Get to the point: Claw morphology impacts frictional interactions on rough substrates.

Claws are a common anatomical feature among limbed amniotes and contribute to a variety of functions including prey capture, locomotion, and attachment. Previous studies of both avian and non-avian reptiles have found correlations between habitat use and claw morphology, suggesting that variation in claw shape permits effective functioning in different microhabitats. How, or if, claw morphology influences attachment performance, particularly in isolation from the rest of the digit, has received little attention. To examine the effects of claw shape on frictional interactions, we isolated the claws of preserved specimens of Cuban knight anoles (Anolis equestris), quantified variation in claw morphology via geometric morphometrics, and measured friction on four different substrates that varied in surface roughness. We found that multiple aspects of claw shape influence frictional interactions, but only on substrates for which asperities are large enough to permit mechanical interlocking with the claw. On such substrates, the diameter of the claw's tip is the most important predictor of friction, with narrower claw tips inducing greater frictional interactions than wider ones. We also found that claw curvature, length, and depth influence friction, but that these relationships depend on the substrate's surface roughness. Our findings suggest that although claw shape plays a critical role in the effective clinging ability of lizards, its relative importance is dependent upon the substrate. Description of mechanical function, as well as ecological function, is critical for a holistic understanding of claw shape variation.

An investigation of gecko attachment on wet and rough substrates leads to the application of surface roughness power spectral density analysis.

The roughness and wettability of surfaces exploited by free-ranging geckos can be highly variable and attachment to these substrates is context dependent (e.g., presence or absence of surface water). Although previous studies focus on the effect of these variables on attachment independently, geckos encounter a variety of conditions in their natural environment simultaneously. Here, we measured maximum shear load of geckos in air and when their toes were submerged underwater on substrates that varied in both surface roughness and wettability. Gecko attachment was greater in water than in air on smooth and rough hydrophobic substrates, and attachment to rough hydrophilic substrates did not differ when tested in air or water. Attachment varied considerably with surface roughness and characterization revealed that routine measurements of root mean square height can misrepresent the complexity of roughness, especially when measured with single instruments. We used surface roughness power spectra to characterize substrate surface roughness and examined the relationship between gecko attachment performance across the power spectra. This comparison suggests that roughness wavelengths less than 70 nm predominantly dictate gecko attachment. This study highlights the complexity of attachment in natural conditions and the need for comprehensive surface characterization when studying biological adhesive system performance.

Biomimicry Training to Promote Employee Engagement in Sustainability.

Employees play a critical role in the success of corporate sustainability initiatives, yet sustained employee engagement is a constant challenge. The psychology literature states that to intrinsically motivate employees to engage in sustainability, there must be opportunity for employees to engage in practices that are directly relevant to their job duties. Traditional ad hoc initiatives such as Earth Week events, recycling challenges and so on, are not sufficient to derive this type of intrinsic motivation. Therefore, the goal of this study was to examine the psychological impact of a biomimicry sustainable innovation training program, to intrinsically motivate R&D employees to reconnect with nature and identify whether this promotes creative thinking and employee engagement. Due to COVID-19 restrictions, the current study conducted virtual workshops with R&D employees and demonstrated that biomimicry training was intrinsically motivating to employees and was valued as a practice that could be incorporated into R&D job duties. In conclusion, this study provides an adaptable procedural template for biomimicry training with a corporate audience. The results demonstrate a strong business case for organizations to experiment with biomimicry by illustrating its potential to create positive change across several business units beyond sustainable innovation to include human resources and sustainable marketing.

Direct evidence of acid-base interactions in gecko adhesion.

While it is generally accepted that van der Waals (vdW) forces govern gecko adhesion, several studies indicate contributions from non-vdW forces and highlight the importance of understanding the adhesive contact interface. Previous work hypothesized that the surface of gecko setae is hydrophobic, with nonpolar lipid tails exposed on the surface. However, direct experimental evidence supporting this hypothesis and its implications on the adhesion mechanism is lacking. Here, we investigate the sapphire-setae contact interface using interface-sensitive spectroscopy and provide direct evidence of the involvement of acid-base interactions between polar lipid headgroups exposed on the setal surface and sapphire. During detachment, a layer of unbound lipids is left as a footprint due to cohesive failure within the lipid layer, which, in turn, reduces wear to setae during high stress sliding. The absence of this lipid layer enhances adhesion, despite a small setal-substrate contact area. Our results show that gecko adhesion is not exclusively a vdW-based, residue-free system.

