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Here, we revealed that O-glycosylated and malonylated flavonols were specifically accumulated in younger leaves of Morus notabilis. A key acyltransferase, namely, MnMaT1, in the flavonol-associated module generated through weighted gene coexpression network analysis (WGCNA), was identified. Subsequent in vitro enzyme assays and in vivo function studies of MnMaT1 revealed its functional role as malonyl-CoA: flavonoid 3\/7-O-glucoside-6\u2033-O-malonyltransferase. A homeobox-leucine zipper protein ATHB51 (MnATHB51) in the same module was also identified and found to directly bind to the MnMaT1 promoter and possess transcriptional activity in mulberry leaves. The results indicated that MnATHB51 is a positive regulator governing the acylated flavonol glucosides biosynthesis in mulberry. Our findings elucidate a key transcriptional mechanism in specialized phenylpropanoid metabolism and provide a molecular foundation for the targeted enhancement of bioactive phytochemicals of mulberry.<\/jats:p>","DOI":"10.1021\/acs.jafc.6c00705","type":"journal-article","created":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T17:02:04Z","timestamp":1786640524000},"page":"26196-26208","source":"Crossref","is-referenced-by-count":0,"title":["A Flavonoid Malonyltransferase\nMnMaT1, Directly Regulated\nby\n                    <i>MnATHB51<\/i>\n                    , Leads to the Biosynthesis of Acylated\nFlavonol Glucosides in\n                    <i>Morus notabilis<\/i>"],"prefix":"10.1021","volume":"74","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2365-9487","authenticated-orcid":true,"given":"Dong","family":"Li","sequence":"first","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fengyao","family":"Chen","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pan","family":"Jiang","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zezhao","family":"Lin","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ting","family":"Lin","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianglian","family":"Yuan","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2750-253X","authenticated-orcid":true,"given":"Zhen","family":"Yang","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pei","family":"Yao","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Biyue","family":"Ding","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7322-7113","authenticated-orcid":true,"given":"Ningjia","family":"He","sequence":"additional","affiliation":[{"name":"Southwest University , , ,","place":["Chongqing, China, 400715"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,8,13]]},"reference":[{"key":"2026090104472683500_cit1","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms3445","article-title":"Draft genome sequence of the mulberry\ntree Morus notabilis","volume":"4","author":"He","year":"2013","journal-title":"Nat. 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Res."},{"issue":"10","key":"2026090104472683500_cit3","doi-asserted-by":"publisher","DOI":"10.1093\/hr\/uhaf171","article-title":"Functional characterization\nof key enzymes involved in the biosynthesis of distinctive flavonoids\nand stilbenoids in Morus notabilis","volume":"12","author":"Hu","year":"2025","journal-title":"Hortic.\nRes."},{"issue":"2","key":"2026090104472683500_cit4","doi-asserted-by":"publisher","first-page":"1307","DOI":"10.1093\/plphys\/kiad065","article-title":"Dehydrogenase\nMnGutB1\ncatalyzes 1-deoxynojirimycin biosynthesis in mulberry","volume":"192","author":"Yang","year":"2023","journal-title":"Plant Physiol."},{"issue":"6","key":"2026090104472683500_cit5","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1080\/09637480310001622288","article-title":"Mulberry\n(Morus alba) leaves as human food:\na new dimension of sericulture","volume":"54","author":"Srivastava","year":"2003","journal-title":"Int. J. Food\nSci. 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Produce Most Acylated Anthocyanins Present in Grape Skins","volume":"169","author":"Rinaldo","year":"2015","journal-title":"Plant Physiol."