Functionally different pads on the same foot allow control of attachment: stick insects have load-sensitive "heel" pads for friction and shear-sensitive "toe" pads for adhesion.

Stick insects (Carausius morosus) have two distinct types of attachment pad per leg, tarsal "heel" pads (euplantulae) and a pre-tarsal "toe" pad (arolium). Here we show that these two pad types are specialised for fundamentally different functions. When standing upright, stick insects rested on their proximal euplantulae, while arolia were the only pads in surface contact when hanging upside down. Single-pad force measurements showed that the adhesion of euplantulae was extremely small, but friction forces strongly increased with normal load and coefficients of friction were [Formula: see text] 1. The pre-tarsal arolium, in contrast, generated adhesion that strongly increased with pulling forces, allowing adhesion to be activated and deactivated by shear forces, which can be produced actively, or passively as a result of the insects' sprawled posture. The shear-sensitivity of the arolium was present even when corrected for contact area, and was independent of normal preloads covering nearly an order of magnitude. Attachment of both heel and toe pads is thus activated partly by the forces that arise passively in the situations in which they are used by the insects, ensuring safe attachment. Our results suggest that stick insect euplantulae are specialised "friction pads" that produce traction when pressed against the substrate, while arolia are "true" adhesive pads that stick to the substrate when activated by pulling forces.

Surface contact and design of fibrillar 'friction pads' in stick insects (Carausius morosus): mechanisms for large friction coefficients and negligible adhesion.

Many stick insects and mantophasmids possess tarsal 'heel pads' (euplantulae) covered by arrays of conical, micrometre-sized hairs (acanthae). These pads are used mainly under compression; they respond to load with increasing shear resistance, and show negligible adhesion. Reflected-light microscopy in stick insects (Carausius morosus) revealed that the contact area of 'heel pads' changes with normal load on three hierarchical levels. First, loading brought larger areas of the convex pads into contact. Second, loading increased the density of acanthae in contact. Third, higher loads changed the shape of individual hair contacts gradually from circular (tip contact) to elongated (side contact). The resulting increase in real contact area can explain the load dependence of friction, indicating a constant shear stress between acanthae and substrate. As the euplantula contact area is negligible for small loads (similar to hard materials), but increases sharply with load (resembling soft materials), these pads show high friction coefficients despite little adhesion. This property appears essential for the pads' use in locomotion. Several morphological characteristics of hairy friction pads are in apparent contrast to hairy pads used for adhesion, highlighting key adaptations for both pad types. Our results are relevant for the design of fibrillar structures with high friction coefficients but small adhesion.