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

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.