Midveins regulate the shape formation of drying leaves

Abstract

Dried leaves in nature often exhibit curled and crumpled morphologies, typically attributed to internal strain gradients that produce dome-like shapes. However, the origin of these strain gradients remains poorly understood. Although leaf veins--particularly the midvein--have been suggested to influence shape formation, their mechanical role has not been systematically investigated. Here, we demonstrate that mechanical constraints imposed by the midvein play a crucial role in generating the diverse morphologies that emerge during leaf drying. Combining numerical simulations and theoretical analysis, we show that a uniformly shrinking leaf lamina constrained by a non-shrinking midvein gives rise to two distinct types of configurations: curling-dominated and folding-dominated morphologies. In the curling-dominated regime, both S-curled and C-curled shapes emerge, with C-curled configurations more commonly observed due to their lower elastic energy. In contrast, the folding-dominated regime features folding accompanied by edge waviness. Theoretical modeling reveals a linear relationship between midvein curvature and mismatch strain, consistent with simulation results. Moreover, we find that the morphological outcome is governed by the ratio of bending stiffnesses between the lamina and the midvein. We construct a comprehensive phase diagram for the transitions between different configurations. These findings provide a mechanical framework for understanding shape formation in drying leaves, offering new insights into natural morphing processes and informing the design of bio-inspired morphable structures.