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Passionate Geneticist, Teacher, and Encyclopaedist

Jim Birchler

George P. Rédei (1921—2008) (Photo courtesy of ASPB)

Passionate Geneticist, Teacher, and Encyclopaedist

The proof of being a classic is that nobody reads it anymore in original, although it is frequently cited and even more frequently misquoted. Some of the perceptions proceed like funeral corteges from author to author; everybody sidesteps the statements and adds some new twists to the myths. Arabidopsis is not an exception to the general situation. There is an essential difference, however, between knowing things and knowing about them.
—G. P. Rédei, 1992

Over the 50 years of his scientific career, George P. Rédei retained his passionate love of genetics and Arabidopsis, his favorite research tool, which found its way into the pantheon of genetic models with fully sequenced genomes after a period of neglect. In shaping the future of plant science, Rédei was one of the most influential geneticists. With his outstanding research papers and recurrent reviews, he persuaded the younger generation of molecular biologists to focus their research on Arabidopsis in the 1980s.

George Rédei was born in Vienna (June 14, 1921, to Kálmán and Margit Rédei), but he grew up in Hungary. His father was an agronomist of a large estate and published studies on hybrid vigor in maize. After finishing high school, the “Benedictine Real Gymnasium” in Pápa, Rédei registered at the College of Agriculture at Magyaróvár, the alma mater of his father, in 1941. However, World War II interrupted his studies. In 1943 he was deported as a forced laborer to a work camp. Luckily he escaped, but upon wandering home, he found that the war took away his beloved parents and brother. As he remained completely alone, he cultivated the family’s land to earn money to finish his agronomy studies and survive the post-war famine. The new Stalinist regime confiscated his farm. In 1951 he moved to Budapest, where he worked in a nursery with the eminent geneticist Vilmos Teichmann. In 1953 he received a student award to prepare his PhD thesis at the Institute of Genetics of Hungarian Academy under the supervision of Barna Györffy. In 1949 Györffy received the highest state award, the Kossuth Prize, but was soon denunciated as he and most of his students, including Rédei, resisted implementing the official Lysenkoist doctrines in their research. Thus, Rédei’s PhD work had to focus on practical breeding problems such as studying the inheritance of tomato fruit weight, hybridization of rye with Triticum turgidum and T. durum, and establishment of a tissue culture system for embryo rescue of endosperm deficient Triticale hybrids. In 1953 he married Magdolna M. Rédei (“Magdi”), who helped him in completing his PhD and with many experiments throughout his life.

In 1955 the political control of research weakened and Rédei began to search for a better experimental tool that would facilitate his biochemical genetic studies. He read publications by Friedrich Laibach, who managed to regenerate flax embryos. However, Laibach favored Arabidopsis for genetic studies, as this plant had only five pairs of chromosomes, a short life cycle, and high seed yield and could be easily crossed and cultivated in vitro. Furthermore, Laibach’s collaborator, Erna Reinholz, succeeded in inducing mutations by X-radiation in Arabidopsis in 1947. Rédei obtained some seed from Laibach and soon shared his opinion that Arabidopsis was indeed well suited as a model for plant genetic studies. In 1956, after Rédei moved to the Agricultural Research Institute in Martonvásár and started X-ray mutagenesis with Arabidopsis, an uprising broke out in the country. As it was not clear how “Morganists” (those recognizing the importance of T. H. Morgan’s genetic work on fruit flies) would be treated by the new political system, Rédei decided to leave his homeland after the Soviet army circled Budapest in November 1956.

In January 1957 Rédei received permission to emigrate from an Austrian refugee camp to the United States, where he obtained a job as assistant professor in the Department of Genetics (later Agronomy) at the University of Missouri—Columbia. In the famous “headquarters” of maize and cytogenetics, the old Curtis Hall, he inherited the former laboratory of Barbara McClintock along with L. J. Stadler’s old X-ray machine and greenhouse. From 1957 to 1967, with his first students, Y. Hirono and S. L. Li, Rédei established a powerful genetic system for Arabidopsis. His most significant scholarly contributions from this period were the isolation of over 200 mutations at five genetic loci controlling thiamine biosynthesis, which represented the first examples of auxotrophic mutations in higher plants; the demonstration of allelic complementation in Arabidopsis; the characterization of unstable mutations affecting the pyrimidine pathway; the identification of nuclear loci that enhance the mutability of extranuclear genomes; the development of genetic techniques for isolation of mutant homoplastidic lines; the identification of mutations affecting flowering time (e.g., ld, co, gi), megaspore differentiation, male transmission, chlorophyll biosynthesis, and leaf development; and the construction of the first rough linkage map for three of five Arabidopsis chromosomes.