Tokay geckos (Gekkonidae: Gekko gecko) preferentially use substrates that elicit maximal adhesive performance.

Gecko substrate use is likely influenced by adhesive performance, yet few studies have demonstrated this empirically. Herein, we examined the substrate use, adhesive performance and vertical clinging behaviour of Gekko gecko in captivity to investigate whether adhesive performance influences patterns of substrate use. We found that geckos were observed significantly more often on the substrate (glass) that elicited maximal adhesive performance relative to its availability within our experimental enclosures, indicating that geckos preferentially use substrates on which their adhesive performance is maximal. Our work here provides additional, yet crucial data establishing connections between adhesive performance and patterns of substrate use in captivity, suggesting the hypothesis that substrate preferences of free-ranging geckos should be correlated with adhesive performance. Clearly, further experimental and field research is necessary to test this hypothesis and identify other parameters that individually and/or collectively influence the habitat use of free-ranging geckos.

The same but different: setal arrays of anoles and geckos indicate alternative approaches to achieving similar adhesive effectiveness.

The functional morphology of squamate fibrillar adhesive systems has been extensively investigated and has indirectly and directly influenced the design of synthetic counterparts. Not surprisingly, the structure and geometry of exemplar fibrils (setae) have been the subject of the bulk of the attention in such research, although variation in setal morphology along the length of subdigital adhesive pads has been implicated to be important in the effective functioning of these systems. Adhesive setal field configuration has been described for several geckos, but that of the convergent Anolis lizards, comprised of morphologically simpler fibrils, remains largely unexplored. Here, we examine setal morphology along the proximodistal axis of the digits of Anolis equestris and compare our findings to those for a model gecko, Gekko gecko. Consistent with previous work, we found that the setae of A. equestris are generally thinner, shorter, and present at higher densities than those of G. gecko and terminate in a single spatulate tip. Contrastingly, the setae of G. gecko are hierarchically branched in structure and carry hundreds of spatulate tips. Although the splitting of contacts into multiple smaller tips is predicted to increase the adhesive performance of a fiber compared to an unbranched one, we posited that the adhesive performance of G. gecko and A. equestris would be relatively similar when the configuration of the setal fields of each was accounted for. We found that, as in geckos, setal morphology of A. equestris follows a predictable pattern along the proximodistal axis of the pad, although there are several critical differences in the configuration of the setal fields of these two groups. Most notably, the pattern of variation in setal length of A. equestris is effectively opposite to that exhibited by G. gecko. This difference in clinal variation mirrors the difference in the direction in which the setal fields of anoles and geckos are peeled from the substrate, consistent with the hypothesis that biomechanical factors are the chief determinants of these patterns of variation. Future empirical work, however, is needed to validate this. Our findings set the stage for future comparative studies investigating the functional morphology of these convergent adhesive apparatuses. Such investigations will lead to an enhanced understanding of the interactions between form, function, and environment of fibril-based biological adhesive systems.

Home-field advantage: native gecko exhibits improved exertion capacity and locomotor ability in structurally complex environments relative to its invasive counterpart.

