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.

During biological evolution, plants have developed a wide variety of body plans and concepts that enable them to adapt to changing environmental conditions. The trade-off between flexural and torsional rigidity is an important example of sometimes conflicting mechanical requirements, the adaptation to which can be quantified by the dimensionless twist-to-bend ratio. Our study considers the triangular flower stalk of Carex pendula, which shows the highest twist-to-bend ratios ever measured for herbaceous plant axes. For an in-depth understanding of this peak value, we have developed geometric models reflecting the 2D setting of triangular cross-sections comprised of a parenchymatous matrix with vascular bundles surrounded by an epidermis. We analysed the mathematical models (using finite elements) to measure the effect of either reinforcements of the epidermal tissue or fibre reinforcements such as collenchyma and sclerenchyma on the twist-to-bend ratio. The change from an epidermis to a covering tissue of corky periderm increases both the flexural and the torsional rigidity and decreases the twist-to-bend ratio. Furthermore, additional individual fibre reinforcement strands located in the periphery of the cross-section and embedded in a parenchymatous ground tissue lead to a strong increase of the flexural and a weaker increase of the torsional rigidity and thus resulted in a marked increase of the twist-to-bend ratio. Within the developed model, a reinforcement by 49 sclerenchyma fibre strands or 24 collenchyma fibre strands is optimal in order to achieve high twist-to-bend ratios. Dependent on the mechanical quality of the fibres, the twist-to-bend ratio of collenchyma-reinforced axes is noticeably smaller, with collenchyma having an elastic modulus that is approximately 20 times smaller than that of sclerenchyma. Based on our mathematical models, we can thus draw conclusions regarding the influence of mechanical requirements on the development of plant axis geometry, in particular the placement of reinforcements.

During biological evolution, plants have developed a wide variety of body plans and concepts that enable them to react or adapt to changing environmental conditions. Their morphological-anatomical and mechanical adaptations to conflicting conditions are especially interesting. A good example is the trade-off between flexural and torsional rigidity, as represented by the dimensionless twist-to-bend ratio. We have developed geometric models of a plant tissue reflecting the 2D situation of triangular cross-sections comprising of a parenchymatous matrix with vascular bundles surrounded by an epidermis and analysed them by using mathematical models (finite element analysis) to measure the effect of either reinforcements of the epidermal tissue or fibre reinforcements such as collenchyma and sclerenchyma on the twist-to-bend ratio. The change from an epidermis to a covering tissue of corky periderm increases both the flexural and the torsional rigidity and decreases the twist-to-bend ratio. Furthermore, additional fibre reinforcement strands in a parenchymatous ground tissue lead to a strong increase of the flexural and a weaker increase of the torsional rigidity and thus resulting in a marked increase of the twist-to-bend ratio. Within the developed model, a reinforcement by 49 sclerenchyma fibre strands or 24 collenchyma fibre strands is optimal in order to achieve high twist-to-bend ratios. Dependent on the mechanical quality of the fibres, the twist-to-bend ratio of collenchyma-reinforced axes is noticeably smaller, with collenchyma having an elastic modulus that is approximately 20 times smaller than that of sclerenchyma.

During the evolution of land plants many body plans have been developed. Differences in the cross-sectional geometry and tissue pattern of plant axes influence their flexural rigidity, torsional rigidity and the ratio of both of these rigidities, the so-called twist-to-bend ratio. For comparison, we have designed artificial cross-sections with various cross-sectional geometries and patterns of vascular bundles, collenchyma or sclerenchyma strands, but fixed percentages for these tissues. Our mathematical model allows the calculation of the twist-to-bend ratio by taking both cross-sectional geometry and tissue pattern into account. Each artificial cross-section was placed into a rigidity chart to provide information about its twist-to-bend ratio. In these charts, artificial cross-sections with the same geometry did not form clusters, whereas those with similar tissue patterns formed clusters characterized by vascular bundles, collenchyma or sclerenchyma arranged as one central strand, as a peripheral closed ring or as distributed individual strands. Generally, flexural rigidity increased the more the bundles or fibre strands were placed at the periphery. Torsional rigidity decreased the more the bundles or strands were separated and the less that they were arranged along a peripheral ring. The calculated twist-to-bend ratios ranged between 0.85 (ellipse with central vascular bundles) and 196 (triangle with individual peripheral sclerenchyma strands).