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Climate change will increase the frequency and intensity of low-salinity (hyposalinity) events in coastal marine habitats. Sea urchins are dominant herbivores in these habitats and are generally intolerant of salinity fluctuations. Their adhesive tube feet are essential for survival, effecting secure attachment and locomotion in high wave energy habitats, yet little is known about how hyposalinity impacts their function. We exposed green sea urchins (Strongylocentrotus droebachiensis) to salinities ranging from ambient (32‰) to severe (14‰) and assessed tube feet coordination (righting response, locomotion) and adhesion [disc tenacity (force per unit area)]. Righting response, locomotion and disc tenacity decreased in response to hyposalinity. Severe reductions in coordinated tube foot activities occurred at higher salinities than those that affected adhesion. The results of this study suggest moderate hyposalinities (24–28‰) have little effect on S. droebachiensis dislodgement risk and survival post-dislodgment, while severe hyposalinity (below 24‰) likely reduces movement and prevents recovery from dislodgment.

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.

Observers ranging from Aristotle to young children have long marvelled at the ability of geckos to cling to walls and ceilings. Detailed studies have revealed that geckos are ‘sticky’ without the use of glue or suction devices. Instead, a gecko's stickiness derives from van der Waals interactions between proteinaceous hairs called setae and substrate. Here, we present surprising evidence that although geckos do not use glue, a residue is transferred on surfaces as they walk—geckos leave footprints. Using matrix-free nano-assisted laser desorption-ionization mass spectrometry, we identified the residue as phospholipids with phosphocholine head groups. Moreover, interface-sensitive sum-frequency generation spectroscopy revealed predominantly hydrophobic methyl and methylene groups and the complete absence of water at the contact interface between a gecko toe pad and the substrate. The presence of lipids has never been considered in current models of gecko adhesion. Our analysis of gecko footprints and the toe pad–substrate interface has significant consequences for models of gecko adhesion and by extension, the design of synthetic mimics.

Regenerating structures critical for survival provide excellent model systems for the study of phenotypic plasticity. These body components must regenerate their morphology and functionality quickly while subjected to different environmental stressors. Sea urchins live in high-energy environments where hydrodynamic conditions pose significant challenges. Adhesive tube feet provide secure attachment to the substratum but can be amputated by predation and hydrodynamic forces. Tube feet display functional and morphological plasticity in response to environmental conditions, but regeneration to their pre-amputation status has not been achieved under quiescent laboratory settings. In this study, we assessed the effect of turbulent water movement, periodic emersion and quiescent conditions on the regeneration process of tube foot morphology (length, disc area) and functionality (maximum disc tenacity, stem breaking force). Disc area showed significant plasticity in response to the treatments; when exposed to emersion and turbulent water movement, disc area was larger than that of tube feet regenerated in quiescent conditions. However, no treatment stimulated regeneration to pre-amputation sizes. Tube foot length was unaffected by treatments and remained shorter than non-amputated tube feet. Stem breaking force for amputated and non-amputated treatments increased in all cases when compared with pre-amputation values. Maximum tenacity (force per unit area) was similar among tube feet subjected to simulated field conditions and amputation treatments. Our results suggest a role of active plasticity of tube foot functional morphology in response to field-like conditions and demonstrate the plastic response of invertebrates to laboratory conditions.

Sea urchins native to the nearshore open coast experience periods of high, repeated wave forces that can result in dislodgement. To remain attached while clinging and locomoting across rocky substrates, sea urchins use adhesive tube feet. Purple sea urchins (Strongylocentrotus purpuratus) adhere to a variety of rock substrates (e.g. sandstone, mudstone, granite), and display morphological plasticity (skeletal morphology) to native substrate. We tested the hypothesis that their adhesive system is also plastic and varies as a function of native population and substrate. The results of our study support our hypothesis. Sea urchins from sandstone adhere less strongly to most substrates than those native to mudstone and granite rock. Sandstone produced the lowest whole animal adhesive force values across all populations, suggesting that this rock type is particularly challenging for sea urchins to adhere to. The number of adhesive tube feet that failed during experimental trials and the area used by sea urchins to attach, matches closely with whole animal adhesive force values: higher forces resulted in more tube foot failure and larger attachment area. On artificial substrates (glass and Plexiglass), differences in adhesion among populations was consistent with differences in adhesion on rock substrates except on glass, where sea urchins native to sandstone adhered more strongly to glass than any other substrate tested. To our knowledge, this study is the first to describe population-level plasticity in a biological adhesive system related to native substrate, and has significant implications for sea urchin ecology, behavior and functional morphology.

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.