BACKGROUND: Invasive species are of substantial concern because they may threaten ecosystem stability and biodiversity worldwide. Not surprisingly, studies examining the drivers of biological invasion have increased in number over the past few decades in an effort to curtail invasive species success by way of informing management decisions. The common house gecko, Hemidactylus frenatus, has successfully invaded the Pacific islands where it appears to thrive in and dominate non-natural habitats offering high food availability (i.e., well-lit human dwellings) compared to native geckos. Previous work demonstrated that H. frenatus can outperform the native gecko, Lepidodactylus lugubris, in terms of maximal sprint speed on relatively simple planar surfaces (e.g., building walls). Lepidodactylus lugubris and other native geckos, however, may have superior locomotor performance in three-dimensional, structurally complex habitats. RESULTS: Here we compared the locomotor behaviour and exertion capacity of the native gecko, Gehyra oceanica, and the invasive gecko, Hemidactylus frenatus, on the island of Mo'orea, French Polynesia, on fabricated structures simulating structurally complex substrates. We found that the native gecko exhibits improved locomotor performance compared to the invasive gecko on structurally complex substrates. We also completed encounter surveys to document free-ranging habitat use and behaviour of these two species. We discovered that H. frenatus were more common in natural habitats than previously observed and used similar substrates as G. oceanica, although G. oceanica appeared to use substrates with greater perch heights (i.e., trees). CONCLUSIONS: Our findings revealed that locomotor performance in complex environments may contribute to the previously observed habitat segregation between native and invasive Pacific island geckos. Furthermore, our locomotor and habitat use data are consistent with the hypothesis that G. oceanica may be resistant to invasion of H. frenatus in natural environments. Our study calls for more detailed ecophysiological and ecomorphological studies of both native and invasive Pacific gecko species.

Digital hyperextension has no influence on the active self-drying of gecko adhesive subdigital pads.

The remarkable properties of the gecko adhesive system have been intensively studied. Although many gecko-inspired synthetic adhesives have been designed and fabricated, few manage to capture the multifunctionality of the natural system. Analogous to previously documented self-cleaning, recent work demonstrated that gecko toe pads dry when geckos take steps on dry substrates (i.e., self-drying). Whether digital hyperextension (DH), the distal to proximal peeling of gecko toe pads, is involved in the self-drying process, had not been determined. Here, the effect of DH on self-drying was isolated by preventing DH from occurring during normal walking locomotion of Gekko gecko after toe pads were wetted. Our initial analysis revealed low statistical power, so we increased our sample size to determine the robustness of our result. We found that neither DH nor the DH-substrate interaction had a significant effect on the maximum shear adhesive force after self-drying. These results suggest that DH is not necessary for self-drying to occur. Interestingly, however, we discovered that shear adhesion is higher on a surface tending hydrophobic compared to a hydrophilic surface, demonstrating that gecko adhesion is sensitive to substrate wettability during the subdigital pad drying process. Furthermore, we also observed frequent damage to the adhesive system during shear adhesion testing post-drying, indicating that water may compromise the structural integrity of the adhesive structures. Our results not only have behavioral and ecological implications for free-ranging geckos but also have the potential to influence the design and fabrication of gecko-inspired synthetic adhesives that can regain adhesion after fouling with water.

A Physical Model Approach to Gecko Adhesion Opportunity and Constraint: How Rough Could It Be?

It has been nearly 20 years since Autumn and colleagues established the central role of van der Waals intermolecular forces in how geckos stick. Much has been discovered about the structure and function of fibrillar adhesives in geckos and other taxa, and substantial success has been achieved in translating natural models into bioinspired synthetic adhesives. Nevertheless, synthetics still cannot match the multidimensional performance observed in the natural gecko system that is simultaneously robust to dirt and water, resilient over thousands of cycles, and purportedly competent on surfaces that are rough at drastically different length scales. Apparent insensitivity of adhesion to variability in roughness is particularly interesting from both a theoretical and applied perspective. Progress on understanding the extent to which and the basis of how the gecko adhesive system is robust to variation in roughness is impeded by the complexity of quantifying roughness of natural surfaces and a dearth of data on free-ranging gecko substrate use. Here we review the main challenges in characterizing rough surfaces as they relate to collecting relevant estimates of variation in gecko adhesive performance across different substrates in their natural habitats. In response to these challenges, we propose a practical protocol (borrowing from thermal biophysical ecological methods) that will enable researchers to design detailed studies of structure-function relationships of the gecko fibrillar system. Employing such an approach will help provide specific hypotheses about how adhesive pad structure translates into a capacity for robust gecko adhesion across large variation in substrate roughness. Preliminary data we present on this approach suggest its promise in advancing the study of how geckos deal with roughness variation. We argue and outline how such data can help advance development of design parameters to improve bioinspired adhesives based on the gecko fibrillar system.

Going Out on a Limb: How Investigation of the Anoline Adhesive System Can Enhance Our Understanding of Fibrillar Adhesion.

