Edward D. Garber, professor emeritus of ecology and evolution at the University of Chicago, died 9 Oct 2004. He was 86. Dr Garber was a polymath geneticist who published more than 150 papers and two books during his long and productive career; Garber’s methods were unusual in the variety of systems he used as experimental models. His research encompassed studies on higher plants (sorghum, peas, Collinsia), lower plants (algae, liverworts), animals (mice, carp, Homo sapiens), bacteria (Erwinia, Pseudomonas, Xanthomonas) and fungi (Aspergillus, Colletotrichum, Fusarium, Penicillium, Microbotryum). The mold systems became his favorite models, and he spent the last three decades of his life focused almost exclusively on filamentous fungi.
Garber was born and raised in New York City; he grew up in a coldwater flat in Manhattan’s Lower East Side and was educated in the city’sPUBLIC schools. With the aid of a scholarship he attended Cornell University, where his zeal for both genetics and fencing was kindled. He was a member of Cornell’s fencing team and was an enthusiast most of his life. Garber graduated in 1940 with a bachelor of science degree in botany and earned his master’s degree in genetics at the University of Minnesota in 1942. He served in the Army during World War II. While stationed in Charleston, South Carolina, he attended a dance given for the soldiers, where he met a quiet, 19 y old southerner named Rosalie Kirshstein, whom he later married. Not long after the wedding he was sent to officers’ candidate school in New Orleans. In 1945, Garber was sent to the European theater. After V-E Day he was reassigned to the Pacific, where he earned the Philippine Liberation Ribbon. Upon his discharge in 1946 the Garbers moved to the West Coast. Garber re-entered graduate school and received his doctorate in genetics from the University of California at Berkeley. His 1949 dissertation on the genetics of sorghum received the John Belling Prize.
Ed’s first job was with the Office of Naval Research in Oakland, California, where he studied variation and drug resistance in Pasteurella pestis (now Yersinia pestis), causative agent of plague. The research required work with an animal colony, and in a satisfying example of Pasteur’s dictum that “chance favors the prepared mind” Garber discovered the first sex-linked gene in mice. During his time at the laboratory his son Joel was born. The University of Chicago recruited him to the faculty of the Botany Department in 1953. Shortly after moving to Chicago the Garbers’ first daughter Martha Ann was born. Garber was well suited to academic life and rose easily through the ranks, becoming associate professor in 1958 and professor in 1961. Janie, the last of the three Garber children, was born in 1959.
During his many decades at the University of Chicago Garber taught genetics to several generations of undergraduates and supervised 11 master’s theses and 20 doctoral dissertations. In the 1980s he helped develop a human genetics curriculum for Chicago’sPUBLIC school system and he was a consultant on the design of the sickle cell anemia exhibit at Chicago’s Museum of Science and Industry. Garber was a first-rate teacher who delivered well organized lectures, in terse sentences laced with humor. He had a staccato, Walter Winchell style of speech, a New York accent and a fair amount of New York attitude. Today, when so much of science instruction consists of flashing PowerPoint images onto a screen, we treasure the memory of Garber’s solving genetics problems on a chalkboard, lecturing at his fast clip and engaging students in after class conversations. His teaching style showed how intellect, personality and passion inspires young minds. He took creative liberties in class, inventing the imaginary organism “darting asterisk” with alleles “ept” and “inept” to illustrate how genetic analysis is symbolic and universal. In 1982 he received the Quantrell Award for Excellence in Undergraduate Teaching, a much-coveted honor among University of Chicago faculty.
Ed was passionate about research. He reveled in the power of classical genetic analysis. He studied many organisms, but two systems in particular are associated with his legacy, the plant genus Collinsia and the smut species Microbotryum violaceum (formerly Ustilago violacea). Garber recognized that scientists needed a small, easily cultivable and convenient model for genetics research on higher plants. In the 1950s and 1960s he focused on the dicot genus Collinsia, a small herb then classified in the Scrophulariaceae (snapdragon family), now classified in an enlarged group called the Plantaginaceae. These annuals have quadripartite flowers that display a variety of colors based on anthocyanin pigments, which aid biochemical studies on the inheritance of color. All 21 Collinsia species studied in the Garber laboratory had seven chromosomes. It was possible to make many interspecific hybrids and in a few cases to isolate triploids. Garber’s group performed elegant cytogenetic analysis of amphidiploids, extracted pigments and enzymes for biochemical analysis and envisioned that this genus would become a kind of plant Drosophila; he was trying to develop an Arabidopsis type model. It is hard to know whether he would have succeeded in making Collinsia what Arabidpsis has become if he had not succumbed to the allure of the even greater experimental convenience of microbial systems. In the 1960s, Garber increasingly turned his attention to filamentous fungi, describing the parasexual cycle in Aspergillus fumigatus, Fusarium oxysporum, Penicillium digitatum, P. expansum and P. italicum; analyzing the relationship between nutrition and phytopathogenicity; and using enzyme electrophoresis as a taxonomic and genetic tool for mycological studies.
