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An important step in SA biosynthesis in Arabidopsis is the conversion of chorismate to isochorismate through the action of isochorismate synthase, encoded by the\n                      <jats:italic>ICS1<\/jats:italic>\n                      gene. Also\n                      <jats:italic>AVR<\/jats:italic>\n                      <jats:sub>\n                        <jats:italic>PPHB<\/jats:italic>\n                      <\/jats:sub>\n                      <jats:italic>SUSCEPTIBLE 3<\/jats:italic>\n                      (\n                      <jats:italic>PBS3<\/jats:italic>\n                      ) plays an important role in SA metabolism, as\n                      <jats:italic>pbs3<\/jats:italic>\n                      mutants accumulate drastically reduced levels of SA-glucoside, a putative storage form of SA. Bioinformatics analysis previously performed by us identified WRKY28 and WRKY46 as possible regulators of\n                      <jats:italic>ICS1<\/jats:italic>\n                      and\n                      <jats:italic>PBS3<\/jats:italic>\n                      .\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Expression studies with\n                      <jats:italic>ICS1 promoter::\u03b2-glucuronidase<\/jats:italic>\n                      (\n                      <jats:italic>GUS<\/jats:italic>\n                      ) genes in\n                      <jats:italic>Arabidopsis thaliana<\/jats:italic>\n                      protoplasts cotransfected with\n                      <jats:italic>35S::WRKY28<\/jats:italic>\n                      showed that over expression of WRKY28 resulted in a strong increase in GUS expression. Moreover, qRT-PCR analyses indicated that the endogenous\n                      <jats:italic>ICS1<\/jats:italic>\n                      and\n                      <jats:italic>PBS3<\/jats:italic>\n                      genes were highly expressed in protoplasts overexpressing WRKY28 or WRKY46, respectively. Electrophoretic mobility shift assays indentified potential WRKY28 binding sites in the\n                      <jats:italic>ICS1<\/jats:italic>\n                      promoter, positioned -445 and -460 base pairs upstream of the transcription start site. Mutation of these sites in protoplast transactivation assays showed that these binding sites are functionally important for activation of the\n                      <jats:italic>ICS1<\/jats:italic>\n                      promoter. Chromatin immunoprecipitation assays with haemagglutinin-epitope-tagged WRKY28 showed that the region of the\n                      <jats:italic>ICS1<\/jats:italic>\n                      promoter containing the binding sites at -445 and -460 was highly enriched in the immunoprecipitated DNA.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>\n                      The results obtained here confirm results from our multiple microarray co-expression analyses indicating that WRKY28 and WRKY46 are transcriptional activators of\n                      <jats:italic>ICS1<\/jats:italic>\n                      and\n                      <jats:italic>PBS3<\/jats:italic>\n                      , respectively, and support this\n                      <jats:italic>in silico<\/jats:italic>\n                      screening as a powerful tool for identifying new components of stress signaling pathways.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/1471-2229-11-89","type":"journal-article","created":{"date-parts":[[2011,5,19]],"date-time":"2011-05-19T15:06:44Z","timestamp":1305817604000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":208,"title":["WRKY Transcription Factors Involved in Activation of SA Biosynthesis Genes"],"prefix":"10.1186","volume":"11","author":[{"given":"Marcel C","family":"van Verk","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John F","family":"Bol","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huub JM","family":"Linthorst","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2011,5,19]]},"reference":[{"key":"875_CR1","doi-asserted-by":"publisher","first-page":"1004","DOI":"10.1126\/science.250.4983.1004","volume":"250","author":"JP M\u00e9traux","year":"1990","unstructured":"M\u00e9traux JP, Signer H, Ryals J, Ward E, Wyss-Benz M, Gaudin J, Raschdorf K, Schmid E, Blum W, Inverardi B: Increase in Salicylic Acid at the Onset of Systemic Acquired Resistance in Cucumber. 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