transport and metabolism.

Prior to the Morgan Medal, Ira had already received substantial recognition from his peers including election to the National Academy of Science in 1986, a MacArthur Foundation Fellowship in 1987, the Genetics Society of America Medal in 1988, and election to the Institute of Medicine in 2002. He served the field at the highest levels including service on the NIH study section, where he had great influence on the articles that appear in this journal. He performed great service on many different editorial boards, including five years as Associate Editor of Genetics, on the Scientific Review Board of the Howard Hughes Institute, and as a jury member for the Albert Lasker Medical Research Board.

I suspect that these and the many other awards and honors he received were of secondary importance to Ira at best. He led a career in which excellence in teaching was always a goal and in which teaching people how to ask the right question is of paramount importance. There can be no better sign of his success than the thousands of students whose appreciation of genetics was born in the classes that Ira taught and the perhaps unprecedented success that Ira's students have had in their own independent careers following their training in his lab. As a community, we warmly applaud the selection of Ira Herskowitz as the recipient of the 2002 Thomas Hunt Morgan Medal of the Genetics Society of America.

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