Extensive collection of femtoliter pad secretion droplets in beetle Leptinotarsa decemlineata allows nanoliter microrheology

Pads of beetles are covered with long, deformable setae, each ending in a micrometric terminal plate coated with secretory fluid. It was recently shown that the layer of the pad secretion covering the terminal plates is responsible for the generation of strong attractive forces. However, less is known about the fluid itself because it is produced in extremely small quantity. We here present a first experimental investigation of the rheological properties of the pad secretion in the Colorado potato beetle {\it Leptinotarsa decemlineata}. Because the secretion is produced in an extremely small amount at the level of the terminal plate, we first develop a procedure based on capillary effects to collect the secretion. We then manage to incorporate micrometric beads, initially in the form of a dry powder, and record their thermal motion to determine the mechanical properties of the surrounding medium. We achieve such a quantitative measurement within the collected volume, much smaller than the $1 {\rm \mu}$l sample volume usually required for this technique. Surprisingly, the beetle secretion was found to behave as a purely viscous liquid, of high viscosity. This suggests that no specific complex fluid behaviour is needed during beetle locomotion. We build a scenario for the contact formation between the spatula at the setal tip and a substrate, during the insect walk. We show that the attachment dynamics of the insect pad computed from the high measured viscosity is in good agreement with observed insect pace. We finally discuss the consequences of the secretion viscosity on the insect adhesion

Evolution of locomotory attachment pads of hexapods

This study shows that, in their evolution, hexapods have convergently developed two distinctly different mechanisms to attach themselves to a variety of substrates during locomotion. The first mechanism is provided by hairy surfaces and the second one by smooth flexible pads. The main similarity of both mechanisms is that the structured pad surface or particular properties of pad materials guarantee a maximum real contact with diverse substrata, regardless of their microsculpture. Ten characters of the two alternative designs were coded and analyzed together with a data matrix containing 105 additional morphological characters of different stages and body parts. The analysis demonstrates that similar structures (arolium, euplantulae, hairy tarsomeres) have evolved independently in several hexapod lineages. The evolution of flight and the associated necessity of being able to cling to vegetation or other substrates are suggested to be major triggers for the evolution of attachment structures

Structural design and biomechanics of friction-based releasable attachment devices in insects

Design of attachment devices in insects varies enormously in relation to different functional loads. Many systems, located on different parts of the body, involve surfaces with particular frictional properties. Such systems evolved to attach parts of the body to each other, or to attach an insect to the substratum by providing fast and reversible attachment/detachment. Among these systems, there are some that deal with predefined surfaces, and others, in which one surface remains unpredictable. The first type of system occurs, for example, in wing-locking devices and head-arresting systems and is called probabilistic fasteners. The second type is mainly represented by insect attachment pads of two alternative designs: hairy and smooth. The relationship between surface patterns and/or mechanical properties of materials of contact pairs results in two main working principles of the frictional devices: mechanical interlocking, or maximization of the contact area. We give an overview of the functional design of two main groups of friction-based attachment devices in insects: probabilistic fasteners and attachment pads