Smooth or with a Snap! Biomechanics of Trap Reopening in the Venus Flytrap (Dionaea muscipula).

Fast snapping in the carnivorous Venus flytrap (Dionaea muscipula) involves trap lobe bending and abrupt curvature inversion (snap-buckling), but how do these traps reopen? Here, the trap reopening mechanics in two different D. muscipula clones, producing normal-sized (N traps, max. ≈3 cm in length) and large traps (L traps, max. ≈4.5 cm in length) are investigated. Time-lapse experiments reveal that both N and L traps can reopen by smooth and continuous outward lobe bending, but only L traps can undergo smooth bending followed by a much faster snap-through of the lobes. Additionally, L traps can reopen asynchronously, with one of the lobes moving before the other. This study challenges the current consensus on trap reopening, which describes it as a slow, smooth process driven by hydraulics and cell growth and/or expansion. Based on the results gained via three-dimensional digital image correlation (3D-DIC), morphological and mechanical investigations, the differences in trap reopening are proposed to stem from a combination of size and slenderness of individual traps. This study elucidates trap reopening processes in the (in)famous Dionaea snap traps - unique shape-shifting structures of great interest for plant biomechanics, functional morphology, and applications in biomimetics, i.e., soft robotics.

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.

Development of a Material Design Space for 4D-Printed Bio-Inspired Hygroscopically Actuated Bilayer Structures with Unequal Effective Layer Widths.

(1) Significance of geometry for bio-inspired hygroscopically actuated bilayer structures is well studied and can be used to fine-tune curvatures in many existent material systems. We developed a material design space to find new material combinations that takes into account unequal effective widths of the layers, as commonly used in fused filament fabrication, and deflections under self-weight. (2) For this purpose, we adapted Timoshenko's model for the curvature of bilayer strips and used an established hygromorphic 4D-printed bilayer system to validate its ability to predict curvatures in various experiments. (3) The combination of curvature evaluation with simple, linear beam deflection calculations leads to an analytical solution space to study influences of Young's moduli, swelling strains and densities on deflection under self-weight and curvature under hygroscopic swelling. It shows that the choice of the ratio of Young's moduli can be crucial for achieving a solution that is stable against production errors. (4) Under the assumption of linear material behavior, the presented development of a material design space allows selection or design of a suited material combination for application-specific, bio-inspired bilayer systems with unequal layer widths.

Snapping mechanics of the Venus flytrap (Dionaea muscipula).

The mechanical principles for fast snapping in the iconic Venus flytrap are not yet fully understood. In this study, we obtained time-resolved strain distributions via three-dimensional digital image correlation (DIC) for the outer and inner trap-lobe surfaces throughout the closing motion. In combination with finite element models, the various possible contributions of the trap tissue layers were investigated with respect to the trap's movement behavior and the amount of strain required for snapping. Supported by in vivo experiments, we show that full trap turgescence is a mechanical-physiological prerequisite for successful (fast and geometrically correct) snapping, driven by differential tissue changes (swelling, shrinking, or no contribution). These are probably the result of the previous accumulation of internal hydrostatic pressure (prestress), which is released after trap triggering. Our research leads to an in-depth mechanical understanding of a complex plant movement incorporating various actuation principles.

Structural stress response of segmented natural shells: a numerical case study on the clypeasteroid echinoid Echinocyamus pusillus.

The skeleton of Echinocyamus pusillus is considered as an exceptional model organism for structural strength and skeletal integrity within the echinoids as demonstrated by the absence of supportive collagenous fibres between single plates and the high preservation potential of their skeletons. The structural principles behind this remarkably stable, multi-plated, light-weight construction remain hardly explored. In this study, high-resolution X-ray micro-computed tomography, finite-element analysis and physical crushing tests are used to examine the structural mechanisms of this echinoid's skeleton. The virtual model of E. pusillus shows that the material is heterogeneously distributed with high material accumulations in the internal buttress system and at the plate boundaries. Finite-element analysis indicates that the heterogeneous material distribution has no effect on the skeleton's strength. This numerical approach also demonstrates that the internal buttress system is of high significance for the overall skeletal stability of this flattened echinoid. Results of the finite-element analyses with respect to the buttress importance were evaluated by physical crushing tests. These uniaxial compression experiments support the results of the simulation analysis. Additionally, the crushing tests demonstrate that organic tissues do not significantly contribute to the skeletal stability. The strength of the echinoid shell, hence, predominantly relies on the structural design.

How the carnivorous waterwheel plant (Aldrovanda vesiculosa) snaps.

The fast motion of the snap-traps of the terrestrial Venus flytrap (Dionaea muscipula) have been intensively studied, in contrast to the tenfold faster underwater snap-traps of its phylogenetic sister, the waterwheel plant (Aldrovanda vesiculosa). Based on biomechanical and functional-morphological analyses and on a reverse biomimetic approach via mechanical modelling and computer simulations, we identify a combination of hydraulic turgor change and the release of prestress stored in the trap as essential for actuation. Our study is the first to identify and analyse in detail the motion principle of Aldrovanda, which not only leads to a deepened understanding of fast plant movements in general, but also contributes to the question of how snap-traps may have evolved and also allows for the development of novel biomimetic compliant mechanisms.