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First published online December 23, 2004; 10.1104/pp.104.054312 Plant Physiology 137:176-189 (2005) © 2005 American Society of Plant Biologists Annotations and Functional Analyses of the Rice WRKY Gene Superfamily Reveal Positive and Negative Regulators of Abscisic Acid Signaling in Aleurone Cells1,[w]Department of Biological Sciences, University of Nevada, Las Vegas, Nevada 89154
The WRKY proteins are a superfamily of regulators that control diverse developmental and physiological processes. This family was believed to be plant specific until the recent identification of WRKY genes in nonphotosynthetic eukaryotes. We have undertaken a comprehensive computational analysis of the rice (Oryza sativa) genomic sequences and predicted the structures of 81 OsWRKY genes, 48 of which are supported by full-length cDNA sequences. Eleven OsWRKY proteins contain two conserved WRKY domains, while the rest have only one. Phylogenetic analyses of the WRKY domain sequences provide support for the hypothesis that gene duplication of single- and two-domain WRKY genes, and loss of the WRKY domain, occurred in the evolutionary history of this gene family in rice. The phylogeny deduced from the WRKY domain peptide sequences is further supported by the position and phase of the intron in the regions encoding the WRKY domains. Analyses for chromosomal distributions reveal that 26% of the predicted OsWRKY genes are located on chromosome 1. Among the dozen genes tested, OsWRKY24, -51, -71, and -72 are induced by abscisic acid (ABA) in aleurone cells. Using a transient expression system, we have demonstrated that OsWRKY24 and -45 repress ABA induction of the HVA22 promoter-
1 This work was supported by the U.S. Department of Agriculture (grant no. 02–35301–12066), by the National Institutes of Health (Biomedical Research Infrastructure Network seed grant no. P20 RR16464), and by the University of Las Vegas, Nevada (start-up funds to Q.J.S.). X.Z. was supported by a National Science Foundation Experimental Program to Stimulate Competitive Research (EPSCoR), Integrative Approaches to Abiotic Stress (EPS–0132556), graduate assistantship and Z.-L.Z. by a National Science Foundation EPSCoR Advanced Computing in Environmental Sciences postdoctoral fellowship. 2 These authors contributed equally to the paper. 3 Present address: Department of Plant and Microbial Biology, University of California, Berkeley, CA 94720 and Plant Gene Expression Center, U.S. Department of Agriculture, Albany, CA 94710. [w] The online version of this article contains Web-only data. Article, publication date, and citation information can be found at www.plantphysiol.org/cgi/doi/10.1104/pp.104.054312. * Corresponding author; e-mail jeffery.shen{at}ccmail.nevada.edu; fax 702–895–3956. Received October 4, 2004; returned for revision November 11, 2004; accepted November 15, 2004. This article has been cited by other articles:
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-glucuronidase construct, while OsWRKY72 and -77 synergistically interact with ABA to activate this reporter construct. This study provides a solid base for functional genomics studies of this important superfamily of regulatory genes in monocotyledonous plants and reveals a novel function for WRKY genes, i.e. mediating plant responses to ABA. 
