Friction modulation in limbless, three-dimensional gaits and heterogeneous terrains

Motivated by a possible convergence of terrestrial limbless locomotion strategies ultimately determined by interfacial effects, we show how both 3D gait alterations and locomotory adaptations to heterogeneous terrains can be understood through the lens of local friction modulation. Via an effective-friction modeling approach, compounded by 3D simulations, the emergence and disappearance of a range of locomotory behaviors observed in nature is systematically explained in relation to inhabited environments. Our approach also simplifies the treatment of terrain heterogeneity, whereby even solid obstacles may be seen as high friction regions, which we confirm against experiments of snakes ‘diffracting’ while traversing rows of posts, similar to optical waves. We further this optic analogy by illustrating snake refraction, reflection and lens focusing. We use these insights to engineer surface friction patterns and demonstrate passive snake navigation in complex topographies. Overall, our study outlines a unified view that connects active and passive 3D mechanics with heterogeneous interfacial effects to explain a broad set of biological observations, and potentially inspire engineering design.

Generation of propulsive force via vertical undulations in snakes

Lateral undulation is the most widespread mode of terrestrial vertebrate limbless locomotion, in which posteriorly propagating horizontal waves press against environmental asperities (e.g. grass, rocks) and generate propulsive reaction forces. We hypothesized that snakes can generate propulsion using a similar mechanism of posteriorly propagating vertical waves pressing against suitably oriented environmental asperities. Using an array of horizontally oriented cylinders, one of which was equipped with force sensors, and a motion capture system, we found snakes generated substantial propulsive force and propulsive impulse with minimal contribution from lateral undulation. Additional tests showed that snakes could propel themselves via vertical undulations from a single suitable contact point, and this mechanism was replicated in a robotic model. Vertical undulations can provide snakes a valuable locomotor tool for taking advantage of vertical asperities in a variety of habitats, potentially in combination with lateral undulation, to fully exploit the 3D structure of the habitat.

Long Limbless Locomotors Over Land: The Mechanics and Biology of Elongate, Limbless Vertebrate Locomotion

Synopsis Elongate, limbless body plans are widespread in nature and frequently converged upon (with over two dozen independent convergences in Squamates alone, and many outside of Squamata). Despite their lack of legs, these animals move effectively through a wide range of microhabitats, and have a particular advantage in cluttered or confined environments. This has elicited interest from multiple disciplines in many aspects of their movements, from how and when limbless morphologies evolve to the biomechanics and control of limbless locomotion within and across taxa to its replication in elongate robots. Increasingly powerful tools and technology enable more detailed examinations of limbless locomotor biomechanics, and improved phylogenies have shed increasing light on the origins and evolution of limblessness, as well as the high frequency of convergence. Advances in actuators and control are increasing the capability of “snakebots” to solve real-world problems (e.g., search and rescue), while biological data have proven to be a potent inspiration for improvements in snakebot control. This collection of research brings together prominent researchers on the topic from around the world, including biologists, physicists, and roboticists to offer new perspective on locomotor modes, musculoskeletal mechanisms, locomotor control, and the evolution and diversity of limbless locomotion.

Burrowing dynamics of aquatic worms in soft sediments

We investigate the dynamics ofLumbriculus variegatusin water-saturated sediment beds to understand limbless locomotion in the benthic zone found at the bottom of lakes and oceans. These slender aquatic worms are observed to perform elongation–contraction and transverse undulatory strokes in both water-saturated sediments and water. Greater drag anisotropy in the sediment medium is observed to boost the burrowing speed of the worm compared to swimming in water with the same stroke using drag-assisted propulsion. We capture the observed speeds by combining the calculated forms based on resistive-force theory of undulatory motion in viscous fluids and a dynamic anchor model of peristaltic motion in the sediments. Peristalsis is found to be effective for burrowing in noncohesive sediments which fill in rapidly behind the moving body inside the sediment bed. Whereas the undulatory stroke is found to be effective in water and in shallow sediment layers where anchoring is not possible to achieve peristaltic motion. We show that such dual strokes occur as well in the earthwormEisenia fetidawhich inhabits moist sediments that are prone to flooding. Our analysis in terms of the rheology of the medium shows that the dual strokes are exploited by organisms to negotiate sediment beds that may be packed heterogeneously and can be used by active intruders to move effectively from a fluid through the loose bed surface layer which fluidizes easily to the well-consolidated bed below.

Correcting for individual quality reveals trade-offs in performance among multiple modes of limbless locomotion in snakes

Trade-offs among performance traits are often difficult to detect despite the physiological and morphological incompatibilities that underlie disparate traits being well understood. However, recent studies that have corrected for individual quality have found trade-offs in human athletes performing various performance tasks. Few studies have found trade-offs among multiple performance tasks after correcting for individual quality in non-human animals because of the difficulty in motivating many animals to perform biomechanically different tasks. We examined potential trade-offs in maximal speeds among ten locomotor conditions that involved the utilization of different locomotor modes in cornsnakes (Pantherophis guttatus). Snakes were assessed during terrestrial lateral undulation, swimming, concertina movements (small and large width) and six conditions of arboreal locomotion (combinations of three perch diameters and two inclines). We found no trade-offs among locomotor conditions when analysing uncorrected speeds or speeds corrected for body condition. However, we found several trade-offs among modes and treatments for speeds corrected for individual quality. Terrestrial lateral undulation speeds were negatively related to speeds of concertina and two of the arboreal locomotion conditions. A trade-off between speeds on large and small perch diameters on a 30° incline was also detected and probably reflects potential conflicts in traits that maximize lateral undulation and concertina.

Forward and inverse problems in the mechanics of soft filaments

Soft slender structures are ubiquitous in natural and artificial systems, in active and passive settings and across scales, from polymers and flagella, to snakes and space tethers. In this paper, we demonstrate the use of a simple and practical numerical implementation based on the Cosserat rod model to simulate the dynamics of filaments that can bend, twist, stretch and shear while interacting with complex environments via muscular activity, surface contact, friction and hydrodynamics. We validate our simulations by solving a number of forward problems involving the mechanics of passive filaments and comparing them with known analytical results, and extend them to study instabilities in stretched and twisted filaments that form solenoidal and plectonemic structures. We then study active filaments such as snakes and other slender organisms by solving inverse problems to identify optimal gaits for limbless locomotion on solid surfaces and in bulk liquids.