The remarkable ability of geckos to adhere to a wide-variety of surfaces has served as an inspiration for hundreds of studies spanning the disciplines of biomechanics, functional morphology, ecology, evolution, materials science, chemistry, and physics. The multifunctional properties (e.g., self-cleaning, controlled releasability, reversibility) and adhesive performance of the gekkotan adhesive system have motivated researchers to design and fabricate gecko-inspired synthetic adhesives of various materials and properties. However, many challenges remain in our attempts to replicate the properties and performance of this complex, hierarchical fibrillar adhesive system, stemming from fundamental, but unanswered, questions about how fibrillar adhesion operates. Such questions involve the role of fibril morphology in adhesive performance and how the gekkotan adhesive apparatus is utilized in nature. Similar fibrillar adhesive systems have, however, evolved independently in two other lineages of lizards (anoles and skinks) and potentially provide alternate avenues for addressing these fundamental questions. Anoles are the most promising group because they have been the subject of intensive ecological and evolutionary study for several decades, are highly speciose, and indeed are advocated as squamate model organisms. Surprisingly, however, comparatively little is known about the morphology, performance, and properties of their convergently-evolved adhesive arrays. Although many researchers consider the performance of the adhesive system of Anolis lizards to be less accomplished than its gekkotan counterpart, we argue here that Anolis lizards are prime candidates for exploring the fundamentals of fibrillar adhesion. Studying the less complex morphology of the anoline adhesive system has the potential to enhance our understanding of fibril morphology and its relationship to the multifunctional performance of fibrillar adhesive systems. Furthermore, the abundance of existing data on the ecology and evolution of anoles provides an excellent framework for testing hypotheses about the influence of habitat microstructure on the performance, behavior, and evolution of lizards with subdigital adhesive pads.

The Ecomechanics of Gecko Adhesion: Natural Surface Topography, Evolution, and Biomimetics.

The study of gecko adhesion is necessarily interdisciplinary due to the hierarchical nature of the adhesive system and the complexity of interactions between the animals and their habitats. In nature, geckos move on a wide range of surfaces including soft sand dunes, trees, and rocks, but much of the research over the past two decades has focused on their adhesive performance on artificial surfaces. Exploring the complex interactions between geckos and their natural habitats will reveal aspects of the adhesive system that can be applied to biomimetic research, such as the factors that facilitate movement on dirty and rough surfaces with varying microtopography. Additionally, contrasting suites of constraints and topographies are found on rocks and plants, likely driving differences in locomotion and morphology. Our overarching goals are to bring to light several aspects of ecology that are important for gecko-habitat interactions, and to propose a framework for how they can inspire material scientists and functional ecologists. We also present new data on surface roughness and topography of a variety of surfaces, and adhesive performance of Phelsuma geckos on surfaces of varying roughness. We address the following key questions: (1) why and how should ecology be incorporated into the study of gecko adhesion? (2) What topographical features of rocks and plants likely drive adhesive performance? (3) How can ecological studies inform material science research? Recent advances in surface replication techniques that eliminate confounding factors among surface types facilitate the ability to address some of these questions. We pinpoint gaps in our understanding and identify key initiatives that should be adopted as we move forward. Most importantly, fine details of locomotor microhabitat use of both diurnal and nocturnal geckos are needed.

Influence of substrate modulus on gecko adhesion.

The gecko adhesion system fascinates biologists and materials scientists alike for its strong, reversible, glue-free, dry adhesion. Understanding the adhesion system's performance on various surfaces can give clues as to gecko behaviour, as well as towards designing synthetic adhesive mimics. Geckos encounter a variety of surfaces in their natural habitats; tropical geckos, such as Gekko gecko, encounter hard, rough tree trunks as well as soft, flexible leaves. While gecko adhesion on hard surfaces has been extensively studied, little work has been done on soft surfaces. Here, we investigate for the first time the influence of macroscale and nanoscale substrate modulus on whole animal adhesion on two different substrates (cellulose acetate and polydimethylsiloxane) in air and find that across 5 orders of magnitude in macroscale modulus, there is no change in adhesion. On the nanoscale, however, gecko adhesion is shown to depend on substrate modulus. This suggests that low surface-layer modulus may inhibit the gecko adhesion system, independent of other influencing factors such as macroscale composite modulus and surface energy. Understanding the limits of gecko adhesion is vital for clarifying adhesive mechanisms and in the design of synthetic adhesives for soft substrates (including for biomedical applications and wearable electronics).

