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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., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

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., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

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., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

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., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

The surface temperature of tree branches in the tropical rainforest canopy can reach up to 55°C. Ants and other small cursorial organisms must maintain adequate attachment in this extreme microenvironment to forage effectively and avoid falling. Ant adhesion depends on liquid secretions that should become less viscous at high temperatures, causing ants to slip. However, tropical arboreal ants have high thermal tolerance and actively forage on hot canopy surfaces, suggesting that these ants can maintain adhesion on hot substrates. We measured tarsal pad shear adhesion of 580 workers (representing 11 species and four subfamilies) of tropical arboreal ants at temperatures spanning the range observed in the field (23-55°C). Adhesive performance among species showed three general trends: (1) a linear decrease with increasing temperature, (2) a non-linear relationship with peak adhesive performance at ca. 30-40°C, and (3) no relationship with temperature. The mechanism responsible for these large interspecific differences remains to be determined, but likely reflects variation in the composition of the secreted adhesive fluid. Understanding such differences will reveal the diverse ways that ants cope with highly variable, and often unpredictable, thermal conditions in the forest canopy., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

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., (© 2016. Published by The Company of Biologists Ltd.)

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., (© 2015. Published by The Company of Biologists Ltd.)

Despite profound interest in the mechanics and performance of the gecko adhesive system, relatively few studies have focused on performance under conditions that are ecologically relevant to the natural habitats of geckos. Because geckos are likely to encounter surfaces that are wet, we used shear force adhesion measurements to examine the effect of surface water and toe pad wetting on the whole-animal performance of a tropical-dwelling gecko (Gekko gecko). To test the effect of surface wetting, we measured the shear adhesive force of geckos on three substrate conditions: dry glass, glass misted with water droplets and glass fully submerged in water. We also investigated the effect of wetting on the adhesive toe pad by soaking the toe pads prior to testing. Finally, we tested for repeatability of the adhesive system in each wetting condition by measuring shear adhesion after each step a gecko made under treatment conditions. Wetted toe pads had significantly lower shear adhesive force in all treatments (0.86 ± 0.09 N) than the control (17.96 ± 3.42 N), as did full immersion in water (0.44 ± 0.03 N). Treatments with droplets of water distributed across the surface were more variable and did not differ from treatments where the surface was dry (4.72 ± 1.59 N misted glass; 9.76 ± 2.81 N dry glass), except after the gecko took multiple steps. These findings suggest that surface water and the wetting of a gecko's adhesive toe pads may have significant consequences for the ecology and behavior of geckos living in tropical environments.