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Outside laboratory conditions and human-made structures, animals rarely encounter flat surfaces. Instead, natural substrates are uneven surfaces with height variation that ranges from the microscopic scale to the macroscopic scale. For walking animals (which we define as encompassing any form of legged movement across the ground, such as walking, running, galloping, etc.), such substrate ‘roughness’ influences locomotion in a multitude of ways across scales, from roughness that influences how each toe or foot contacts the ground, to larger obstacles that animals must move over or navigate around. Historically, the unpredictability and variability of natural environments has limited the ability to collect data on animal walking biomechanics. However, recent technical advances, such as more sensitive and portable cameras, biologgers, laboratory tools to fabricate rough terrain, as well as the ability to efficiently store and analyze large variable datasets, have expanded the opportunity to study how animals move under naturalistic conditions. As more researchers endeavor to assess walking over rough terrain, we lack a consistent approach to quantifying roughness and contextualizing these findings. This Review summarizes existing literature that examines non-human animals walking on rough terrain and presents a metric for characterizing the relative substrate roughness compared with animal size. This framework can be applied across terrain and body scales, facilitating direct comparisons of walking over rough surfaces in animals ranging in size from ants to elephants. 

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.

 

Synopsis Arboreal ants must navigate variably sized and inclined linear structures across a range of substrate roughness when foraging tens of meters above the ground. To achieve this, arboreal ants use specialized adhesive pads and claws to maintain effective attachment to canopy substrates. Here, we explored the effect of substrate structure, including small and large-scale substrate roughness, substrate diameter, and substrate orientation (inclination), on adhesion and running speed of workers of one common, intermediately-sized, arboreal ant species. Normal (orthogonal) and shear (parallel) adhesive performance varied on sandpaper and natural leaf substrates, particularly at small size scales, but running speed on these substrates remained relatively constant. Running speed also varied minimally when running up and down inclined substrates, except when the substrate was positioned completely vertical. On vertical surfaces, ants ran significantly faster down than up. Ant running speed was slower on relatively narrow substrates. The results of this study show that variation in the physical properties of tree surfaces differentially affects arboreal ant adhesive and locomotor performance. Specifically, locomotor performance was much more robust to surface roughness than was adhesive performance. The results provide a basis for understanding how performance correlates of functional morphology contribute to determining local ant distributions and foraging decisions in the tropical rainforest canopy. 

Abstract Geckos are remarkable in their ability to reversibly adhere to smooth vertical, and even inverted surfaces. However, unraveling the precise mechanisms by which geckos do this has been a long process, involving various approaches over the last two centuries. Our understanding of the principles by which gecko adhesion operates has advanced rapidly over the past 20 years and, with this knowledge, material scientists have attempted to mimic the system to create artificial adhesives. From a biological perspective, recent studies have examined the diversity in morphology, performance, and real-world use of the adhesive apparatus. However, the lack of multidisciplinarity is likely a key roadblock to gaining new insights. Our goals in this paper are to 1) present a historical review of gecko adhesion research, 2) discuss the mechanisms and morphology of the adhesive apparatus, 3) discuss the origin and performance of the system in real-world contexts, 4) discuss advancement in bio-inspired design, and 5) present grand challenges in gecko adhesion research. To continue to improve our understanding, and to more effectively employ the principles of gecko adhesion for human applications, greater intensity and scope of interdisciplinary research are necessary.