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There are several types of capitulum in the Asteraceae due to different combinations of florets varying in corolla shape and stamen development. Previous studies have shown that the formation of ray florets on a radiate capitulum may be related to the parallel co-option of  CYC2 -like genes among independent Asteraceae lineages. The present work tests that hypothesis and attempts to shed light on the pattern of evolution of the Asteraceae capitulum and floral heteromorphism under the regulation of  CYC2 -like genes. In this study, the evolutionary history of  CYC2 -like genes in the Asterales was reconstructed and their expression patterns were examined in species representing different capitulum types and several major Asteraceae lineages. To clarify the role of CYC2d clade genes in morphogenesis of ray flowers, overexpression of  ClCYC2d  was conducted in  Chrysanthemum lavandulifolium . Our results show that there are six  CYC2 -like members in the Asteraceae; they are results of five duplication events starting from a single-copy gene in the common ancestor of the Goodeniaceae-Calyceraceae-Asteraceae group and completing before the divergence of the subfamily Carduoideae of Asteraceae. Spatial expression pattern of each of the Asteraceae  CYC2 -like members is conserved across the family. All the six members contribute to the development of the complexity of a capitulum: To form a ray floret, either  CYC2c  or  CYC2g  plays an essential role, while  CYC2d  represses the development of dorsal corolla lobes and stamens of the floret. In sum, the developmental program of making a ray flower is conserved involving functionally divergent  CYC2 -like genes. Based on extensive species sampling, this study provides an overview of the mode of regulation of  CYC2 -like genes that patterns the capitulum architectures and their transitions.

Patterning the Asteraceae Capitulum: Duplications and Differential Expression of the Flower Symmetry CYC2-Like Genes