},{"issue":"3","key":"2026090104472683500_cit28","doi-asserted-by":"publisher","first-page":"683","DOI":"10.1111\/tpj.16166","article-title":"FaMYB123\ninteracts with FabHLH3 to regulate the late steps of anthocyanin and\nflavonol biosynthesis during ripening","volume":"114","author":"Marti\u0301nez-Rivas","year":"2023","journal-title":"Plant\nJ."},{"issue":"3","key":"2026090104472683500_cit29","doi-asserted-by":"publisher","DOI":"10.1093\/plphys\/kiaf567","article-title":"An HDG11-mediated\ntranscriptional network regulates 1-deoxynojirimycin biosynthesis\nin mulberry","volume":"199","author":"Yang","year":"2025","journal-title":"Plant Physiol."},{"issue":"5","key":"2026090104472683500_cit30","doi-asserted-by":"publisher","first-page":"1858","DOI":"10.1111\/nph.17453","article-title":"AaWRKY9 contributes\nto light- and jasmonate-mediated to regulate the biosynthesis of artemisinin\nin Artemisia annua","volume":"231","author":"Fu","year":"2021","journal-title":"New Phytol."},{"key":"2026090104472683500_cit31","doi-asserted-by":"publisher","DOI":"10.1093\/database\/baaa011","article-title":"MMHub,\na database for the mulberry metabolome","volume":"2020","author":"Li","year":"2020","journal-title":"Database"},{"issue":"D1","key":"2026090104472683500_cit32","doi-asserted-by":"publisher","first-page":"D1040","DOI":"10.1093\/nar\/gkw982","article-title":"PlantTFDB 4.0: toward\na central hub\nfor transcription factors and regulatory interactions in plants","volume":"45","author":"Jin","year":"2017","journal-title":"Nucleic Acids Res."},{"issue":"D1","key":"2026090104472683500_cit33","doi-asserted-by":"publisher","first-page":"D1104","DOI":"10.1093\/nar\/gkz1020","article-title":"PlantRegMap:\ncharting functional regulatory maps in plants","volume":"48","author":"Tian","year":"2020","journal-title":"Nucleic Acids Res."},{"issue":"18","key":"2026090104472683500_cit34","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21186541","article-title":"The Beneficial Effects\nof Morusin, an Isoprene Flavonoid Isolated from the Root Bark of Morus","volume":"21","author":"Choi","year":"2020","journal-title":"Int. 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Ethnopharmacol."},{"issue":"21","key":"2026090104472683500_cit37","doi-asserted-by":"publisher","DOI":"10.3390\/molecules27217580","article-title":"Synthesis,\nBiosynthesis, and Biological Activity of Diels-Alder Adducts from Morus Genus: An Update","volume":"27","author":"Tortora","year":"2022","journal-title":"Molecules"},{"issue":"7","key":"2026090104472683500_cit38","doi-asserted-by":"publisher","first-page":"1399","DOI":"10.1007\/s00210-023-02434-4","article-title":"Morus alba: a comprehensive phytochemical and pharmacological review","volume":"396","author":"Batiha","year":"2023","journal-title":"Naunyn-Schmiedeberg\u2019s Arch. 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Plant."},{"issue":"5","key":"2026090104472683500_cit48","doi-asserted-by":"publisher","first-page":"2303","DOI":"10.1111\/tpj.16918","article-title":"The HD-ZIP\nIV transcription factor GLABRA2 acts as an activator for proanthocyanidin\nbiosynthesis in Medicago seed coat","volume":"119","author":"Gu","year":"2024","journal-title":"Plant J."},{"issue":"5","key":"2026090104472683500_cit49","doi-asserted-by":"publisher","first-page":"1668","DOI":"10.1093\/jxb\/erab511","article-title":"An EjbHLH14-EjHB1-EjPRX12\nmodule is involved in methyl jasmonate alleviation of chilling-induced\nlignin deposition in loquat fruit","volume":"73","author":"Zhang","year":"2022","journal-title":"J. Exp. Bot."},{"key":"2026090104472683500_cit50","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1016\/j.plaphy.2019.11.033","article-title":"Functional characterization\nof a HD-ZIP IV transcription factor NtHDG2 in regulating flavonols\nbiosynthesis in Nicotiana tabacum","volume":"146","author":"Wang","year":"2020","journal-title":"Plant Physiol. 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