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Background  The proanthocyanidins (PAs), a subgroup of flavonoids, accumulate to levels of approximately 10% total dry weight of cacao seeds. PAs have been associated with human health benefits and also play important roles in pest and disease defense throughout the plant.    Results  To dissect the genetic basis of PA biosynthetic pathway in cacao ( Theobroma cacao ), 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  TcANR ,  TcANS  and  TcLAR  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  TcANR  in an Arabidopsis  ban  mutant complemented the PA deficient phenotype in seeds and resulted in reduced anthocyanidin levels in hypocotyls. Overexpression of  TcANS  in tobacco resulted in increased content of both anthocyanidins and PAs in flower petals. Overexpression of  TcANS  in an Arabidopsis  ldox  mutant complemented its PA deficient phenotype in seeds. Recombinant TcLAR protein converted leucoanthocyanidin to catechin  in vitro . Transgenic tobacco overexpressing  TcLAR  had decreased amounts of anthocyanidins and increased PAs. Overexpressing  TcLAR  in Arabidopsis  ldox  mutant also resulted in elevated synthesis of not only catechin but also epicatechin.    Conclusion  Our results confirm the  in vivo  function of cacao  ANS  and  ANR  predicted based on sequence homology to previously characterized enzymes from other species. In addition, our results provide a clear functional analysis of a  LAR  gene  in vivo .

Proanthocyanidin synthesis in Theobroma cacao: genes encoding anthocyanidin synthase, anthocyanidin reductase, and leucoanthocyanidin reductase