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PAs have been associated with human health benefits and also play important roles in pest and disease defense throughout the plant.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      To dissect the genetic basis of PA biosynthetic pathway in cacao (\n                      <jats:italic>Theobroma cacao<\/jats:italic>\n                      ), we have isolated three genes encoding key PA synthesis enzymes, anthocyanidin synthase (ANS), anthocyanidin reductase (ANR) and leucoanthocyanidin reductase (LAR). We measured the expression levels of\n                      <jats:italic>TcANR<\/jats:italic>\n                      ,\n                      <jats:italic>TcANS<\/jats:italic>\n                      and\n                      <jats:italic>TcLAR<\/jats:italic>\n                      and PA content in cacao leaves, flowers, pod exocarp and seeds. In all tissues examined, all three genes were abundantly expressed and well correlated with PA accumulation levels, suggesting their active roles in PA synthesis. Overexpression of\n                      <jats:italic>TcANR<\/jats:italic>\n                      in an Arabidopsis\n                      <jats:italic>ban<\/jats:italic>\n                      mutant complemented the PA deficient phenotype in seeds and resulted in reduced anthocyanidin levels in hypocotyls. Overexpression of\n                      <jats:italic>TcANS<\/jats:italic>\n                      in tobacco resulted in increased content of both anthocyanidins and PAs in flower petals. Overexpression of\n                      <jats:italic>TcANS<\/jats:italic>\n                      in an Arabidopsis\n                      <jats:italic>ldox<\/jats:italic>\n                      mutant complemented its PA deficient phenotype in seeds. Recombinant TcLAR protein converted leucoanthocyanidin to catechin\n                      <jats:italic>in vitro<\/jats:italic>\n                      . Transgenic tobacco overexpressing\n                      <jats:italic>TcLAR<\/jats:italic>\n                      had decreased amounts of anthocyanidins and increased PAs. Overexpressing\n                      <jats:italic>TcLAR<\/jats:italic>\n                      in Arabidopsis\n                      <jats:italic>ldox<\/jats:italic>\n                      mutant also resulted in elevated synthesis of not only catechin but also epicatechin.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusion<\/jats:title>\n                    <jats:p>\n                      Our results confirm the\n                      <jats:italic>in vivo<\/jats:italic>\n                      function of cacao\n                      <jats:italic>ANS<\/jats:italic>\n                      and\n                      <jats:italic>ANR<\/jats:italic>\n                      predicted based on sequence homology to previously characterized enzymes from other species. In addition, our results provide a clear functional analysis of a\n                      <jats:italic>LAR<\/jats:italic>\n                      gene\n                      <jats:italic>in vivo<\/jats:italic>\n                      .\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/1471-2229-13-202","type":"journal-article","created":{"date-parts":[[2013,12,5]],"date-time":"2013-12-05T08:12:53Z","timestamp":1386231173000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":107,"title":["Proanthocyanidin synthesis in Theobroma cacao: genes encoding anthocyanidin synthase, anthocyanidin reductase, and leucoanthocyanidin reductase"],"prefix":"10.1186","volume":"13","author":[{"given":"Yi","family":"Liu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zi","family":"Shi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Siela","family":"Maximova","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mark J","family":"Payne","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mark J","family":"Guiltinan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2013,12,5]]},"reference":[{"key":"1404_CR1","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1111\/j.1469-8137.2004.01217.x","volume":"165","author":"RA Dixon","year":"2005","unstructured":"Dixon RA, Xie DY, Sharma SB: Proanthocyanidins - a final frontier in flavonoid research?. 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