SUMMARY Conventional pressure sensitive adhesives (PSAs) are fabricated from soft viscoelastic materials that satisfy Dahlquist's criterion for tack with a Young's modulus ( E ) of 100 kPa or less at room temperature and 1 Hz. In contrast, the adhesive on the toes of geckos is made of β-keratin, a stiff material with E at least four orders of magnitude greater than the upper limit of Dahlquist's criterion. Therefore, one would not expect aβ-keratin structure to function as a PSA by deforming readily to make intimate molecular contact with a variety of surface profiles. However, since the gecko adhesive is a microstructure in the form of an array of millions of high aspect ratio shafts (setae), the effective elastic modulus( E eff ) is much lower than E of bulkβ-keratin. In the first test of the E eff of a gecko setal adhesive, we measured the forces resulting from deformation of isolated arrays of tokay gecko ( Gekko gecko ) setae during vertical compression, and during tangential compression at angles of +45° and-45°. We tested the hypothesis that E eff of gecko setae falls within Dahlquist's criterion for tack, and evaluated the validity of a model of setae as cantilever beams. Highly linear forces of deformation under all compression conditions support the cantilever model. E eff of setal arrays during vertical and +45°compression (along the natural path of drag of the setae) were 83±4.0 kPa and 86±4.4 kPa (means ± s.e.m.), respectively. Consistent with the predictions of the cantilever model, setae became significantly stiffer when compressed against the natural path of drag: E eff during -45° compression was 110±4.7 kPa. Unlike synthetic PSAs, setal arrays act as Hookean elastic solids; setal arrays function as a bed of springs with a directional stiffness, assisting alignment of the adhesive spatular tips with the contact surface during shear loading.
SUMMARY Directional arrays of branched microscopic setae constitute a dry adhesive on the toes of pad-bearing geckos, nature's supreme climbers. Geckos are easily and rapidly able to detach their toes as they climb. There are two known mechanisms of detachment: (1) on the microscale, the seta detaches when the shaft reaches a critical angle with the substrate, and (2) on the macroscale, geckos hyperextend their toes, apparently peeling like tape. This raises the question of how geckos prevent detachment while inverted on the ceiling, where body weight should cause toes to peel and setal angles to increase. Geckos use opposing feet and toes while inverted, possibly to maintain shear forces that prevent detachment of setae or peeling of toes. If detachment occurs by macroscale peeling of toes, the peel angle should monotonically decrease with applied force. In contrast, if adhesive force is limited by microscale detachment of setae at a critical angle, the toe detachment angle should be independent of applied force. We tested the hypothesis that adhesion is increased by shear force in isolated setal arrays and live gecko toes. We also tested the corollary hypotheses that (1) adhesion in toes and arrays is limited as on the microscale by a critical angle, or (2)on the macroscale by adhesive strength as predicted for adhesive tapes. We found that adhesion depended directly on shear force, and was independent of detachment angle. Therefore we reject the hypothesis that gecko toes peel like tape. The linear relation between adhesion and shear force is consistent with a critical angle of release in live gecko toes and isolated setal arrays, and also with our prior observations of single setae. We introduced a new model,frictional adhesion, for gecko pad attachment and compared it to existing models of adhesive contacts. In an analysis of clinging stability of a gecko on an inclined plane each adhesive model predicted a different force control strategy. The frictional adhesion model provides an explanation for the very low detachment forces observed in climbing geckos that does not depend on toe peeling.