Jan-Henning Dirks

A comparative study of the effects of vein-joints on the mechanical behaviour of insect wings: I. Single joints

The flight performance of insects is strongly affected by the deformation of the wing during a stroke cycle. Many insects therefore use both active and passive mechanisms to control the deformation of their wings in flight. Several studies have focused on the wing kinematics, and plenty is known about the mechanism of their passive deformability. However, given the small size of the vein-joints, accurate direct mechanical experiments are almost impossible to perform. We therefore developed numerical models to perform a comparative and comprehensive investigation of the mechanical behaviour of the vein-joints under external loading conditions. The results illustrate the effect of the geometry and the presence of the rubberlike protein resilin on the flexibility of the joints. Our simulations further show the contribution of the spikes to the anisotropic flexural stiffness in the dorsal and ventral directions. In addition, our results show that the cross veins, only in one joint type, help to transfer the stress to the thicker longitudinal veins. The deformation pattern and the stress distribution in each vein-joint are discussed in detail. This study provides a strong background for further realistic modelling of the dragonfly wing deformation.

Physical principles of fluid-mediated insect attachment - Shouldn’t insects slip?

Summary Insects use either hairy or smooth adhesive pads to safely adhere to various kinds of surfaces. Although the two types of adhesive pads are morphologically different, they both form contact with the substrate via a thin layer of adhesive fluid. To model adhesion and friction forces generated by insect footpads often a simple “wet adhesion” model is used, in which two flat undeformable substrates are separated by a continuous layer of fluid. This review summarizes the key physical and tribological principles that determine the adhesion and friction in such a model. Interestingly, such a simple wet-adhesion model falls short in explaining several features of insect adhesion. For example, it cannot predict the observed high static friction forces of the insects, which enable them to cling to vertical smooth substrates without sliding. When taking a closer look at the “classic” attachment model, one can see that it is based on several simplifications, such as rigid surfaces or continuous layers of Newtonian fluids. Recent experiments show that these assumptions are not valid in many cases of insect adhesion. Future tribological models for insect adhesion thus need to incorporate deformable adhesive pads, non-Newtonian properties of the adhesive fluid and/or partially “dry” or solid-like contact between the pad and the substrate.

Bridging the gap: wound healing in insects restores mechanical strength by targeted cuticle deposition

If an insect is injured, can it repair its skeleton in a manner which is mechanically strong and viable? Previous work has described the biological processes that occur during repair of insect cuticle, but until now, there has been no biomechanical assessment of the repaired area. We analysed the biomechanics of the injury repair process in the desert locust (Schistocerca gregaria). We show that after an incision, a healing process occurred which almost doubled the mechanical strength of locust tibial cuticle, restoring it to 66% of the original, intact strength. This repair process occurred by targeted cuticle deposition, stimulated by the presence of the injury. The cut surfaces remained unrepaired, but a patch of endocuticle was deposited, reinforcing the area and thus increasing the effective fracture toughness. The deposition rate of endocuticle inside the tibia increased fourfold compared with uninjured controls, but only on the dorsal side, where the incision was placed. The limb is highly loaded during jumping, so this partial restoration of strength will have a profound effect on the fitness of the insect. A finite-element model provided insights into the mechanics of the repair, predicting that the patch material reaches its ultimate strength before the fracture toughness of the existing cuticle is exceeded.

Wing cross veins: an efficient biomechanical strategy to mitigate fatigue failure of insect cuticle

Locust wings are able to sustain millions of cycles of mechanical loading during the lifetime of the insect. Previous studies have shown that cross veins play an important role in delaying crack propagation in the wings. Do cross veins thus also influence the fatigue behaviour of the wings? Since many important fatigue parameters are not experimentally accessible in a small biological sample, here we use the finite element (FE) method to address this question numerically. Our FE model combines a linear elastic material model, a direct cyclic approach and the Paris law and shows results which are in very good agreement with previously reported experimental data. The obtained results of our study show that cross veins indeed enhance the durability of the wings by temporarily stopping cracks. The cross veins further distribute the stress over a larger area and therefore minimize stress concentrations. In addition, our work indicates that locust hind wings have an endurance limit of about 40% of the ultimate tensile strength of the wing material, which is comparable to many engineering materials. The comparison of the results of the computational study with predictions of two most commonly used fatigue failure criteria further indicates that the Goodman criterion can be used to roughly predict the failure of the insect wing. The methodological framework presented in our study could provide a basis for future research on fatigue of insect cuticle and other biological composite structures.

Both stiff and compliant: morphological and biomechanical adaptations of stick insect antennae for tactile exploration

Active tactile exploration behaviour is constrained to a large extent by the morphological and biomechanical properties of the animals somatosensory system. In the model organism Carausius morosus, the main tactile sensory organs are long, thin, seemingly delicate, but very robust antennae. Previous studies have shown that these antennae are compliant under contact, yet stiff enough to maintain a straight shape during active exploration. Overcritical damping of the flagellum, on the other hand, allows for a rapid return to the straight shape after release of contact. Which roles do the morphological and biomechanical adaptations of the flagellum play in determining these special mechanical properties? To investigate this question, we used a combination of biomechanical experiments and numerical modelling. A set of four finite-element (FE) model variants was derived to investigate the effect of the distinct geometrical and material properties of the flagellum on its static (bending) and dynamic (damping) characteristics. The results of our numerical simulations show that the tapered shape of the flagellum had the strongest influence on its static biomechanical behaviour. The annulated structure and thickness gradient affected the deformability of the flagellum to a lesser degree. The inner endocuticle layer of the flagellum was confirmed to be essential for explaining the strongly damped return behaviour of the antenna. By highlighting the significance of two out of the four main structural features of the insect flagellum, our study provides a basis for mechanical design of biomimetic touch sensors tuned to become maximally flexible while quickly resuming a straight shape after contact.