Tree frogs are remarkable for their capacity to cling to smooth surfaces using large toe pads. The adhesive skin of tree frog toe pads is characterized by peg-studded hexagonal cells separated by deep channels into which mucus glands open. The pads are completely wetted with watery mucus, which led previous authors to suggest that attachment is solely due to capillary and viscous forces generated by the fluid-filled joint between the pad and the substrate. Here, we present evidence from single-toe force measurements, laser tweezer microrheometry of pad mucus and interference reflection microscopy of the contact zone inLitoria caerulea, that tree frog attachment forces are significantly enhanced by close contacts and boundary friction between the pad epidermis and the substrate, facilitated by the highly regular pad microstructure.

SUMMARY: Many arthropods and vertebrates possess tarsal adhesive pads densely covered with setae. The striking morphological convergence of `hairy' pads in lizards, spiders and several insect orders demonstrates the advantage of this design for substrate adhesion. Early functional explanations of hairy adhesive organs focused on the performance on rough substrates, where flexible setae can make more intimate contact. Recent theoretical and experimental work shows that the hairy design can also help to achieve self-cleaning properties,controllable detachment and increased adhesion. Several arguments have been proposed to explain why adhesive forces are maximised. First, the `Force scaling' hypothesis states that when adhesive forces scale linearly with the dimensions of the contact, adhesion is increased by dividing the contact zone into many microscopic subunits. Second, the `Fracture mechanics' argument implies that adhesion is maximised when the size of adhesive contacts is smaller than the critical crack length. Third, the `Work of adhesion' model suggests that adhesion increases due to the bending and stretching of setae and associated energy losses during detachment.Several morphological traits of hairy adhesive pads can be explained by the need to maximise the work of adhesion, while avoiding the sticking of setae to each other (self-matting). Firstly, if setae are oblique and convex toward the foot tip as typical of most hairy pads, arrays should achieve greater adhesion. Secondly, a branched seta morphology not only confers the advantage that setae can adapt to roughness at different length scales but also prevents self-matting and increases the work of adhesion.It is predicted from the `Work of adhesion' model that adhesion of pads with unbranched setae cannot be increased by subdividing the contact zone into ever finer subcontacts, because this would increasingly cause self-matting. However, contact splitting can increase adhesion if setae are branched. The greater density of setae in large animals has been interpreted by `Force scaling'. However, the existing data can be explained by the effect of seta branching and by a fundamental difference between `wet' and `dry' adhesive systems. As insects employ adhesive fluids, they can cope with small-scale surface roughness even with relatively blunt seta tips, whereas the dry systems of lizards and spiders require extremely fine endings.

Adhesive pads on the legs of animals can be classified as either `smooth' or `hairy' (fibrillar). It has been proposed that the hairy design conveys superior and controllable adhesion. However, no study has yet compared the basic performance of both systems. As such, we measured single-pad friction and adhesion forces in sample hairy (Gastrophysa viridula) and smooth (Carausius morosus) pads and simultaneously recorded contact area. Adhesion and friction forces per unit pad area were very similar in smooth and hairy systems. Insect pads of both types adhere via a thin film of liquid secretion. As found previously for the smooth system, forces in the fibrillar system strongly decreased with larger amounts of fluid secretion present, suggesting that the fluid mainly serves to maximize contact on rough substrates. One essential prerequisite for the control of surface attachment during locomotion is the direction-dependence of adhesive pads. We compared the mechanisms of direction-dependence in smooth and hairy systems by performing proximal and distal slides. Both types of pad exhibited a large drop in friction when moved away from the body, although this effect was more extreme for the hairy system. Direction-dependence is explained in both smooth and fibrillar systems by the instability of the tarsal chain, causing the whole pad to peel off. In the fibrillar pads, anisotropy additionally arises from the direction-dependence of individual setae.

Attachment devices are essential adaptations for climbing animals and valuable models for synthetic adhesives. A major unresolved question for both natural and bioinspired attachment systems is how attachment performance depends on size. Here, we discuss how contact geometry and mode of detachment influence the scaling of attachment forces for claws and adhesive pads, and how allometric data on biological systems can yield insights into their mechanism of attachment. Larger animals are expected to attach less well to surfaces, due to their smaller surface-to-volume ratio, and because it becomes increasingly difficult to distribute load uniformly across large contact areas. In order to compensate for this decrease of weight-specific adhesion, large animals could evolve overproportionally large pads, or adaptations that increase attachment efficiency (adhesion or friction per unit contact area). Available data suggest that attachment pad area scales close to isometry within clades, but pad efficiency in some animals increases with size so that attachment performance is approximately size-independent. The mechanisms underlying this biologically important variation in pad efficiency are still unclear. We suggest that switching between stress concentration (easy detachment) and uniform load distribution (strong attachment) via shear forces is one of the key mechanisms enabling the dynamic control of adhesion during locomotion.

Adhesive organs on the legs of arthropods and vertebrates are strongly direction dependent, making contact only when pulled towards the body but detaching when pushed away from it. Here we show that the two types of attachment pads found in cockroaches (Nauphoeta cinerea), tarsal euplantulae and pretarsal arolium, serve fundamentally different functions. Video recordings of vertical climbing revealed that euplantulae are almost exclusively engaged with the substrate when legs are pushing, whereas arolia make contact when pulling. Thus, upward-climbing cockroaches used front leg arolia and hind leg euplantulae, whereas hind leg arolia and front leg euplantulae were engaged during downward climbing. Single-leg friction force measurements showed that the arolium and euplantulae have an opposite direction dependence. Euplantulae achieved maximum friction when pushed distally, whereas arolium forces were maximal during proximal pulls. This direction dependence was not explained by the variation of shear stress but by different contact areas during pushing or pulling. The changes in contact area result from the arrangement of the flexible tarsal chain, tending to detach the arolium when pushing and to peel off euplantulae when in tension. Our results suggest that the euplantulae in cockroaches are notadhesiveorgans but ‘frictionpads’, mainly providing the necessary traction during locomotion.