JWB’s relationship with the Garber lab began when she entered the Botany Department 1963. She had planned to become a plant cytogeneticist and Garber was the only cytogeneticist in the department. Without any lab rotations he automatically became her mentor, and guided her transition from green plants to filamentous fungi.
By the 1970s a smut then called Ustilago violacea, now named Microbotryum violaceum, became the model of choice for the Garber laboratory. This basidiomycete is the causative agent of anther smut disease in susceptible species of the Caryophyllaceae (carnation or pink family). Teliospores of Microbotryum are diploid. They can be collected from smutted anthers and germinated on laboratory medium where they undergo meiosis. The resultant tetrad of basidiospores produces four clones of sporidia that each form a yeast-like colony through budding. When a conjugation bridge forms between appropriate strains, the resulting dikaryon initiates the infection cycle. The species has a parasexual cycle, and complementation tests can be performed in diploid sporidia. The Garber lab collected smutted anthers from plants in the field as well as herbarium specimens. Strains with white, yellow, pumpkin and pink sporidial colony colors provided a rich phenotypic resource for studying carotene biosynthesis and became the basis for transmission genetic systems in this smut.
Michael Baird was a graduate student from 1973–1978, during the early years of smut research. At first he started a project on grasshoppers but “soon realized it was more difficult to keep them alive than to study their genetics.” He then moved to a project on Ustilago violacea. Writing many years later he spoke fondly of Garber’s tutelage in the basics of being a good geneticist. Some of Dr Baird’s fondest memories however involved coffee breaks. Most afternoons Ed came to see whether Michael needed to stretch his legs. “This inevitably meant first a stop in the bookstore where Ed would read the cartoons in Punch and get a good laugh,” Baird said. “This would carry us through the rest of the day.” All of Garber’s former students valued his easy mix of science and sociability. He particularly enjoyed introducing undergraduates to the pleasures of laboratory research and helped guide many of them into medical or scientific careers. Garber’s open-mindedness and wry sense of humor was fortunate for the undergraduate (JAK) who decided to wait until after he had started working with colored colonies before mentioning that he was colorblind.
Ed had an insight into biology that bordered on the mystical. When pink sporidial colonies turned out not to be due to a carotenoid, he guessed that the coloration had to be caused by a heme-containing pigment, such as cytochrome c. And that is exactly what it turned out to be. In later years his investment in a microbial system using colored markers proved prescient when sectored colonies gave genetic evidence for the presence of transposable elements.
Garber also had exceptional writing and editorial skills. Garber was a master of elegant scientific prose. His decades of laboratory productivity resulted in numerous tautly written papers. Manfred Ruddat, a longtime colleague, said that he would not have been surprised if Garber someday had managed to write a paper that was so beautifully concise that it would “fit on a postage stamp.” Garber was fond of using Roman numerals to connect his work on a given experimental model, with more than 25 papers on Collinsia and a similar number on Microbotryum (most of them published when the organism was still called Ustilago). Further he wrote a short text titled Cytogenetics: An Introduction for McGraw Hill in 1972 and edited a collection of papers titled Genetic Perspectives in Biology and Medicine for the University of Chicago Press in 1985. In 1972 he became editor in chief of the Botanical Gazette, helped supervise the revitalization of the journal and its name change to International Journal of Plant Sciences and served as co-editor in chief with Manfred Ruddat 1992–2000.
Ed was not fond of travel and only rarely attended scientific meetings. However in the 1970s, under the auspices of the United States-Israel Binational Science Foundation, the Garbers spent a spring and two summers at the Hebrew University in Jerusalem. Projects included a study of prawns that would flourish in brackish water ponds, breeding hybrid carp, and the use of yeast as a human protein supplement.
In 1968 the Botany Department was merged with several other units to become the Department of Biology. Garber faced mandatory retirement in 1988 but stayed on as professor emeritus of ecology and evolution after yet another reorganization of biology disciplines. Instead of finding a quiet place to settle back and bask in justifiable pride after retirement he continued to do research, teach undergraduate genetics and edit the International Journal of Plant Sciences. In 1992 the Medical and Biological Sciences Alumni Association awarded him the Gold Key Award.
Ed maintained offices and laboratories in both the former botany building (now the Erman Biology Center) and in Barnes Laboratory. Both buildings were within walking distance of home. “Ed loved the lab,” Rosalie Garber said. “Given his druthers he would have lived at the lab and just dropped in at home now and then.” In the late 1990s the university needed their home for expansion of its hospital complex. They sold their property in 1999 and moved to Skokie, Illinois, to be near their daughters and grandchildren. The move coincided with escalating health problems for Garber. His body became increasingly weak, but his mind showed undiminished acuity. It became difficult for him to travel to the University of Chicago campus so he worked from home. At the time of his death he was working on a paper in which he tried to unite the gene of classical eukaryotic transmission genetics with the gene concept of modern molecular analysis. In 2004 he made the decision to stop dialysis. His wife of 61 years, his three children and his grandchildren, Becky and Matt, all were nearby when he passed away quietly at the Hospice of the North Shore.
Fig. 1. Edward Garber at Botany Pond, University of Chicago campus.