In the 1960s Rédei, alone in his appreciation of the extraordinary value of Arabidopsis in the United States, contacted Laibach’s lab in Frankfurt and also developed interactions with all other Arabidopsis researchers in Europe and Australia. With G. Röbbelen, Rédei edited a regular newsletter, the Arabidopsis Information Service, and organized the first Arabidopsis conference in 1965 to celebrate Laibach’s retirement and the establishment of a common stock collection. Despite publishing in Science, Nature, Genetics, and other high-ranking journals, the years following 1969 were very bitter for Rédei. The NSF declined funding for Arabidopsis research, whereas USDA and DOE shared the view of those breeders who believed that, despite some differences in the cost of cultivation, plant genetics could be done perfectly with maize, tomato, or wheat without a need for an Arabidopsis model. Rédei was promoted to professor in 1969 and, because of lack of funding, invested part of his salary into his experiments. Because this was not much, he also began working on theoretical aspects of genetics and writing university textbooks. One of these, the handbook Genetics (Macmillan, 1982), was translated into Chinese and Hungarian and used by thousands of students. Rédei’s best friend, Ernie Sears, the outstanding wheat cytogeneticist, encouraged him to concentrate on other important occupations as well, such as the organization of 14 Stadler Genetics Symposia, which brought many excellent geneticists to the University of Missouri for more than three decades. In 1974 Rédei published several important papers on the mathematical theory of planning mutagenesis experiments and a fabulous compilation on the history of genetics titled “Steps in the Evolution of Genetic Concepts” (Biol. Zentralblatt 93:385–424). In the late 1970s he participated in a large-scale international mutagen-testing project and demonstrated the extreme sensitivity of Arabidopsis mutagen assay system. In 1970 and 1975 he published two seminal reviews on the genetics and biology of Arabidopsis and its value as a model system, respectively (Bibliographia Genet. 20:1–151; Annu. Rev. Genet. 9:111–27). The latter attracted particular attention because it purged a major scientific forgery of L. Ledoux and associates. These researchers reported in Nature that they achieved a correction of Rédei’s thiamine auxotrophy mutations to wild type by soaking Arabidopsis seeds in a solution of transducing lambda phage DNA carrying the E. coli thiamine locus. These and related reports gave the hope for funding agencies and breeders that transformation of plants could be easily and rapidly implemented. However, a simple segregation experiment performed by Rédei eliminated these hopes, unfortunately together with the hope of better support for Arabidopsis genetics.

After the second Arabidopsis meeting in 1976, most of Rédei’s European colleagues, including A. Müller, G. Röbbelen, J. H. van der Veen, and others, shifted to work with other plant species. During these dark years for Arabidopsis, Rédei was inspired by and closely followed the development of new plant transformation technologies, which used the transferred DNA (T-DNA) of Agrobacterium Ti and Ri plasmids and led ultimately to the birth of plant molecular biology. Jeff Schell, a pioneer of these developments, invited Rédei as guest professor to the Max-Planck Institute of Plant Breeding Research (Cologne, Germany) in 1986, where he established basic methods of tissue culture, regeneration, and Agrobacterium-mediated transformation of Arabidopsis. He was 66 when Arabidopsis finally entered the long-awaited triumphal path at the third International Arabidopsis meeting, organized by Chris Somerville at Michigan State University in 1987. Rédei et al. and three other laboratories reported on Agrobacterium-mediated transformation of Arabidopsis at the meeting. This opened the way to molecular studies of Arabidopsis gene functions, first using recombinant DNA constructs and later by high-frequency Agrobacterium T-DNA–mediated insertion mutagenesis. Elliot Meyerowitz, who followed Rédei’s footsteps by proposing general acceptance of Arabidopsis as a plant model in genetics (Annu. Rev. Genet. [1987] 21:91–111), reported at the same meeting that the Arabidopsis genome size was in the range of 100–120Mb, the smallest known in dicots. Thus, it became evident that Arabidopsis was also an excellent choice for genome sequencing. The long-awaited success of Arabidopsis and an unexpected smaller grant from NSF revitalized Rédei, who asked the university authorities to prolong his active research time. He retired in 1991, but continued to collaborate as professor emeritus with the MPI in Cologne on the analysis of T-DNA–tagged insertion mutants, generation of new mutant collections, study of the T-DNA integration mechanism, and confirmation of hormonal functions of brassinosteroids. By 2006 Rédei had published 250 research papers, notes, letters, book chapters, and books. By his retirement he had deposited 6,728 specimens at the Arabidopsis Biological Resource Center.

For recognition of his life work, Rédei was elected foreign member of the Hungarian Academy of Sciences in 1990. The Arabidopsis community dedicated to him the first practical training course in Arabidopsis Molecular Genetics (1992, Cologne, supported by the European Molecular Biology Organization and European Commission) and the first handbook on Methods in Arabidopsis Research (1992). In 2004 the University of Missouri dedicated a section of its local Plant Growth Facilities to him. From 1994 to 1997 Rédei returned nearly every year to Hungary for a few months as visiting professor, teaching at the Institute of Genetics in the Eötvös Lóránd University of Budapest. In 1996 he taught as Fulbright lecturer and edited a book titled Genetic Bases of Physiological Responses to Environmental Effects in Plants at Pannon Agricultural University at Keszthely.