The effect of temperature and humidity on adhesion of a gecko-inspired adhesive: implications for the natural system.

The adhesive system of geckos has inspired hundreds of synthetic adhesives. While this system has been used relentlessly as a source of inspiration, less work has been done in reverse, where synthetics are used to test questions and hypotheses about the natural system. Here we take such an approach. We tested shear adhesion of a mushroom-tipped synthetic gecko adhesive under conditions that produced perplexing results in the natural adhesive system. Synthetic samples were tested at two temperatures (12 °C and 32 °C) and four different humidity levels (30%, 55%, 70%, and 80% RH). Surprisingly, adhesive performance of the synthetic samples matched that of living geckos, suggesting that uncontrolled parameters in the natural system, such as surface chemistry and material changes, may not be as influential in whole-animal performance as previously thought. There was one difference, however, when comparing natural and synthetic adhesive performance. At 12 °C and 80% RH, adhesion of the synthetic structures was lower than expected based on the natural system's performance. Our approach highlights a unique opportunity for both biologists and material scientists, where new questions and hypotheses can be fueled by joint comparisons of the natural and synthetic systems, ultimately improving knowledge of both.

Superhydrophobicity of the gecko toe pad: biological optimization versus laboratory maximization.

While many gecko-inspired hierarchically structured surfaces perform as well as or better than the natural adhesive system, these designs often fail to function across a variety of contexts. For example, the gecko can adhere to rough, wet and dirty surfaces; however, most synthetic mimics cannot maintain function when faced with a similar situation. The solution to this problem lies in a more thorough investigation of the natural system. Here, we review the adhesive system of the gecko toe pad, as well as the far less-well-studied anti-adhesive system that results from the chemistry and structure of the toe pad (superhydrophobicity). This paradoxical relationship serves as motivation to study functional optimization at the system level. As an example, we experimentally investigate the role of surface lipids in adhesion and anti-adhesion, and find a clear performance trade-off related to shear adhesion in air on a hydrophilic surface. This represents the first direct investigation of the role of surface lipids in gecko adhesion and anti-adhesion, and supports the argument that a system-level approach is necessary to elucidate optimization in biological systems. Without such an approach, bioinspired designs will be limited in functionality and context, especially compared to the natural systems they mimic.This article is part of the themed issue 'Bioinspired hierarchically structured surfaces for green science'.

Sticking to the story: outstanding challenges in gecko-inspired adhesives.

The natural clinging ability of geckos has inspired hundreds of studies seeking design principles that could be applied to creating synthetic adhesives with the same performance capabilities as the gecko: adhesives that use no glue, are self-cleaning and reusable, and are insensitive to a wide range of surface chemistries and roughness. Important progress has been made, and the basic mechanics of how 'hairy' adhesives work have been faithfully reproduced, advancing theory in surface science and portending diverse practical applications. However, after 15 years, no synthetic mimic can yet perform as well as a gecko and simultaneously meet of all the criteria listed above. Moreover, processes for the production of inexpensive and scalable products are still not clearly in view. Here, we discuss our perspective on some of the gaps in understanding that still remain; these gaps in our knowledge should stimulate us to turn to deeper study of the way in which free-ranging geckos stick to the variety of surfaces found in their natural environments and to a more complete analysis of the materials composing the gecko toe pads.

Gecko Adhesion on Wet and Dry Patterned Substrates.

Perhaps one of the most astounding characteristics of the gecko adhesive system is its versatility. Geckos can locomote across complex substrates in a variety of conditions with apparent ease. In contrast, many of our synthetic pressure sensitive adhesives fail on substrates that are dirty, wet or rough. Although many studies have investigated the effect of environmental challenges on performance, the interaction of multiple, potentially compromising variables is studied less often. Here we focus on substrate structure and surface water, both of which are highly relevant to the biological system and to synthetic design. To do this we utilized a highly controlled, patterned substrate (Sharklet®, by Sharklet® Technologies Inc.). This allowed us to test independently and jointly the effects of reduced surface area substrates, with a defined pattern, on adhesion in both air and water. Our results show that adhesion is not significantly impaired in air, whereas surface area and pattern significantly affect adhesion in water. These findings highlight the need to study multiple parameters that are relevant to the gecko adhesive system to further improve our understanding of the biological system and to design better, more versatile synthetics.

