Model for perianth formation in orchids

Orchidaceae, the orchid family under the order Asparagales, contains more than 20,000 accepted species in approximately 880 genera(1-3). In contrast to most flowers of actinomorphic symmetry, orchid flowers typically have zygomorphic symmetry with a striking well-differentiated labellum (lip) that acts as the main pollinator attractant by employing visual, fragrance and tactile cues(4-7). Genetics models controlling patterning formation of actinomorphic flowers, such as Arabidopsis, are well known. However, the mechanisms of sepal/petal/lip determination remain obscure. Here, we demonstrate a conserved principle, called the Perianth (P) code, which involves competition between two protein complexes containing different AP3/AGL6 homologues to determine the formation of the complex perianth patterns in orchids. In the P code, the higher-order heterotetrameric SP (sepal/petal) complex (OAP3-1/OAGL6-1/OAGL6-1/OPI) specifies sepal/petal formation, whereas the L (lip) complex (OAP3-2/OAGL6-2/OAGL6-2/OPI) is exclusively required for lip formation. This model is validated by the conversion of lips into sepal/petal structures in Oncidium and Phalaenopsis orchids through the suppression of the proposed L complex activity in lips using the virus-induced gene silencing (VIGS) strategy. A comprehensive examination of four different subfamilies of Orchidaceae further validates the P code and significantly extends the current knowledge regarding the mechanism and pathways of perianth formation in orchids.