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Here, we show that acute PEG\u2010induced osmotic stress induces rapid caffeine accumulation, accompanied by transcriptional activation of the key biosynthetic gene\n                    <jats:italic>CsTCS1<\/jats:italic>\n                    . Through integrative analyses, including GWAS and co\u2010expression network modeling, we identified a\n                    <jats:italic>CsBPC6<\/jats:italic>\n                    \u2013\n                    <jats:italic>CsNACX2<\/jats:italic>\n                    \u2013\n                    <jats:italic>CsTCS1<\/jats:italic>\n                    module as a central regulatory hub. Mechanistically,\n                    <jats:italic>CsNACX2<\/jats:italic>\n                    directly binds the promoter of the caffeine synthase gene\n                    <jats:italic>CsTCS1<\/jats:italic>\n                    , activating its transcription and promoting caffeine accumulation. Moreover, the drought\u2010responsive transcription factor\n                    <jats:italic>CsBPC6<\/jats:italic>\n                    acts upstream to transactivate\n                    <jats:italic>CsNACX2<\/jats:italic>\n                    . Functional validation using gene silencing and transient overexpression assays confirmed that the\n                    <jats:italic>CsBPC6<\/jats:italic>\n                    \u2013\n                    <jats:italic>CsNACX2<\/jats:italic>\n                    \u2013\n                    <jats:italic>CsTCS1<\/jats:italic>\n                    module positively regulates caffeine biosynthesis under PEG\u2010induced drought conditions. Collectively, our findings uncover a previously uncharacterized drought\u2010responsive transcriptional module governing caffeine biosynthesis, providing new mechanistic insights into how environmental cues regulate secondary metabolism in tea plants.\n                  <\/jats:p>","DOI":"10.1111\/pce.70805","type":"journal-article","created":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T05:47:02Z","timestamp":1786081622000},"page":"9054-9069","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["<i>CsBPC6<\/i>\n                    \u2013\n                    <i>CsNACX2<\/i>\n                    \u2013\n                    <i>CsTCS1<\/i>\n                    Module Positively Regulates Caffeine Biosynthesis Under PEG\u2010Induced Drought Stress in Tea Plants\n                    <i>(Camellia sinensis)<\/i>"],"prefix":"10.1111","volume":"49","author":[{"given":"Shiyu","family":"Zhang","sequence":"first","affiliation":[{"name":"College of Life Sciences Guizhou University Guiyang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhouzhuoer","family":"Chen","sequence":"additional","affiliation":[{"name":"College of Life Sciences Guizhou University Guiyang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinzhuan","family":"Yao","sequence":"additional","affiliation":[{"name":"College of Tea Sciences\/The Key Laboratory of Plant Resources Conservation and Germplasm Innovation in Mountainous Region (Ministry of Education) Guizhou University Guiyang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shilai","family":"Tian","sequence":"additional","affiliation":[{"name":"College of Life Sciences Guizhou University Guiyang 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