Avoiding unfavorable situations is a vital skill and a constant task for any animal. Situations can be unfavorable because they feature something that the animal wants to escape from, or because they do not feature something that it seeks to obtain. We investigate whether the microbehavioral mechanisms by which these two classes of aversion come about are shared or distinct. We find that larval Drosophila avoid odors either previously associated with a punishment, or previously associated with the lack of a reward. These two classes of conditioned aversion are found to be strikingly alike at the microbehavioral level. In both cases larvae show more head casts when oriented toward the odor source than when oriented away, and direct fewer of their head casts toward the odor than away when oriented obliquely to it. Thus, conditioned aversion serving two qualitatively different functions-escape from a punishment or search for a reward-is implemented by the modulation of the same microbehavioral features. These features also underlie conditioned approach, albeit with opposite sign. That is, the larvae show conditioned approach toward odors previously associated with a reward, or with the lack of a punishment. In order to accomplish both these classes of conditioned approach the larvae show fewer head casts when oriented toward an odor, and direct more of their head casts toward it when they are headed obliquely. Given that the Drosophila larva is a genetically tractable model organism that is well suited to study simple circuits at the single-cell level, these analyses can guide future research into the neuronal circuits underlying conditioned approach and aversion, and the computational principles of conditioned search and escape.
Peltate- or umbrella- shaped leaves are characterised by a petiole more or less centrally attached to the lamina on the abaxial side. The transition from the petiole to lamina in peltate leaves resembles a significant and abrupt geometrical change from a beam to a plate in a very compact shape. Since these leaves have not been subject of many studies, the distribution of that specific leaf morphology in the plant kingdom was investigated. Furthermore, the connection between the petiole and lamina of several peltate species was studied anatomically and morphologically, focusing on the reinforcing fibre strands. We found peltate leaves in 357 species representing 25 orders, 40 families and 99 genera. The majority are herbaceous perennials growing in shady, humid to wet habitats mainly distributed in the subtropical–tropical zones. Detailed anatomical investigation of 41 species revealed several distinct principles of how the transition zone between the petiole and lamina is organised. In-depth analysis of these different types accompanied by finite element-modelling could serve as inspiration for supporting structures in lightweight construction.
Stephania japonica is a slender climbing plant with peltate, triangular-ovate leaves. Not many research efforts have been devoted to investigate the anatomy and the mechanical properties of this type of leaf shape. In this study, displacement driven tensile tests with three cycles on different displacement levels are performed on petioles, venation and intercostal areas of the Stephania japonica leaves. Furthermore, compression tests in longitudinal direction are performed on petioles. The mechanical experiments are combined with light microscopy and X-ray tomography. The experiments show, that these plant organs and tissues behave in the finite strain range in a viscoelastic manner. Based on the results of the light microscopy and X-ray tomography, the plant tissue can be considered as a matrix material reinforced by fibers. Therefore, a continuum mechanical anisotropic viscoelastic material model at finite deformations is proposed to model such behavior. The anisotropy is specified as the so-called transverse isotropy, where the behavior in the plane perpendicular to the fibers is assumed to be isotropic. The model is obtained by postulating a Helmholtz free energy, which is split additively into an elastic and an inelastic part. Both parts of the energy depend on structural tensors to account for the transversely isotropic material behavior. The evolution equations for the internal variables, e.g. inelastic deformations, are chosen in a physically meaningful way that always fulfills the second law of thermodynamics. The proposed model is calibrated against experimental data, and the material parameters are identified. The model can be used for finite element simulations of this type of leaf shape, which is left open for the future work.
Plants are exposed to various external stresses influencing physiology, anatomy, and morphology. Shape, geometry, and size of shoots and leaves are particularly affected. Among the latter, peltate leaves are not very common and so far, only few studies focused on their properties. In this case study, four Begonia species with different leaf shapes and petiole attachment points were analyzed regarding their leaf morphology, anatomy, and biomechanical properties. One to two plants per species were examined. In all four species, the petiole showed differently sized vascular bundles arranged in a peripheral ring and subepidermal collenchyma. These anatomical characteristics, low leaf dry mass, and low amount of lignified tissue in the petiole point toward turgor pressure as crucial for leaf stability. The petiole-lamina transition zone shows a different organization in leaves with a more central (peltate) and lateral petiole insertion. While in non-peltate leaves simple fiber branching is present, peltate leaves show a more complex reticulate fiber arrangement. Tensile and bending tests revealed similar structural Young’s moduli in all species for intercostal areas and venation, but differences in the petiole. The analysis of the leaves highlights the properties of petiole and the petiole-lamina transition zone that are needed to resist external stresses.
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