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To test whether theanine biosynthesis can be reconstructed in a nontea crop and whether this metabolic network contributes to stress tolerance, we heterologously expressed the tea alanine decarboxylase gene CsAlaDC in tomato (Solanum lycopersicum cv. Micro-Tom). The resulting OE-CsAlaDC lines accumulated ethylamine and synthesized theanine without introducing a tea theanine synthase gene, demonstrating that endogenous tomato glutamine synthetase supports theanine formation. Transgenic plants exhibited distinct morphological changes, including dwarfism and dark-green leaves, while their fruits showed accelerated development, elevated levels of theanine and GABA, and improved quality-related traits such as enhanced lycopene accumulation. Under heat stress, OE-CsAlaDC plants maintained higher PSII efficiency, reduced membrane damage and reactive oxygen species accumulation, and stronger antioxidant enzyme activities than wild-type plants. Exogenous theanine further enhanced thermotolerance, promoted SlGAD1\/2 expression, and increased GABA accumulation, whereas silencing SlGAD1 and SlGAD2 markedly diminished the protective effect of theanine. Exogenous ethylamine also conferred partial heat protection, but theanine showed a stronger association with GAD-dependent GABA biosynthesis. Collectively, these findings demonstrate that heterologous expression of CsAlaDC establishes a functional ethylamine\u2013theanine metabolic branch in tomato and enhances thermotolerance through coordination with the GABA metabolic network, offering a promising strategy to improve both stress resilience and nutritional quality in crops.<\/jats:p>","DOI":"10.1093\/plphys\/kiag543","type":"journal-article","created":{"date-parts":[[2026,7,24]],"date-time":"2026-07-24T11:40:51Z","timestamp":1784893251000},"source":"Crossref","is-referenced-by-count":0,"title":["Heterologous expression of\n                    <i>CsAlaDC<\/i>\n                    enhances thermotolerance through a functional ethylamine\u2013theanine\u2013GABA metabolic network in tomato"],"prefix":"10.1093","volume":"201","author":[{"given":"Qianying","family":"Wang","sequence":"first","affiliation":[{"name":"Key Laboratory of Tea Quality and Safety Control, Ministry of Agriculture and Rural Affairs, Tea Research Institute , Chinese Academy of Agricultural Sciences, Hangzhou 310008 ,","place":["China"]},{"name":"Department of Horticulture, Zijingang Campus, Zhejiang University , Hangzhou 310058 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-8520-1504","authenticated-orcid":false,"given":"Jingbo","family":"Yu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tea Quality and Safety Control, Ministry of Agriculture and Rural Affairs, Tea Research Institute , Chinese Academy of Agricultural Sciences, Hangzhou 310008 ,","place":["China"]},{"name":"Tea Science Research Institute, Natural Small-Molecule Pesticides Laboratory, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-6650-5094","authenticated-orcid":false,"given":"Peng","family":"Mao","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tea 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310008 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-8100-4665","authenticated-orcid":false,"given":"Fengshui","family":"Yang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tea Quality and Safety Control, Ministry of Agriculture and Rural Affairs, Tea Research Institute , Chinese Academy of Agricultural Sciences, Hangzhou 310008 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-8216-5688","authenticated-orcid":false,"given":"Dongmiao","family":"Zhai","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tea Quality and Safety Control, Ministry of Agriculture and Rural Affairs, Tea Research Institute , Chinese Academy of Agricultural Sciences, Hangzhou 310008 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