{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T12:48:14Z","timestamp":1792068494779,"version":"4.4.0"},"reference-count":51,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T00:00:00Z","timestamp":1792022400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T00:00:00Z","timestamp":1792022400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["JIPB"],"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Apple\n                    <jats:italic>Alternaria<\/jats:italic>\n                    blotch, caused by\n                    <jats:italic>Alternaria alternata<\/jats:italic>\n                    f. sp.\n                    <jats:italic>mali<\/jats:italic>\n                    (\n                    <jats:italic>A. alternata<\/jats:italic>\n                    ), is a devastating fungal disease threatening global apple production. However, the early signaling pathway underlying apple resistance to\n                    <jats:italic>A. alternata<\/jats:italic>\n                    remains unclear. In this study, through transcriptomic analysis, we identified that\n                    <jats:italic>MdWRKY9<\/jats:italic>\n                    is a key positive regulator of apple resistance to\n                    <jats:italic>A. alternata<\/jats:italic>\n                    , whose expression is induced in the resistant \u201cHanfu\u201d while being reduced in the susceptible \u201cGala\u201d following\n                    <jats:italic>A. alternata<\/jats:italic>\n                    infection. Stable overexpression of\n                    <jats:italic>MdWRKY9<\/jats:italic>\n                    enhanced apple resistance to\n                    <jats:italic>A. alternata<\/jats:italic>\n                    , whereas RNA interference (RNAi)\u2010mediated knockdown of\n                    <jats:italic>MdWRKY9<\/jats:italic>\n                    increased susceptibility. We further demonstrated that MdWRKY9 directly represses the expression of\n                    <jats:italic>MdJAZ8<\/jats:italic>\n                    , a key repressor of jasmonate (JA) signaling. Accordingly, stable overexpression of\n                    <jats:italic>MdJAZ8<\/jats:italic>\n                    in apple calli reduced resistance to\n                    <jats:italic>A. alternata<\/jats:italic>\n                    and suppressed JA signaling. Consistently, transient overexpression of\n                    <jats:italic>MdJAZ8<\/jats:italic>\n                    in apple leaves decreased resistance to\n                    <jats:italic>A. alternata<\/jats:italic>\n                    , whereas transient silencing of\n                    <jats:italic>MdJAZ8<\/jats:italic>\n                    via RNAi enhanced disease resistance. Furthermore, we identified that the receptor\u2010like cytoplasmic kinase MdBIK1 (\n                    <jats:italic>BOTRYTIS<\/jats:italic>\n                    \u2010INDUCED KINASE1, a core component of the PTI pathway) interacts with and phosphorylates MdWRKY9 at Thr\u20102. The phosphorylation of MdWRKY9 enhances resistance to\n                    <jats:italic>A. alternata<\/jats:italic>\n                    by further suppressing\n                    <jats:italic>MdJAZ8<\/jats:italic>\n                    and activating JA signaling. Together, our findings define a complete PTI\u2010associated signaling module in which the MdBIK1\u2013MdWRKY9 complex enhances JA\u2010mediated immunity against\n                    <jats:italic>A. alternata<\/jats:italic>\n                    by directly repressing\n                    <jats:italic>MdJAZ8<\/jats:italic>\n                    . This study refines the regulatory mechanism by which the BIK1 family kinases modulate JA signaling, providing new insights into the plant immune signaling network.\n                  <\/jats:p>","DOI":"10.1111\/jipb.70414","type":"journal-article","created":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T11:57:30Z","timestamp":1792065450000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["The BIK1\u2013WRKY9 module activates jasmonate signaling to confer apple resistance to\n                    <i>Alternaria<\/i>\n                    blotch"],"prefix":"10.1111","author":[{"ORCID":"https:\/\/orcid.org\/0009-0004-1153-761X","authenticated-orcid":false,"given":"Yixue","family":"Bai","sequence":"first","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-0815-5049","authenticated-orcid":false,"given":"Dongqian","family":"Shan","sequence":"additional","affiliation":[{"name":"College of Horticulture Northwest A &amp; F University Yangling 712100 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-4494-9945","authenticated-orcid":false,"given":"Chanyu","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-8982-1065","authenticated-orcid":false,"given":"Zehui","family":"Hu","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-0175-1925","authenticated-orcid":false,"given":"Tianci","family":"Yan","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3009-1770","authenticated-orcid":false,"given":"Tong","family":"Zhang","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-3377-0962","authenticated-orcid":false,"given":"Xin","family":"Liu","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-1849-4416","authenticated-orcid":false,"given":"Handong","family":"Song","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2178-9525","authenticated-orcid":false,"given":"Zhaoyang","family":"Zhou","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3255-269X","authenticated-orcid":false,"given":"Jun","family":"Liu","sequence":"additional","affiliation":[{"name":"College of Plant Protection China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6955-8008","authenticated-orcid":false,"given":"Yan","family":"Guo","sequence":"additional","affiliation":[{"name":"College of Biological Sciences China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0198-4852","authenticated-orcid":false,"given":"Wei","family":"Li","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-5786-5848","authenticated-orcid":false,"given":"Xiaodong","family":"Zheng","sequence":"additional","affiliation":[{"name":"College of Horticulture Qingdao Agricultural University Qingdao 266000 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0828-7208","authenticated-orcid":false,"given":"Jin","family":"Kong","sequence":"additional","affiliation":[{"name":"College of Horticulture China Agricultural University Beijing 100193 China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,10,15]]},"reference":[{"key":"e_1_2_9_2_1","first-page":"12747","article-title":"Appliance of fungicides on the Alternaria leaf spot disease of red delicious apple (Malus domestica Borkh.) caused by the pathogen Alternaria mali","volume":"5","author":"Aalum K.","year":"2016","journal-title":"Int. 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