Integrating biomechanics in evolutionary studies, with examples from the amphidromous goby model system.

The functional capacities of animals are a primary factor determining survival in nature. In this context, understanding the biomechanical performance of animals can provide insight into diverse aspects of their biology, ranging from ecological distributions across habitat gradients to the evolutionary diversification of lineages. To survive and reproduce in the face of environmental pressures, animals must perform a wide range of tasks, some of which entail tradeoffs between competing demands. Moreover, the demands encountered by animals can change through ontogeny as they grow, sexually mature or migrate across environmental gradients. To understand how mechanisms that underlie functional performance contribute to survival and diversification across challenging and variable habitats, we have pursued diverse studies of the comparative biomechanics of amphidromous goby fishes across functional requirements ranging from prey capture and fast-start swimming to adhesion and waterfall climbing. The pan-tropical distribution of these fishes has provided opportunities for repeated testing of evolutionary hypotheses. By synthesizing data from the lab and field, across approaches spanning high-speed kinematics, selection trials, suction pressure recordings, mechanical property testing, muscle fiber-type measurements and physical modeling of bioinspired designs, we have clarified how multiple axes of variation in biomechanical performance associate with the ecological and evolutionary diversity of these fishes. Our studies of how these fishes meet both common and extreme functional demands add new, complementary perspectives to frameworks developed from other systems, and illustrate how integrating knowledge of the mechanical underpinnings of diverse aspects of performance can give critical insights into ecological and evolutionary questions.

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.

Wading through water: effects of water depth and speed on the drag and kinematics of walking Chilean flamingos, Phoenicopterus chilensis.

Wading behaviours, in which an animal walks while partially submerged in water, are present in a variety of taxa including amphibians, reptiles, mammals and birds. Despite the ubiquity of wading behaviours, few data are available to evaluate how animals adjust their locomotion to accommodate changes in water depth. Because drag from water might impose additional locomotor costs, wading animals might be expected to raise their feet above the water up to a certain point until such behaviours lead to awkward steps and are abandoned. To test for such mechanisms, we measured drag on models of the limbs of Chilean flamingos (Phoenicopterus chilensis) and measured their limb and body kinematics as they walked and waded through increasing depths of water in a zoo enclosure. Substantial drag was incurred by models of both open- and closed-toed feet, suggesting that flamingos could avoid some locomotor costs by stepping over water, rather than through it, during wading. Step height was highest while wading through intermediate water depths and while wading at a faster speed. Stride length increased with increasing water depth and velocity, and the limb joints generally flexed more while moving through intermediate water depths. However, movements of the head and neck were not strongly correlated with water depth or velocity. Our results show a wide range of kinematic changes that occur to allow wading birds to walk through different water depths, and have implications for better understanding the locomotor strategies employed by semi-aquatic species.

Sticking to it: testing passive pull-off forces in waterfall-climbing fishes across challenging substrates.

The pelvic sucker of Hawaiian waterfall climbing gobies allows these fishes to attach to substrates while climbing waterfalls tens to hundreds of meters tall. Climbing ability varies by species and may be further modulated by the physical characteristics of the waterfall substrate. In this study, we investigated the influence of surface wettability (hydrophobic versus hydrophilic surface charges) and substrate roughness on the passive adhesive system of four species of gobies with different climbing abilities. Overall, passive adhesive performance varied by species and substrate, with the strongest climbers showing the highest shear pull-off forces, particularly on rough surfaces. Thus, differences in passive adhesive performance may help to explain the ability of some species to migrate further upstream than others and contribute to their ability to invade new habitats.

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.