In 1996 Rédei learned practical computing from his daughter, Mari. In addition to communicating from his home in Columbia, Mo., with his beloved grandchildren, Paige, Grace, and Anne in Nashville, Tenn., Rédei used the computer to initiate a huge project based on his extremely wide and deep knowledge in genetics and other fields of biology. He collected more than 18,000 genetic concepts, 600 illustrations, and thousands of references to books and databases in a Genetics Manual, and then doubled these in a revised encyclopedic dictionary printed by Wiley in 2003. After five years of exhausting work, he finished correcting the third edition of his extraordinary lexicon, titled Encyclopedia of Genetics, Genomics, Proteomics, and Informatics, this past summer, which was published by Springer in two volumes covering 3,335 pages. In August he said, “It seems to me unrealistic to continue the encyclopedia because a single person cannot track any more properly the new developments in genomics, proteomics, and systems biology. The problem is that the encyclopedia gave me some meaningful work, without which I feel useless.” Soon after, his health declined unexpectedly fast and he died on November 10, 2008, in Nashville.

With the passing of George Rédei, we lost a legendary geneticist and teacher who devoted his life to promoting the progress of plant genetics and, in particular, Arabidopsis research—a task that should remain our responsibility.

Contributed by Csaba Koncz
Max-Planck Institute for Plant
Breeding Research

A Formative Encounter with George Rédei

In the spring of 1978 we were living in Paris, thinking about what we should do next. The writings of people like Paul Ehrlich and Norman Borlaug had impressed on us the related facts that human population growth was creating environmental problems and technology could help avert some aspects of the problem by intensifying production, thereby reducing the demand for undeveloped land. Therefore, we were thinking about how we might participate in bringing new technology to plant improvement. We spent our mornings reading in the beautiful, small library at the Institute Pierre and Marie Curie, where Antoine Danchin had graciously permitted us to visit, and our afternoons in the cafés of Paris discussing what we had read.

Just before arriving in Paris we had been playing around with a gift of EcoRI from Howard Goodman and had read a paper from Mary Dell Chilton and collaborators proposing that Agrobacterium tumefaciens transferred DNA into the host genome during pathogenesis. We inferred that it was going to be possible to transform plants using the Ti plasmid and began thinking about what that meant. One insight we had was that in order to exploit the emerging tools of molecular biology, plant biologists needed a better model organism for molecular genetics; something diploid and small with a rapid life cycle and a low DNA content that was suited to laboratory work. This led us to a compelling article written the year before by George Rédei for Annual Reviews of Genetics, extolling the virtues of Arabidopsis for plant genetics. The fact that George had been able to identify auxotrophic mutants and had good numbers for the frequency with which such mutants could be isolated was quite exciting for us because we took it as an indication that it was going to be relatively easy to isolate mutations in a wide variety of genes. During the next several months we tracked down and read as many of George’s papers as we could obtain access to in the Rothschild and Institute Pasteur libraries. George’s papers were extremely helpful because they had good numbers about frequencies of genetic events and detailed methods sections. On the basis of his papers, we were able to envision how to do experiments without having ever seen the plant; we were able to carry out gedankenexperiments that led us to the ideas we eventually carried to Bill Ogren’s lab at Illinois.

After our money ran out, we returned to the University of Alberta, where we had both been graduate students, to wait for our visas and to write fellowship proposals to work with Bill. The graduate students had funding to invite a speaker and we convinced them to invite George Rédei and to let us be his hosts.We cannot remember how it happened, but somehow George ended up visiting for almost three days. Presumably in our enthusiasm for his work, we talked him into coming for an extended visit. We could not get enough faculty members to meet with him to fill three days, so, to our delight, we had the better part of several days with George, during which we talked through everything he knew about Arabidopsis in particular and plant genetics in general. We talked about the details of every manipulation, such as crossing the minuscule flowers, and we tested our ideas against his view of reality and feasibility. It was really a wonderful and memorable occasion—for us. We think George also enjoyed it. He was passionate about his work and had probably not experienced a level of interest in his work comparable to ours. It probably helped that we had read all of his papers that were available in the libraries we had access to. He was certainly very patient with us, and after he returned home he showered us with resources, such as the marker lines he had developed for mapping and our first M2 population, which allowed us to begin screening for mutants immediately. We think back to that time now as our micro-postdoc with George.

We never had the opportunity to spend a lot of time with George after that brief but formative encounter. However, from our correspondence and chance encounters at meetings, we had the impression that he took considerable satisfaction in seeing the explosion of Arabidopsis research and in having directly helped many of the early members of the Arabidopsis community get started. The use of the Columbia wild type (named by George for Columbia, Mo.) as the standard accession for Arabidopsis research memorializes George’s founding contributions.

Chris Somerville and Shauna Somerville
University of California at Berkeley