Tempo and mode of performance evolution across multiple independent origins of adhesive toe pads in lizards.

Understanding macroevolutionary dynamics of trait evolution is an important endeavor in evolutionary biology. Ecological opportunity can liberate a trait as it diversifies through trait space, while genetic and selective constraints can limit diversification. While many studies have examined the dynamics of morphological traits, diverse morphological traits may yield the same or similar performance and as performance is often more proximately the target of selection, examining only morphology may give an incomplete understanding of evolutionary dynamics.

Modeling observed animal performance using the Weibull distribution.

To understand how organisms adapt, researchers must link performance and microhabitat. However, measuring performance, especially maximum performance, can sometimes be difficult. Here, we describe an improvement over previous techniques that only consider the largest observed values as maxima.

Stick or grip? Co-evolution of adhesive toepads and claws in Anolis lizards.

Exploring the relationship between phenotype and performance in an ecological and evolutionary context is crucial to understanding the adaptive nature of phenotypic traits. Despite their ubiquity in vertebrates, few studies have examined the functional and ecological significance of claw morphologies. Here we examine the adhesive toepad and claw system of Anolis lizards.

Geckoprinting: assembly of microelectronic devices on unconventional surfaces by transfer printing with isolated gecko setal arrays.

Developing electronics in unconventional forms provides opportunities to expand the use of electronics in diverse applications including bio-integrated or implanted electronics. One of the key challenges lies in integrating semiconductor microdevices onto unconventional substrates without glue, high pressure or temperature that may cause damage to microdevices, substrates or interfaces. This paper describes a solution based on natural gecko setal arrays that switch adhesion mechanically on and off, enabling pick and place manipulation of thin microscale semiconductor materials onto diverse surfaces including plants and insects whose surfaces are usually rough and irregular.

Experimental evidence for friction-enhancing integumentary modifications of chameleons and associated functional and evolutionary implications.

The striking morphological convergence of hair-like integumentary derivatives of lizards and arthropods (spiders and insects) demonstrates the importance of such features for enhancing purchase on the locomotor substrate. These pilose structures are responsible for the unique tractive abilities of these groups of animals, enabling them to move with seeming ease on overhanging and inverted surfaces, and to traverse inclined smooth substrates. Three groups of lizards are well known for bearing adhesion-promoting setae on their digits: geckos, anoles and skinks.

Dry self-cleaning properties of hard and soft fibrillar structures.

Recently, gecko-inspired synthetic adhesives (GSAs) have been made using a variety of fabrication techniques and materials, with one made from a hard polymer having been reported to recover its shear adhesion after fouling by normal use, or "dry self-clean", a feature useful for applications in wall crawling robots, reusable adhesives, microfabrication and solar panel cleaning. This paper investigates the impact of two design parameters on the dry self-cleaning capability of GSAs by experimentally testing two GSAs after fouling with small (1 μm), medium (3-10 μm), and large (40-50 μm) particles. We found that a GSA made from a hard thermoplastic with nanoscopic fibers was able to recover 96-115% of its shear adhesion after fouling with small and large but not medium particles, while a GSA made from a soft polymer and microscopic fibers recovered 40-55% on medium and large particles, with SEM imaging revealing particles embedding within the polymer.

Macroscale adhesion of gecko setae reflects nanoscale differences in subsurface composition.

Surface energies are commonly used to determine the adhesion forces between materials. However, the component of surface energy derived from long-range forces, such as van der Waals forces, depends on the material's structure below the outermost atomic layers. Previous theoretical results and indirect experimental evidence suggest that the van der Waals energies of subsurface layers will influence interfacial adhesion forces.

Design and fabrication of gecko-inspired adhesives.

Recently, there has been significant interest in developing dry adhesives mimicking the gecko adhesive system, which offers several advantages compared to conventional pressure-sensitive adhesives. Specifically, gecko adhesive pads have anisotropic adhesion properties; the adhesive pads (spatulae) stick strongly when sheared in one direction but are non-adherent when sheared in the opposite direction. This anisotropy property is attributed to the complex topography of the array of fine tilted and curved columnar structures (setae) that bear the spatulae.

Changes in materials properties explain the effects of humidity on gecko adhesion.

Geckos owe their remarkable stickiness to millions of dry setae on their toes, and the mechanism of adhesion in gecko setae has been the topic of scientific scrutiny for over two centuries. Previously, we demonstrated that van der Waals forces are sufficient for strong adhesion and friction in gecko setae, and that water-based capillary adhesion is not required. However, recent studies demonstrated that adhesion increases with relative humidity (RH) and proposed that surface hydration and capillary water bridge formation is important or even necessary.

Gecko adhesion pad: a smart surface?

Recently, it has been shown that humidity can increase the adhesion of the spatula pads that form the outermost (adhesive) surface of the tokay gecko feet by 50% relative to the main adhesion mechanism (i.e. van der Waals adhesive forces), although the mechanism by which the enhancement is realized is still not well understood.

Role of tilted adhesion fibrils (setae) in the adhesion and locomotion of gecko-like systems.

