A molecular framework underlying the compound leaf pattern of Medicago truncatula

Compound leaves show more complex patterns than simple leaves, and this is mainly because of a specific morphogenetic process (leaflet initiation and arrangement) that occurs during their development. How the relevant morphogenetic activity is established and modulated to form a proper pattern of leaflets is a central question. Here we show that the trifoliate leaf pattern of the model leguminous plant Medicago truncatula is controlled by the BEL1-like homeodomain protein PINNATE-LIKE PENTAFOLIATA1 (PINNA1). We identify PINNA1 as a determinacy factor during leaf morphogenesis that directly represses transcription of the LEAFY (LFY) orthologue SINGLE LEAFLET1 (SGL1), which encodes an indeterminacy factor key to the morphogenetic activity maintenance. PINNA1 functions alone in the terminal leaflet region and synergizes with another determinacy factor, the C2H2 zinc finger protein PALMATE-LIKE PENTAFOLIATA1 (PALM1), in the lateral leaflet regions to define the spatiotemporal expression of SGL1, leading to an elaborate control of morphogenetic activity. This study reveals a framework for trifoliate leaf-pattern formation and sheds light on mechanisms generating diverse leaf forms.