Ingo Scholz

Slippery surfaces of pitcher plants: Nepenthes wax crystals minimize insect attachment via microscopic surface roughness

SUMMARY Pitcher plants of the genus Nepenthes efficiently trap and retain insect prey in highly specialized leaves. Besides a slippery peristome which inhibits adhesion of insects they employ epicuticular wax crystals on the inner walls of the conductive zone of the pitchers to hamper insect attachment by adhesive devices. It has been proposed that the detachment of individual crystals and the resulting contamination of adhesive organs is responsible for capturing insects. However, our results provide evidence in favour of a different mechanism, mainly based on the stability and the roughness of the waxy surface. First, we were unable to detect a large quantity of crystal fragments on the pads of insects detached from mature pitcher surfaces of Nepenthes alata. Second, investigation of the pitcher surface by focused ion beam treatment showed that the wax crystals form a compact 3D structure. Third, atomic force microscopy of the platelet-shaped crystals revealed that the crystals are mechanically stable, rendering crystal detachment by insect pads unlikely. Fourth, the surface profile parameters of the wax layer showed striking similarities to those of polishing paper with low grain size. By measuring friction forces of insects on this artificial surface we demonstrate that microscopic roughness alone is sufficient to minimize insect attachment. A theoretical model shows that surface roughness within a certain length scale will prevent adhesion by being too rough for adhesive pads but not rough enough for claws.

Determination of the Young's modulus of the epicuticle of the smooth adhesive organs of Carausius morosus using tensile testing

Adhesive organs like arolia of insects allow these animals to climb on different substrates by creating high adhesion forces. According to the Dahlquist criterion, adhesive organs must be very soft, exhibiting an effective Youngs modulus of below 100 kPa to adhere well to substrates. Such a low effective Youngs modulus allows the adhesive organs to make almost direct contact with the substrate and results in van der Waals forces along with capillary forces. In previous studies, the effective Youngs moduli of adhesive organs were determined using indentation tests, revealing their structure to be very soft. However, adhesive organs show a layered structure, thus the measured values comprise the effective Youngs moduli of several layers of the adhesive organs. In this study, a new approach is illustrated to measure the Youngs modulus of the outermost layer of the arolium, i.e. of the epicuticle, of the stick insect Carausius morosus. As a result of the epicuticle being supported by upright fibres, tensile tests allow the determination of the Youngs modulus of the epicuticle with hardly influence from subjacent layers. In our tensile tests, arolia of stick insects adhering on a latex membrane were stretched by stretching the membrane while the elongation of the contact area between an arolium and the membrane was recorded. For analysis, mathematical models of the mechanical system were developed. When fed with the observed elongations, these models yield estimates for the Youngs modulus of the epicuticle of approximately 100 MPa. Thus, in arolia, a very thin layer (~225 nm) of a rather stiff material, which is less susceptible to abrasion, makes contact with the substrates, whereas the inner fibrous structure of arolia is responsible for their softness.

Cribellate thread production in spiders: Complex processing of nano-fibres into a functional capture thread

Spider silk production has been studied intensively in the last years. However, capture threads of cribellate spiders employ an until now often unnoticed alternative of thread production. This thread in general is highly interesting, as it not only involves a controlled arrangement of three types of threads with one being nano-scale fibres (cribellate fibres), but also a special comb-like structure on the metatarsus of the fourth leg (calamistrum) for its production. We found the cribellate fibres organized as a mat, enclosing two parallel larger fibres (axial fibres) and forming the typical puffy structure of cribellate threads. Mat and axial fibres are punctiform connected to each other between two puffs, presumably by the action of the median spinnerets. However, this connection alone does not lead to the typical puffy shape of a cribellate thread. Removing the calamistrum, we found a functional capture thread still being produced, but the puffy shape of the thread was lost. Therefore, the calamistrum is not necessary for the extraction or combination of fibres, but for further processing of the nano-scale cribellate fibres. Using data from Uloborus plumipes we were able to develop a model of the cribellate thread production, probably universally valid for cribellate spiders.

Functional morphology of the adhesive organs of stick insects( Carausius morosus )

Adhesive organs enable insects to cling to various substrates. During locomotion, a very fast but reliable change of adhesion and detachment is realised. To reveal the detailed underlying mechanisms of this impressive performance, we analysed the ultrastructure of adhesive organs of the stick insect C. morosus using scanning electron microscopy (SEM), transmission electron microscopy (TEM) and confocal laser scanning microscopy (CLSM).

Morphological adaptation of the calamistrum to the cribellate spinning process in Deinopoidae (Uloboridae, Deinopidae)

Spiders are famous for their silk with fascinating mechanical properties. However, some can further produce, process and handle nano fibres, which are used as capture threads. These ‘cribellate spiders’ bear a specialized setae comb on their metatarsus (calamistrum), which modifies cribellate nano fibres to assemble a puffy structure within the capture thread. Among different species, the calamistrum morphology can differ remarkably. Although a model of thread production has been established for Uloborus plumipes, it is not resolved if/how different shaped calamistra influence the production process. We were able to transfer the model without restrictions to spiders with different shaped calamistra. Fibres are not locked between setae but are passing across a rather smooth surface-like area on the calamistrum. This area can be relocated, explaining the first morphological difference between calamistra, without changing the influence of the calamistrum on fibres. By performing an elongated leg movement, contact between fibres and calamistrum could be adjusted after finishing thread production. This movement has to bring the thread in contact with the second morphological peculiarity: cribellate teeth. We suggest these teeth are used to handle the thread independently of the spinnerets, a feature only necessary for spiders, which do not move during web construction.