Orchestrated Movement Sequences and Shape-Memory-like Effects in Pine Cones.

Hygroscopic seed-scale movement is responsible for the weather-adaptive opening and closing of pine cones and for facilitating seed dispersal under favorable environmental conditions. Although this phenomenon has long been investigated, many involved processes are still not fully understood. To gain a deeper mechanical and structural understanding of the cone and its functional units, namely the individual seed scales, we have investigated their desiccation- and wetting-induced movement processes in a series of analyses and manipulative experiments. We found, for example, that the abaxial scale surface is responsible for the evaporation of water from the closed cone and subsequent cone opening. Furthermore, we tested the capability of dry and deformed scales to restore their original shape and biomechanical properties by wetting. These results shed new light on the orchestration of scale movement in cones and the involved forces and provide information about the functional robustness and resilience of cones, leading to a better understanding of the mechanisms behind hygroscopic pine cone opening, the respective ecological framework, and, possibly, to the development of smart biomimetic actuators.

The Structural and Mechanical Basis for Passive-Hydraulic Pine Cone Actuation.

The opening and closing of pine cones is based on the hygroscopic behavior of the individual seed scales around the cone axis, which bend passively in response to changes in environmental humidity. Although prior studies suggest a bilayer architecture consisting of lower actuating (swellable) sclereid and upper restrictive (non- or lesser swellable) sclerenchymatous fiber tissue layers to be the structural basis of this behavior, the exact mechanism of how humidity changes are translated into global movement are still unclear. Here, the mechanical and hydraulic properties of each structural component of the scale are investigated to get a holistic picture of their functional interplay. Measurements of the wetting behavior, water uptake, and mechanical measurements are used to analyze the influence of hydration on the different tissues of the cone scales. Furthermore, their dimensional changes during actuation are measured by comparative micro-computed tomography (µ-CT) investigations of dry and wet scales, which are corroborated and extended by 3D-digital image correlation-based displacement and strain analyses, biomechanical testing of actuation force, and finite element simulations. Altogether, a model allowing a detailed mechanistic understanding of pine cone actuation is developed, which is a prime concept generator for the development of biomimetic hygromorphic systems.