Evolution of CYCLOIDEA-like genes in Fabales: Insights into duplication patterns and the control of floral symmetry.

Cycloidea-like (CYC-like) genes are the key regulatory factors in the development of flower symmetry. Duplication and/or reduction of CYC-like genes have occurred several times in various angiosperm groups and are hypothesized to be correlated with the evolution of flower symmetry, which in turn has contributed to the evolutionary success of these groups. However, less is known about the evolutionary scenario of CYC-like genes in the whole Fabales, which contains four families with either symmetric or actinomorphic flowers. Here we investigated the evolution of CYC-like genes in all the four families of Fabales and recovered one to nine CYC-like genes (CYC1, CYC2, and CYC3) depending on which lineages, but the CYC3 genes were most likely lost in the ancestor of Leguminosae. Phylogenetic analysis suggested that the CYC-like genes could have undergone multiple duplications and losses in different plant lineages and formed distinct paralogous/orthologous clades. The ancestor of the Papilionoideae and Caesalpinioideae may possess two paralogs of CYC1 genes but one of them was subsequently lost in Papilionoideae and was retained only in several species of Caesalpinioideae. CYC2 genes were more frequently duplicated in Papilionoideae than in other legumes. We propose that the diversification patterns of both CYC1 and CYC2 genes are not related to the floral symmetry in non-papilionoid Fabales groups, however, gene duplication and functional divergence of CYC2 are essential for the floral zygomorphy of Papilionoideae. This is the first systematic analysis of the CYC-like genes in Fabales and could form the basis for further study of molecular mechanisms controlling floral symmetry in non-model plants of Fabales.