Hook Shape of Growing Leaves Results from an Active Regulation

Abstract

The rachis of most growing compound leaves observed in nature exhibit a stereotyped hook shape. In this study, we focus on the canonical case of Averrhoa carambola. Combining kinematics and mechanical investigation, we characterize this hook shape and shed light on its establishment and maintenance. We show quantitatively that the hook shape is a conserved bent zone propagating at constant velocity and constant distance from the apex throughout development. A simple mechanical test first reveals non-zero spontaneous curvature profiles for the growing leaves, indicating that the hook shape is actively regulated. It then evidences the robust spatial organization of growth, curvature, rigidity and lignification and their interplay. Regulation processes appear to be specifically localized: in particular, differential growth occurs where the elongation rate drops. Finally, impairing the graviception of the leaf on a clinostat led to reduced hook curvatures but not to its loss. Altogether our results suggest a role for proprioception in the regulation of the apical hook, likely mediated via mechanical strain.