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Cytochrome P450 hydroxylases require electron-transfer partners, but their coordination during ripening remains unclear. Integrated transcriptomic and targeted metabolomic analyses identified the endoplasmic reticulum-localized cytochrome b5 VvCB5D as a VvMYBA1 co-expressed gene in grape (Vitis vinifera). VvMYBA1 directly activated VvCB5D transcription by binding its promoter, and VvCB5D interacted with VvF3\u2032H and VvF3\u20325\u2032H, preferentially with VvF3\u20325\u2032H. Genetic manipulation in grape berries, tobacco leaves, and transgenic tomatoes showed that VvCB5D directed flavonoid flux toward the 3\u2032,4\u2032,5\u2032-trihydroxylated branch, increasing delphinidin, petunidin, and malvidin derivatives. These results support a model in which VvMYBA1 coordinates flavonoid-pathway gene expression with a cytochrome b5 partner and identify VvCB5D as a candidate for modifying grape flavonoid composition.<\/jats:p>","DOI":"10.1021\/acs.jafc.6c08391","type":"journal-article","created":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T04:01:59Z","timestamp":1791518519000},"page":"32599-32614","source":"Crossref","is-referenced-by-count":0,"title":["The VvMYBA1-Activated\nElectron Carrier VvCB5D Directs\nFlavonoid Metabolic Flux toward the Trihydroxylated Branch in Grape"],"prefix":"10.1021","volume":"74","author":[{"given":"Ziyan","family":"Guo","sequence":"first","affiliation":[{"name":"Nanjing Agricultural University , , ,","place":["Nanjing, China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mengyu","family":"Xu","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , ,","place":["Nanjing, China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yang","family":"Xiao","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , ,","place":["Nanjing, China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinggui","family":"Fang","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , ,","place":["Nanjing, China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5876-8819","authenticated-orcid":true,"given":"Suwen","family":"Lu","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , ,","place":["Nanjing, China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,10,8]]},"reference":[{"key":"2026102109080487600_cit1","doi-asserted-by":"publisher","first-page":"132531","DOI":"10.1016\/j.foodchem.2022.132531","article-title":"Plant flavonoids: Classification, distribution, biosynthesis, and\nantioxidant activity","volume":"383","author":"Shen","year":"2022","journal-title":"Food Chem."},{"issue":"1","key":"2026102109080487600_cit2","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1104\/pp.105.073262","article-title":"Identification of\nthe Flavonoid Hydroxylases from Grapevine and Their Regulation during\nFruit Development","volume":"140","author":"Bogs","year":"2006","journal-title":"Plant Physiol."},{"issue":"5","key":"2026102109080487600_cit3","doi-asserted-by":"publisher","first-page":"1003","DOI":"10.1007\/s00425-014-2166-3","article-title":"The B-ring hydroxylation pattern\nof anthocyanins can be determined through activity of the flavonoid\n3\u2032-hydroxylase on leucoanthocyanidins","volume":"240","author":"Schwinn","year":"2014","journal-title":"Planta"},{"issue":"9","key":"2026102109080487600_cit4","doi-asserted-by":"publisher","first-page":"1001","DOI":"10.3390\/horticulturae9091001","article-title":"Anthocyanin\nAccumulation and Its Corresponding Gene Expression, Total Phenol,\nAntioxidant Capacity, and Fruit Quality of \u2018Crimson Seedless\u2019\nGrapevine (Vitis vinifera L.) in Response to Grafting and Pre-Harvest\nApplications","volume":"9","author":"Salama","year":"2023","journal-title":"Horticulturae"},{"issue":"10","key":"2026102109080487600_cit5","doi-asserted-by":"publisher","first-page":"909","DOI":"10.1021\/acs.biochem.2c00002","article-title":"Biochemical Investigation\nof Membrane-Bound Cytochrome b5 and the\nCatalytic Domain of Cytochrome b5 Reductase from Arabidopsis thaliana","volume":"61","author":"Iqbal","year":"2022","journal-title":"Biochemistry"},{"issue":"2","key":"2026102109080487600_cit6","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.ade4389","article-title":"Tissue-preferential\nrecruitment of electron transfer chains for cytochrome P450-catalyzed\nphenolic biosynthesis","volume":"9","author":"Zhao","year":"2023","journal-title":"Sci. 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