title: SENSITIVE TO PROTON RHIZOTOXICITY1, CALMODULIN BINDING TRANSCRIPTION ACTIVATOR2, and other transcription factors are involved in ALUMINUM-ACTIVATED MALATE TRANSPORTER1 expression.
abstract: In Arabidopsis (Arabidopsis thaliana) the root apex is protected from aluminum (Al) rhizotoxicity by excretion of malate, an Al chelator, by ALUMINUM-ACTIVATED MALATE TRANSPORTER1 (AtALMT1). AtALMT1 expression is fundamentally regulated by the SENSITIVE TO PROTON RHIZOTOXICITY1 (STOP1) zinc finger protein, but other transcription factors have roles that enable Al-inducible expression with a broad dynamic range. In this study, we characterized multiple cis-elements in the AtALMT1 promoter that interact with transcription factors. In planta complementation assays of AtALMT1 driven by 5' truncated promoters of different lengths showed that the promoter region between -540 and 0 (the first ATG) restored the Al-sensitive phenotype of atalm1 and thus contains cis-elements essential for AtALMT1 expression for Al tolerance. Computation of overrepresented octamers showed that eight regions in this promoter region contained potential cis-elements involved in Al induction and STOP1 regulation. Mutation in a position around -297 from the first ATG completely inactivated AtALMT1 expression and Al response. In vitro binding assays showed that this region contained the STOP1 binding site, which accounted for the recognition by four zinc finger domains of the protein. Other positions were characterized as cis-elements that regulated expression by repressors and activators and a transcription factor that determines root tip expression of AtALMT1. From the consensus of known cis-elements, we identified CALMODULIN-BINDING TRANSCRIPTION ACTIVATOR2 to be an activator of AtALMT1 expression. Al-inducible expression of AtALMT1 changed transcription starting sites, which increased the abundance of transcripts with a shortened 5' untranslated region. The present analyses identified multiple mechanisms that regulate AtALMT1 expression.
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan.
authors: Mutsutomo
authors: M
authors: Tokizawa
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan.
authors: Yuriko
authors: Y
authors: Kobayashi
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan.
authors: Tatsunori
authors: T
authors: Saito
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan.
authors: Masatomo
authors: M
authors: Kobayashi
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan.
authors: Satoshi
authors: S
authors: Iuchi
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan.
authors: Mika
authors: M
authors: Nomoto
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan.
authors: Yasuomi
authors: Y
authors: Tada
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan koyama@gifu-u.ac.jp yyy@gifu-u.ac.jp.
authors: Yoshiharu Y
authors: YY
authors: Yamamoto
authors: Applied Biological Sciences, Gifu University, Gifu 501-1193, Japan (M.T., Y.K., T.S., Y.Y.Y., H.K.);RIKEN BioResource Center, Ibaraki 305-0074, Japan (M.K., S.I.); andDivision of Biological Science, Graduate School of Science (M.N.), and The Center for Gene Research, Division of Biological Science (Y.T.), Nagoya University, Aichi 464-8602, Japan koyama@gifu-u.ac.jp yyy@gifu-u.ac.jp.
authors: Hiroyuki
authors: H
authors: Koyama
mesh_terms: D000222:Adaptation, Physiological / Q000187:drug effects; D000535:Aluminum / Q000633:toxicity; D017360:Arabidopsis / Q000187:drug effects / Q000235:genetics* / Q000378:metabolism; D029681:Arabidopsis Proteins / Q000737:chemistry / Q000235:genetics / Q000378:metabolism*; D001483:Base Sequence; D020869:Gene Expression Profiling; D018506:Gene Expression Regulation, Plant / Q000187:drug effects; D005816:Genetic Complementation Test; D005966:Glucuronidase / Q000378:metabolism; D008954:Models, Biological; D008969:Molecular Sequence Data; D009154:Mutation / Q000235:genetics; D027361:Organic Anion Transporters / Q000235:genetics / Q000378:metabolism*; D030821:Plants, Genetically Modified; D011401:Promoter Regions, Genetic; D011485:Protein Binding / Q000187:drug effects; D017434:Protein Structure, Tertiary; D012333:RNA, Messenger / Q000235:genetics / Q000378:metabolism; D013194:Staining and Labeling; D014157:Transcription Factors / Q000737:chemistry / Q000235:genetics / Q000378:metabolism*; D024363:Transcription Initiation Site; D014158:Transcription, Genetic / Q000187:drug effects; D016335:Zinc Fingers
chemical_list: C512361:ALMT1 protein, Arabidopsis; D029681:Arabidopsis Proteins; D027361:Organic Anion Transporters; D012333:RNA, Messenger; C525886:STOP1 protein, Arabidopsis; D014157:Transcription Factors; D000535:Aluminum; D005966:Glucuronidase
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