Geckos are super climbers: they can readily and rapidly stick to almost any surface, whether hydrophilic or hydrophobic, rough or smooth, in dry or wet conditions, and detach with equal rapidity within tens of milliseconds. In this paper, we discuss the rapid switching between the strong adhesion/friction (attached) state and zero adhesion/friction (detached) state, and present a finite element analysis of gecko setae in terms of their adhesion and friction forces. The analysis shows why the asymmetric, naturally curved setae with a directional tilt play a crucial role in the gecko's articulation mechanism, consistent with recent experimental studies of gecko setal arrays.

Microscopic modeling of the dynamics of frictional adhesion in the gecko attachment system.

We present a simple microscopic model describing the unique friction behavior of gecko setal arrays as they are dragged on smooth surfaces. Unlike other solids of high elastic modulus that do not stick under van der Waals forces alone, the gecko setal arrays do not require a compressive force to display a drag resistance but rather develop a tensile normal force when they are dragged (J. Experim.

Rate-dependent frictional adhesion in natural and synthetic gecko setae.

Geckos owe their remarkable stickiness to millions of dry, hard setae on their toes. In this study, we discovered that gecko setae stick more strongly the faster they slide, and do not wear out after 30,000 cycles. This is surprising because friction between dry, hard, macroscopic materials typically decreases at the onset of sliding, and as velocity increases, friction continues to decrease because of a reduction in the number of interfacial contacts, due in part to wear.

A microfabricated wedge-shaped adhesive array displaying gecko-like dynamic adhesion, directionality and long lifetime.

Gecko adhesion has become a paradigmatic example of bio-inspired engineering, yet among the many gecko-like synthetic adhesives (GSAs), truly gecko-like performance remains elusive. Many GSAs have previously demonstrated one or two features of the gecko adhesive. We present a new wedge-shaped GSA that exhibits several gecko-like properties simultaneously: directional features; zero force at detachment; high ratio of detachment force to preload force; non-adhesive default state; and the ability to maintain performance while sliding, even after thousands of cycles.

Adhesion and friction force coupling of gecko setal arrays: implications for structured adhesive surfaces.

The extraordinary climbing ability of geckos is partially attributed to the fine structure of their toe pads, which contain arrays consisting of thousands of micrometer-sized stalks (setae) that are in turn terminated by millions of fingerlike pads (spatulae) having nanoscale dimensions. Using a surface forces apparatus (SFA), we have investigated the dynamic sliding characteristics of setal arrays subjected to various loading, unloading, and shearing conditions at different angles. Setal arrays were glued onto silica substrates and, once installed into the SFA, brought toward a polymeric substrate surface and then sheared.

Convergent setal morphology in sand-covering spiders suggests a design principle for particle capture.

Sicarius and Homalonychus are unrelated, desert-dwelling spiders that independently evolved the ability to cover themselves in fine sand particles, making them cryptic against their background. Observations that particles associate with these spiders' setae inspired us to investigate the role of setal microstructure in particle capture and retention. Here we report that Sicarius and Homalonychus convergently evolved numerous high aspect ratio, flexible fibres that we call 'hairlettes' protruding from the setal shaft.

Frictional and elastic energy in gecko adhesive detachment.

Geckos use millions of adhesive setae on their toes to climb vertical surfaces at speeds of over 1 m s(-1). Climbing presents a significant challenge for an adhesive since it requires both strong attachment and easy, rapid removal. Conventional pressure-sensitive adhesives are either strong and difficult to remove (e.

Analysis of the locomotor activity of a nocturnal desert lizard (Reptilia: Gekkonidae: Teratoscincus scincus) under varying moonlight.

1. This project seeks to identify determinants of the variation observed in the foraging behavior of predatory animals, especially in moonlight, using a lizard as a model. 2.

Ultrahydrophobicity indicates a non-adhesive default state in gecko setae.

Geckos may represent the world's most demanding adhesives application. The adhesive setae on the toes of climbing geckos must adhere strongly yet avoid fouling or attachment at inappropriate times. We tested the hypothesis that gecko setae are non-adhesive in their unloaded default state by comparing the water droplet contact angle (theta) of isolated setal arrays to the smooth surface of eye spectacle scales of tokay geckos (Gekko gecko).

Increased capacity for sustained locomotion at low temperature in parthenogenetic geckos of hybrid origin.

The evolution of parthenogenesis is typically associated with hybridization and polyploidy. These correlates of parthenogenesis may have important physiological consequences that need be taken into account in understanding the relative merits of sexual and parthenogenetic reproduction. We compared the thermal sensitivity of aerobically sustained locomotion in hybrid/triploid parthenogenetic races of the gecko Heteronotia binoei and their diploid sexual progenitors.

Integrating historical and mechanistic biology enhances the study of adaptation.

Adding a causal, mechanistic dimension to the study of character evolution will increase the strength of inferences regarding the evolutionary history of characters and their adaptive consequences. This approach has the advantage of illuminating mechanism and testing evolutionary hypotheses rigorously. We consider the advantages of combining mechanistic and historical biology in the study of behavior, physiology, and development.

Mechanisms of adhesion in geckos.

The extraordinary adhesive capabilities of geckos have challenged explanation for millennia, since Aristotle first recorded his observations. We have discovered many of the secrets of gecko adhesion, yet the millions of dry, adhesive setae on the toes of geckos continue to generate puzzling new questions and valuable answers. Each epidermally-derived, keratinous seta ends in hundreds of 200 nm spatular tips, permitting intimate contact with rough and smooth surfaces alike.