Robust self-cleaning and micromanipulation capabilities of gecko spatulae and their bio-mimics.

Geckos have the extraordinary ability to prevent their sticky feet from fouling while running on dusty walls and ceilings. Understanding gecko adhesion and self-cleaning mechanisms is essential for elucidating animal behaviours and rationally designing gecko-inspired devices. Here we report a unique self-cleaning mechanism possessed by the nano-pads of gecko spatulae. The difference between the velocity-dependent particle-wall adhesion and the velocity-independent spatula-particle dynamic response leads to a robust self-cleaning capability, allowing geckos to efficiently dislodge dirt during their locomotion. Emulating this natural design, we fabricate artificial spatulae and micromanipulators that show similar effects, and that provide a new way to manipulate micro-objects. By simply tuning the pull-off velocity, our gecko-inspired micromanipulators, made of synthetic microfibers with graphene-decorated micro-pads, can easily pick up, transport, and drop-off microparticles for precise assembling. This work should open the door to the development of novel self-cleaning adhesives, smart surfaces, microelectromechanical systems, biomedical devices, and more.

Run don't walk: locomotor performance of geckos on wet substrates.

The gecko adhesive system has been under particular scrutiny for over a decade, as the field has recently attracted attention for its application to bio-inspired design. However, little is known about how the adhesive system behaves in ecologically relevant conditions. Geckos inhabit a variety of environments, many of which are characterized by high temperature, humidity and rain. The van der Waals-based gecko adhesive system should be particularly challenged by wet substrates because water can disrupt the intimate contact necessary for adhesion. While a few previous studies have focused on the clinging ability of geckos on wet substrates, we tested a dynamic performance characteristic, sprint velocity. To better understand how substrate wettability and running orientation affect locomotor performance of multiple species on wet substrates, we measured average sprint velocity of five species of gecko on substrates that were either hydrophilic or intermediately wetting and oriented either vertically or horizontally. Surprisingly, we found no indication that wet substrates impact average sprint velocity over 1 m, and rather, in some species, sprint velocity was increased on wet substrates rather than reduced. When investigating physical characteristics and behavior that may be associated with running on wet substrates, such as total number of stops, slips and wet toes at the completion of a race, we found that there may be habitat-related differences between some species. Our results show that in general, unlike clinging and walking, geckos running along wet substrates suffer no significant loss in locomotor performance over short distances.

Adhesive interactions of geckos with wet and dry fluoropolymer substrates.

Fluorinated substrates like Teflon® (poly(tetrafluoroethylene); PTFE) are well known for their role in creating non-stick surfaces. We showed previously that even geckos, which can stick to most surfaces under a wide variety of conditions, slip on PTFE. Surprisingly, however, geckos can stick reasonably well to PTFE if it is wet. In an effort to explain this effect, we have turned our attention to the role of substrate surface energy and roughness when shear adhesion occurs in media other than air. In this study, we removed the roughness component inherent to commercially available PTFE and tested geckos on relatively smooth wet and dry fluoropolymer substrates. We found that roughness had very little effect on shear adhesion in air or in water and that the level of fluorination was most important for shear adhesion, particularly in air. Surface energy calculations of the two fluorinated substrates and one control substrate using the Tabor-Winterton approximation and the Young-Dupré equation were used to determine the interfacial energy of the substrates. Using these interfacial energies we estimated the ratio of wet and dry normal adhesion for geckos clinging to the three substrates. Consistent with the results for rough PTFE, our predictions show a qualitative trend in shear adhesion based on fluorination, and the quantitative experimental differences highlight the unusually low shear adhesion of geckos on dry smooth fluorinated substrates, which is not captured by surface energy calculations. Our work has implications for bioinspired design of synthetics that can preferentially stick in water but not in air.