Biomechanics of the fibrillar adhesive system in insects
Many animals are able to scale smooth surfaces using adhesive structures on their feet. These organs are either soft pads with a relatively smooth surface or dense arrays of microscopic adhesive hairs with both designs having independently evolved in diverse taxa of arthropods and vertebrates. Biological adhesive pads out-perform conventional adhesives in many respects, making them important models for biomimetics. Hairy pads have attracted particular attention, because it has become feasible to fabricate similar synthetic microstructures. Nevertheless, the detailed performance and functional properties have not been characterised for any natural fibrillar adhesive system, and many fundamental aspects are still not understood. The aim of this thesis was therefore to investigate the fibrillar adhesive system of leaf beetles as a model. To investigate the functional implications of hairy pad design, the attachment performance between hairy pads of the leaf beetle Gastrophysa viridula and smooth pads of stick insects (Carausius morosus) was compared. Adhesive and frictional stresses were found to be similar in smooth and hairy pads, inconsistent with contact splitting theory, which predicts higher adhesive stresses for fibrillar adhesives. Hairy pads showed a greater direction-dependence of friction forces than smooth pads, confirming the importance of the asymmetric design of individual setae for effortless detachment. Experiments with contaminating particles also showed that hairy pads removed contamination more rapidly and efficiently than smooth pads. Self-cleaning ability had not been previously documented for adhesive organs of insects. To investigate to what extent the hairy system is able to compensate for surface roughness, whole-body attachment forces were measured for varying roughness levels. Attachment was reduced for all length scales of surface roughness, but in particular for asperity sizes smaller than the diameter of individual seta tips. Leaf beetles possess adhesive pads on three tarsal segments, which vary in setal morphology. However, the functional implications of this variation are unknown. The mechanical and adhesive properties of individual pads were therefore tested and their use during climbing observed. Proximal pads were shown to be stiffer than distal pads, conferring stability during pushing. In contrast, the softer distal pads allowed better attachment to rough surfaces. Hence the morphological variation is explained by an effective division of labour between the pads. To investigate an extreme example of pushing in a hairy system, pad use was studied during jumping in flea beetles. The pushing forces needed during take-off were exclusively produced by the proximal pads, again confirming the division of labour. To characterise the effects of different hair morphologies and to understand how individual setae contribute to array and whole-animal performance, single hair forces were measured using a glass capillary cantilever. Male-specific discoidal hairs were shown to be both stiffer and more adhesive than pointed and spatula-tipped setae, likely affecting overall pad stability and attachment. This thesis has shown that hairy pads are similar to smooth pads in the magnitude of adhesive stress supported yet outperform them in detachability and self-cleaning. It was also demonstrated that there are considerable differences in design and performance even within setal arrays of the same insect, indicating the limitations of general models of fibrillar adhesion and underlining the importance of specialised adaptations.