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A revised model for flavonoid biosynthesis in\n                      <jats:italic>Arabidopsis thaliana<\/jats:italic>\n                      is proposed that integrates the regulatory roles of these R2R3\u2013MYBs across early and late biosynthetic steps.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      Flavonoids are a large group of specialized plant metabolites. Their biosynthesis is mainly transcriptionally regulated by a sophisticated network of different transcription factors from various families, with R2R3\u2013MYB factors being the main determinant of specific flavonoid class formation. The early biosynthetic steps, leading to the formation of non-visible flavonoids, have been proposed to be regulated by three R2R3\u2013MYBs, PRODUCTION OF FLAVONOL GYLCOSIDE1-3 (PFG1-3), while the later biosynthetic steps leading to the formation of visible anthocyanin and proanthocyanidin pigments are reported to be regulated by four R2R3\u2013MYBs, PRODUCTION OF ANTHOCYANIN PIGMENT1-4 (PAP1-4) and TRANSPARENT TESTA2 (TT2), respectively. Several studies have indicated that this model for the transcriptional regulation of flavonoid biosynthesis may be incomplete. To address this issue, especially regarding the regulation of the early biosynthesis genes by PAP1-4 and TT2, we generated several multiple\n                      <jats:italic>r2r3-myb<\/jats:italic>\n                      mutant lines. We characterized the\n                      <jats:italic>pfg1-3<\/jats:italic>\n                      ,\n                      <jats:italic>pfg1-3 tt2<\/jats:italic>\n                      and\n                      <jats:italic>pfg1-3 pap1-4<\/jats:italic>\n                      mutants and did comparative metabolite profiling. This revealed that only the\n                      <jats:italic>pfg1-3 tt2<\/jats:italic>\n                      mutant was deficient in proanthocyanidins and only the\n                      <jats:italic>pfg1-3 pap1-4<\/jats:italic>\n                      mutant was deficient in anthocyanins. We demonstrate that PAP and TT2 R2R3\u2013MYBs are also capable of activating the early biosynthesis genes required for dihydroflavonol formation. Our results provide evidence that the traditional view of distinct branch-specific R2R3\u2013MYB regulators in flavonoid biosynthesis is overly simplistic. We, therefore, propose a revised model for the transcriptional regulation of flavonoid biosynthesis.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1007\/s00425-026-04938-8","type":"journal-article","created":{"date-parts":[[2026,2,6]],"date-time":"2026-02-06T13:49:03Z","timestamp":1770385743000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Combined R2R3\u2013MYB transcription factor mutants reveal the regulatory structure of the Arabidopsis thaliana flavonoid biosynthesis pathway"],"prefix":"10.1007","volume":"263","author":[{"ORCID":"https:\/\/orcid.org\/0009-0002-5987-6579","authenticated-orcid":false,"given":"Lennart Malte","family":"Sielmann","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-1787-7490","authenticated-orcid":false,"given":"Timo","family":"Denecke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7635-3473","authenticated-orcid":false,"given":"Bernd","family":"Weisshaar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9261-2279","authenticated-orcid":false,"given":"Ralf","family":"Stracke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,2,6]]},"reference":[{"issue":"5","key":"4938_CR1","doi-asserted-by":"publisher","first-page":"955","DOI":"10.1007\/s00425-014-2088-0","volume":"240","author":"I Appelhagen","year":"2014","unstructured":"Appelhagen I, Thiedig K, Nordholt N, Schmidt N, Huep G, Sagasser M, Weisshaar B (2014) Update on transparent testa mutants from Arabidopsis thaliana: characterisation of new alleles from an isogenic collection. 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