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First published online October 23, 2003; 10.1104/pp.103.028530 Plant Physiology 133:1732-1746 (2003) © 2003 American Society of Plant Biologists Tissue-Specific and Developmentally Regulated Expression of a Cluster of Tandemly Arrayed Cell Wall-Associated Kinase-Like Kinase Genes in Arabidopsis1Department of Biology, San Francisco State University, 1600 Holloway Avenue, San Francisco, California 94132
The Arabidopsis cell wall-associated kinase (WAK) and WAK-like kinase (WAKL) family of receptor-like kinase genes encodes transmembrane proteins with a cytoplasmic serine/threonine kinase domain and an extracellular region containing epidermal growth factor-like repeats. Previous studies have suggested that some WAK members are involved in plant defense and heavy metal responses, whereas others are required for cell elongation and plant development. The WAK/WAKL gene family consists of 26 members in Arabidopsis and can be divided into four groups. Here, we describe the characterization of group 2 members that are composed of a cluster of seven tandemly arrayed WAKL genes. The predicted WAKL proteins are highly similar in their cytoplasmic region but are more divergent in their predicted extracellular ligand-binding region. WAKL7 encodes a truncated WAKL isoform that is predicted to be secreted from the cytoplasm. Ratios of nonsynonymous to synonymous substitutions suggest that the extracellular region is subject to diversifying selection. Comparison of the WAKL and WAK gene clusters suggests that they arose independently. Protein gel-blot and immunolocalization analyses suggest that WAKL6 is associated with the cell wall. Histochemical analyses of WAKL promoters fused with the
Article, publication date, and citation information can be found at www.plantphysiol.org/cgi/doi/10.1104/pp.103.028530. 1 This work was supported by the National Science Foundation (grant no. MCB 9985135). Additional support for this work was provided by the National Institutes of Health (grant nos. MBRS SCORE 5 SO6 GM52533 and RIMI P20 RR11805). 2 Present address: Department of Horticulture, College of Agricultural Sciences, Penn State University, 103 Tyson Building, University Park, PA 16802–4200. 3 Present address: Department of Plant and Microbial Biology, University of California, Berkeley, CA 94720. * Corresponding author; e-mail zhe{at}sfsu.edu; fax 415–338–2295. Received June 12, 2003; returned for revision July 22, 2003; accepted September 2, 2003. This article has been cited by other articles:
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-glucuronidase reporter gene have shown that the expressions of WAKL members are developmentally regulated and tissue specific. Unlike WAK members whose expressions were found predominately in green tissues, WAKL genes are highly expressed in roots and flowers. The expression of WAKL5 and WAKL7 can be induced by wounding stress and by the salicylic acid analog 2,6-dichloroisonicotinic acid in an nonexpressor of pathogenesis-related gene 1-dependent manner, suggesting that they, like some WAK members, are wound inducible and can be defined as pathogenesis-related genes. 
