Within the framework of a biomimetic top-down approach, our study started with the technical question of the development of a hinge-free and compliant actuator inspired by plant movements. One meaningful biological concept generator was the opening and closing movements of the leaf halves of grasses. Functional morphological investigations were carried out on the selected model plant Sesleria nitida . The results formed the basis for further clarifying the functional movement principle with a particular focus on the role of turgor changes in bulliform cells on kinetic amplification. All findings gained from the investigations of the biological model were incorporated into a finite-element analysis, as a prerequisite for the development of a pneumatic cellular actuator. The first prototype consisted of a row of single cells positioned on a plate. The cells were designed in such a way that the entire structure bent when the pneumatic pressure applied to each individual cell was increased. The pneumatic cellular actuator thus has the potential for applications on an architectural scale. It has subsequently been integrated into the midrib of the facade shading system Flectofold in which the bending of its midrib controls the hoisting of its wings.
Mechanical optimisation plays a key role in living beings either as an immediate response of individuals or as an evolutionary adaptation of populations to changing environmental conditions. Since biological structures are the result of multifunctional evolutionary constraints, the dimensionless twist-to-bend ratio is particularly meaningful because it provides information about the ratio of flexural rigidity to torsional rigidity determined by both material properties (bending and shear modulus) and morphometric parameters (axial and polar second moment of area). The determination of the mutual contributions of material properties and structural arrangements (geometry) or their ontogenetic alteration to the overall mechanical functionality of biological structures is difficult. Numerical methods in the form of gradient flows of phase field functionals offer a means of addressing this question and of analysing the influence of the cross-sectional shape of the main load-bearing structures on the mechanical functionality. Three phase field simulations were carried out showing good agreement with the cross-sections found in selected plants: (i) U-shaped cross-sections comparable with those of Musa sp. petioles, (ii) star-shaped cross-sections with deep grooves as can be found in the lianoid wood of Condylocarpon guianense stems, and (iii) flat elliptic cross-sections with one deep groove comparable with the cross-sections of the climbing ribbon-shaped stems of Bauhinia guianensis.
10 December 2021 in Journal of Experimental Botany
Plants are exposed to various environmental stresses. Leaves immediately respond to mechano-stimulation, such as wind and touch, by bending and twisting or acclimate over a longer time period by thigmomorphogenetic changes of mechanical and geometrical properties. We selected the peltate leaves of Pilea peperomioides for a comparative analysis of mechano-induced effects on morphology, anatomy, and biomechanics of petiole and transition zone. The plants were cultivated for 6 weeks in a phytochamber divided into four treatment groups: control (no stimulus), touch stimulus (brushing every 30 s), wind stimulus (constant air flow of 4.6 m s−1), and a combination of touch and wind stimuli. Comparing the four treatment groups, neither the petiole nor the transition zone showed significant thigmomorphogenetic acclimations. However, comparing the petiole and the transition zone, the elastic modulus (E), the torsional modulus (G), the E/G ratio, and the axial rigidity (EA) differed significantly, whereas no significant difference was found for the torsional rigidity (GK). The twist-to-bend ratios (EI/GK) of all petioles ranged between 4.33 and 5.99, and of all transition zones between 0.67 and 0.78. Based on the twist-to-bend ratios, we hypothesize that bending loads are accommodated by the petiole, while torsional loads are shared between the transition zone and petiole.
The ability to perform hygroscopic movements has evolved in many plant lineages and relates to a multitude of different functions such as seed burial, flower protection or regulation of diaspore release. In most mosses, spore release is controlled by hygroscopic movements of the peristome teeth and also of the spore capsule. Our study presents, for the first time, temporally and spatially well-resolved kinematic analyses of these complex shape changes in response to humidity conditions and provides insights into the sophisticated functional morphology and anatomy of the peristome teeth. In Brachythecium populeum the outer teeth of the peristome perform particularly complex hygroscopic movements during hydration and desiccation. Hydration induces fast inward dipping followed by partial re-straightening of the teeth. In their final shape, wet teeth close the capsule. During desiccation, the teeth perform an outward flicking followed by a re-straightening which opens the capsule. We present a kinematic analysis of these shape changes and of the underlying functional anatomy of the teeth. These teeth are shown to be composed of two layers which show longitudinal gradients in their material composition, structure and geometry. We hypothesize that these gradients result in (i) differences in swelling/shrinking capacity and velocity between the two layers composing the teeth, and in (ii) a gradient of velocity of swelling and shrinking from the tip to the base of the teeth. We propose these processes explain the observed movements regulating capsule opening or closing. This hypothesis is corroborated by experiments with isolated layers of peristome teeth. During hydration and desiccation, changes to the shape and mass of the whole spore capsule accompany the opening and closing. Results are discussed in relation to their significance for humidity-based regulation of spore release.
Although both the petiole and lamina of foliage leaves have been thoroughly studied, the transition zone between them has often been overlooked. We aimed to identify objectively measurable morphological and anatomical criteria for a generally valid definition of the petiole–lamina transition zone by comparing foliage leaves with various body plans (monocotyledons vs. dicotyledons) and spatial arrangements of petiole and lamina (two-dimensional vs. three-dimensional configurations). Cross-sectional geometry and tissue arrangement of petioles and transition zones were investigated via serial thin-sections and µCT. The changes in the cross-sectional geometries from the petiole to the transition zone and the course of the vascular bundles in the transition zone apparently depend on the spatial arrangement, while the arrangement of the vascular bundles in the petioles depends on the body plan. We found an exponential acropetal increase in the cross-sectional area and axial and polar second moments of area to be the defining characteristic of all transition zones studied, regardless of body plan or spatial arrangement. In conclusion, a variety of terms is used in the literature for describing the region between petiole and lamina. We prefer the term “petiole–lamina transition zone” to underline its three-dimensional nature and the integration of multiple gradients of geometry, shape, and size.
From a mechanical viewpoint, petioles of foliage leaves are subject to contradictory mechanical requirements. High flexural rigidity guarantees support of the lamina and low torsional rigidity ensures streamlining of the leaves in wind. This mechanical trade-off between flexural and torsional rigidity is described by the twist-to-bend ratio. The safety factor describes the maximum load capacity. We selected four herbaceous species with different body plans (monocotyledonous, dicotyledonous) and spatial configurations of petiole and lamina (2-dimensional, 3-dimensional) and carried out morphological-anatomical studies, two-point bending tests and torsional tests on the petioles to analyze the influence of geometry, size and shape on their twist-to-bend ratio and safety factor. The monocotyledons studied had significantly higher twist-to-bend ratios (23.7 and 39.2) than the dicotyledons (11.5 and 13.3). High twist-to-bend ratios can be geometry-based, which is true for the U-profile of Hosta x tardiana with a ratio of axial second moment of area to torsion constant of over 1.0. High twist-to-bend ratios can also be material-based, as found for the petioles of Caladium bicolor with a ratio of bending elastic modulus and torsional modulus of 64. The safety factors range between 1.7 and 2.9, meaning that each petiole can support about double to triple the leaf’s weight.