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In apple (<jats:italic>Malus\u00a0\u00d7\u00a0domestica<\/jats:italic>), inbreeding has resulted in severe decreases in the genetic diversity and flavonoid content. Over the last decade, we have focused on the genetic improvement of apple using wild red\u2010fleshed apple resources (<jats:italic>Malus sieversii<\/jats:italic> f<jats:italic>. niedzwetzkyana<\/jats:italic>). Here, we found that the <jats:styled-content style=\"fixed-case\">MYB<\/jats:styled-content> transcription factors (<jats:styled-content style=\"fixed-case\">TF<\/jats:styled-content>s) involved in the synthesis of proanthocyanidins can be classified into <jats:styled-content style=\"fixed-case\">TT<\/jats:styled-content>2 and <jats:styled-content style=\"fixed-case\">PA<\/jats:styled-content>1 types. We characterized a <jats:styled-content style=\"fixed-case\">PA<\/jats:styled-content>1\u2010type <jats:styled-content style=\"fixed-case\">MYB<\/jats:styled-content> transcription factor, <jats:italic>Md<jats:styled-content style=\"fixed-case\">MYBPA<\/jats:styled-content>1<\/jats:italic>, from red\u2010fleshed apple and identified its role in flavonoid biosynthesis using overexpression and knockdown\u2010expression transgenes in apple calli. We explored the relationship between <jats:styled-content style=\"fixed-case\">TT<\/jats:styled-content>2\u2010 and <jats:styled-content style=\"fixed-case\">PA<\/jats:styled-content>1\u2010type <jats:styled-content style=\"fixed-case\">MYB TF<\/jats:styled-content>s, and found that <jats:italic>Md<jats:styled-content style=\"fixed-case\">MYB<\/jats:styled-content>9\/11\/12<\/jats:italic> bind the <jats:italic>Md<jats:styled-content style=\"fixed-case\">MYBPA<\/jats:styled-content>1<\/jats:italic> promoter. In addition, <jats:italic>Md<jats:styled-content style=\"fixed-case\">MYBPA<\/jats:styled-content>1<\/jats:italic> responded to low temperature by redirecting the flavonoid biosynthetic pathway from proanthocyanidin to anthocyanin production. In binding analyses, <jats:italic>Mdb<jats:styled-content style=\"fixed-case\">HLH<\/jats:styled-content>33<\/jats:italic> directly bound to the low\u2010temperature\u2010responsive (<jats:styled-content style=\"fixed-case\">LTR<\/jats:styled-content>) <jats:italic>cis<\/jats:italic>\u2010element of the <jats:italic>Md<jats:styled-content style=\"fixed-case\">MYBPA<\/jats:styled-content>1<\/jats:italic> promoter and promotes its activity. In addition, the calli expressing both <jats:italic>Md<jats:styled-content style=\"fixed-case\">MYBPA<\/jats:styled-content>1<\/jats:italic> and <jats:italic>Mdb<jats:styled-content style=\"fixed-case\">HLH<\/jats:styled-content>33<\/jats:italic>, which together form a complex, produced more anthocyanin under low temperature. Our findings shed light on the essential roles of <jats:styled-content style=\"fixed-case\">PA<\/jats:styled-content>1\u2010type <jats:styled-content style=\"fixed-case\">TF<\/jats:styled-content>s in the metabolic network of proanthocyanidin and anthocyanin synthesis in plants. Studies on red\u2010fleshed wild apple are beneficial for breeding new apple varieties with high flavonoid contents.<\/jats:p>","DOI":"10.1111\/tpj.14013","type":"journal-article","created":{"date-parts":[[2018,7,6]],"date-time":"2018-07-06T08:25:40Z","timestamp":1530865540000},"page":"39-55","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":211,"title":["The proanthocyanidin\u2010specific transcription factor Md<scp>MYBPA<\/scp>1 initiates anthocyanin synthesis under low\u2010temperature conditions in red\u2010fleshed apples"],"prefix":"10.1111","volume":"96","author":[{"given":"Nan","family":"Wang","sequence":"first","affiliation":[{"name":"State Key Laboratory of Crop Biology Shandong Agricultural University No. 61 Daizong Road 271018 Tai'an Shandong China"},{"name":"College of Horticulture Sciences Shandong Agricultural University No. 61 Daizong Road 271018 Tai'an Shandong 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West Road 100094 Beijing China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haifeng","family":"Xu","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Biology Shandong Agricultural University No. 61 Daizong Road 271018 Tai'an Shandong China"},{"name":"College of Horticulture Sciences Shandong Agricultural University No. 61 Daizong Road 271018 Tai'an Shandong China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongcheng","family":"Fang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Biology Shandong Agricultural University No. 61 Daizong Road 271018 Tai'an Shandong China"},{"name":"College of Horticulture Sciences Shandong Agricultural University No. 61 Daizong Road 271018 Tai'an Shandong China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mengyu","family":"Su","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Biology Shandong Agricultural 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