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First published online June 15, 2006; 10.1104/pp.106.077388 Plant Physiology 141:1363-1375 (2006) © 2006 American Society of Plant Biologists Characterization of the Class IV Homeodomain-Leucine Zipper Gene Family in Arabidopsis1,[W]Division of Biological Sciences, Graduate School of Science, Hokkaido University, Sapporo 060–0810, Japan (M.N., H.K., M.A., K.T.Y.); Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Tokyo 113–0033, Japan (N.Y., Y.K.); and Division of Bioscience, Graduate School of Natural Science and Technology, Okayama University, Okayama 700–8530, Japan (T.T.)
The Arabidopsis (Arabidopsis thaliana) genome contains 16 genes belonging to the class IV homeodomain-Leucine zipper gene family. These include GLABRA2, ANTHOCYANINLESS2, FWA, ARABIDOPSIS THALIANA MERISTEM LAYER1 (ATML1), and PROTODERMAL FACTOR2 (PDF2). Our previous study revealed that atml1 pdf2 double mutants have severe defects in the shoot epidermal cell differentiation. Here, we have characterized additional members of this gene family, which we designated HOMEODOMAIN GLABROUS1 (HDG1) through HDG12. Analyses of transgenic Arabidopsis plants carrying the gene-specific promoter fused to the bacterial
1 This work was supported by a Grant-in-Aid for Scientific Research on Priority Areas (15031201, 17027021 to T.T.) from the Ministry of Education, Culture, Sports, Science, and Technology of Japan. 2 Present address: Department of Integrated Genetics, National Institute of Genetics, Mishima, Shizuoka 411–8540, Japan. 3 Present address: Department of Botany, Graduate School of Science, Kyoto University, Kyoto 606–8502, Japan. The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy described in the Instructions for Authors (www.plantphysiol.org) is: Taku Takahashi (perfect{at}cc.okayama-u.ac.jp). [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.106.077388. * Corresponding author; e-mail perfect{at}cc.okayama-u.ac.jp; fax 81–86–251–7876. Received January 21, 2006; returned for revision June 4, 2006; accepted June 6, 2006. This article has been cited by other articles:
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-glucuronidase reporter gene revealed that some of the promoters have high activities in the epidermal layer of the shoot apical meristem and developing shoot organs, while others are temporarily active during reproductive organ development. Expression profiles of highly conserved paralogous gene pairs within the family were found to be not necessarily overlapping. Analyses of T-DNA insertion mutants of these HDG genes revealed that all mutants except hdg11 alleles exhibit no abnormal phenotypes. hdg11 mutants show excess branching of the trichome. This phenotype is enhanced in hdg11 hdg12 double mutants. Double mutants were constructed for other paralogous gene pairs and genes within the same subfamily. However, novel phenotypes were observed only for hdg3 atml1 and hdg3 pdf2 mutants that both exhibited defects in cotyledon development. These observations suggest that some of the class IV homeodomain-Leucine zipper members act redundantly with other members of the family during various aspects of cell differentiation. DNA-binding sites were determined for two of the family members using polymerase chain reaction-assisted DNA selection from random oligonucleotides with their recombinant proteins. The binding sites were found to be similar to those previously identified for ATML1 and PDF2, which correspond to the pseudopalindromic sequence 5'-GCATTAAATGC-3' as the preferential binding site. 
