title: The Responses of Arabidopsis Early Light-Induced Protein2 to Ultraviolet B, High Light, and Cold Stress Are Regulated by a Transcriptional Regulatory Unit Composed of Two Elements.
abstract: The Arabidopsis (Arabidopsis thaliana) Early Light-Induced Protein (ELIP) is thought to act as a photoprotectant, reducing the damaging effects of high light (HL). Expression of ELIP2 is activated by multiple environmental stresses related to photoinhibition. We have identified putative regulatory elements in an ELIP2 promoter using an octamer-based frequency comparison method, analyzed the role of these elements using synthetic promoters, and revealed a key transcriptional regulatory unit for ultraviolet B (UV-B) radiation, HL, and cold stress responses. The unit is composed of two elements, designated as Elements A (TACACACC) and B (GGCCACGCCA), and shows functionality only when paired. Our genome-wide correlation analysis between possession of these elements in the promoter region and expression profiles in response to UV-B, HL, and cold suggests that Element B receives and integrates these multiple stress signals. In vitro protein-DNA binding assays revealed that LONG HYPOCOTYL5 (HY5), a basic domain-Leucine zipper transcription factor, directly binds to Element B. In addition, mutant analysis of HY5 showed partial involvement in the UV-B and HL responses but not in the cold stress response. These results suggest that signals for UV-B, HL, and cold stress join at Element B, which recognizes the signals of multiple transcription factors, including HY5.
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Natsuki
authors.initials: N
authors.lastname: Hayami
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Yusaku
authors.initials: Y
authors.lastname: Sakai
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Mitsuhiro
authors.initials: M
authors.lastname: Kimura
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Tatsunori
authors.initials: T
authors.lastname: Saito
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Mutsutomo
authors.initials: M
authors.lastname: Tokizawa
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Satoshi
authors.initials: S
authors.lastname: Iuchi
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Yukio
authors.initials: Y
authors.lastname: Kurihara
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Minami
authors.initials: M
authors.lastname: Matsui
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Mika
authors.initials: M
authors.lastname: Nomoto
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.).
authors.forename: Yasuomi
authors.initials: Y
authors.lastname: Tada
authors.affiliation: Faculty of Applied Biological Sciences (N.H., Y.S., T.S., Y.Y.Y.) and United Graduate School of Agricultural Science (M.T., Y.Y.Y.), Gifu University, Gifu 501-1103, Japan; Department of Frontier Research, Kazusa DNA Research Institute, Kazusa-kamatari 2-6-7, Kisarazu, Chiba 292-0818, Japan (M.K.);RIKEN Bioresource Center, Koyadai 3-1-1, Tsukuba, Ibaraki 305-0074, Japan (S.I.);RIKEN Center for Sustainable Resource Science, Suehiro-cho 1-7-22, Tsurumi-ku, Yokohama City, Kanagawa 230-0045, Japan (Y.K., M.M., Y.Y.Y.); andCenter for Gene Research, Nagoya University, Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan (M.N., Y.T.) yyy@gifu-u.ac.jp.
authors.forename: Yoshiharu Y
authors.identifier: http://orcid.org/0000-0002-9667-0572
authors.initials: YY
authors.lastname: Yamamoto
mesh_terms: D017360:Arabidopsis / Q000235:genetics / Q000528:radiation effects; D029681:Arabidopsis Proteins / Q000235:genetics* / Q000378:metabolism; D050976:Basic-Leucine Zipper Transcription Factors / Q000378:metabolism; D002940:Circadian Rhythm / Q000528:radiation effects; D003080:Cold Temperature; D020869:Gene Expression Profiling; D018506:Gene Expression Regulation, Plant / Q000528:radiation effects*; D008156:Luciferases / Q000378:metabolism; D008957:Models, Genetic; D009687:Nuclear Proteins / Q000378:metabolism; D020411:Oligonucleotide Array Sequence Analysis; D009928:Organ Specificity / Q000235:genetics; D011401:Promoter Regions, Genetic; D011485:Protein Binding; D013312:Stress, Physiological / Q000528:radiation effects*; D014158:Transcription, Genetic / Q000528:radiation effects; D014466:Ultraviolet Rays
chemical_list: D029681:Arabidopsis Proteins; D050976:Basic-Leucine Zipper Transcription Factors; C481743:ELIP2 protein, Arabidopsis; C109254:HY5 protein, Arabidopsis; D009687:Nuclear Proteins; D008156:Luciferases