Surprising simplicities and syntheses in limbless self-propulsion in sand.

Animals moving on and in fluids and solids move their bodies in diverse ways to generate propulsion and lift forces. In fluids, animals can wiggle, stroke, paddle or slap, whereas on hard frictional terrain, animals largely engage their appendages with the substrate to avoid slip. Granular substrates, such as desert sand, can display complex responses to animal interactions. This complexity has led to locomotor strategies that make use of fluid-like or solid-like features of this substrate, or combinations of the two. Here, we use examples from our work to demonstrate the diverse array of methods used and insights gained in the study of both surface and subsurface limbless locomotion in these habitats. Counterintuitively, these seemingly complex granular environments offer certain experimental, theoretical, robotic and computational advantages for studying terrestrial movement, with the potential for providing broad insights into morphology and locomotor control in fluids and solids, including neuromechanical control templates and morphological and behavioral evolution. In particular, granular media provide an excellent testbed for a locomotion framework called geometric mechanics, which was introduced by particle physicists and control engineers in the last century, and which allows quantitative analysis of alternative locomotor patterns and morphology to test for control templates, optimality and evolutionary alternatives. Thus, we posit that insights gained from movement in granular environments can be translated into principles that have broader applications across taxa, habitats and movement patterns, including those at microscopic scales.

Emergence of the advancing neuromechanical phase in a resistive force dominated medium.

Undulatory locomotion, a gait in which thrust is produced in the opposite direction of a traveling wave of body bending, is a common mode of propulsion used by animals in fluids, on land, and even within sand. As such, it has been an excellent system for discovery of neuromechanical principles of movement. In nearly all animals studied, the wave of muscle activation progresses faster than the wave of body bending, leading to an advancing phase of activation relative to the curvature toward the tail. This is referred to as "neuromechanical phase lags" (NPL). Several multiparameter neuromechanical models have reproduced this phenomenon, but due to model complexity, the origin of the NPL has proved difficult to identify. Here, we use perhaps the simplest model of undulatory swimming to predict the NPL accurately during sand-swimming by the sandfish lizard, with no fitting parameters. The sinusoidal wave used in sandfish locomotion, the friction-dominated and noninertial granular resistive force environment, and the simplicity of the model allow detailed analysis, and reveal the fundamental mechanism responsible for the phenomenon: the combination of synchronized torques from distant points on the body and local traveling torques. This general mechanism should help explain the NPL in organisms in other environments; we therefore propose that sand-swimming could be an excellent system with which to generate and test other neuromechanical models of movement quantitatively. Such a system can also provide guidance for the design and control of robotic undulatory locomotors in complex environments.

Controlled preparation of wet granular media reveals limits to lizard burial ability.

Many animals move within ground composed of granular media (GM); the resistive properties of such substrates can depend on water content and compaction, but little is known about how such parameters affect locomotion or the physics of drag and penetration. Using apparatus to control compaction of GM, our recent studies of movement in dry GM have revealed locomotion strategies of specialized dry-sand-swimming reptiles. However, these animals represent a small fraction of the diversity and presumed burial strategies of fossorial reptilian fauna. Here we develop a system to create states of wet GM of varying moisture content and compaction in quantities sufficient to study the burial and subsurface locomotion of the Ocellated skink (C. ocellatus), a generalist lizard. X-ray imaging revealed that in wet and dry GM the lizard slowly buried (≈30 s) propagating a wave from head to tail, while moving in a start-stop motion. During forward movement, the head oscillated, and the forelimb on the convex side of the body propelled the animal. Although body kinematics and 'slip' were similar in both substrates, the burial depth was smaller in wet GM. Penetration and drag force experiments on smooth cylinders revealed that wet GM was ≈4× more resistive than dry GM. In total, our measurements indicate that while the rheology of the dry and wet GM differ substantially, the lizard's burial motor pattern is conserved across substrates, while its burial depth is largely constrained by environmental resistance.

Locomotor benefits of being a slender and slick sand swimmer.

Squamates classified as 'subarenaceous' possess the ability to move long distances within dry sand; body elongation among sand and soil burrowers has been hypothesized to enhance subsurface performance. Using X-ray imaging, we performed the first kinematic investigation of the subsurface locomotion of the long, slender shovel-nosed snake (Chionactis occipitalis) and compared its biomechanics with those of the shorter, limbed sandfish lizard (Scincus scincus). The sandfish was previously shown to maximize swimming speed and minimize the mechanical cost of transport during burial. Our measurements revealed that the snake also swims through sand by propagating traveling waves down the body, head to tail. Unlike the sandfish, the snake nearly followed its own tracks, thus swimming in an approximate tube of self-fluidized granular media. We measured deviations from tube movement by introducing a parameter, the local slip angle, ßs, which measures the angle between the direction of movement of each segment and body orientation. The average ßs was smaller for the snake than for the sandfish; granular resistive force theory (RFT) revealed that the curvature utilized by each animal optimized its performance. The snake benefits from its slender body shape (and increased vertebral number), which allows propagation of a higher number of optimal curvature body undulations. The snake's low skin friction also increases performance. The agreement between experiment and RFT combined with the relatively simple properties of the granular 'frictional fluid' make subarenaceous swimming an attractive system to study functional morphology and bauplan evolution.

Environmental interaction influences muscle activation strategy during sand-swimming in the sandfish lizard Scincus scincus.

Animals like the sandfish lizard (Scincus scincus) that live in desert sand locomote on and within a granular medium whose resistance to intrusion is dominated by frictional forces. Recent kinematic studies revealed that the sandfish utilizes a wave of body undulation during swimming. Models predict that a particular combination of wave amplitude and wavelength yields maximum speed for a given frequency, and experiments have suggested that the sandfish targets this kinematic waveform. To investigate the neuromechanical strategy of the sandfish during walking, burial and swimming, here we use high-speed X-ray and visible light imaging with synchronized electromyogram (EMG) recordings of epaxial muscle activity. While moving on the surface, body undulation was not observed and EMG showed no muscle activation. During subsurface sand-swimming, EMG revealed an anterior-to-posterior traveling wave of muscle activation which traveled faster than the kinematic wave. Muscle activation intensity increased as the animal swam deeper into the material but was insensitive to undulation frequency. These findings were in accord with empirical force measurements, which showed that resistance force increased with depth but was independent of speed. The change in EMG intensity with depth indicates that the sandfish targets a kinematic waveform (a template) that models predict maximizes swimming speed and minimizes the mechanical cost of transport as the animal descends into granular media. The differences in the EMG pattern compared with EMG of undulatory swimmers in fluids can be attributed to the friction-dominated intrusion forces of granular media.

Mechanics of undulatory swimming in a frictional fluid.

The sandfish lizard (Scincus scincus) swims within granular media (sand) using axial body undulations to propel itself without the use of limbs. In previous work we predicted average swimming speed by developing a numerical simulation that incorporated experimentally measured biological kinematics into a multibody sandfish model. The model was coupled to an experimentally validated soft sphere discrete element method simulation of the granular medium. In this paper, we use the simulation to study the detailed mechanics of undulatory swimming in a "granular frictional fluid" and compare the predictions to our previously developed resistive force theory (RFT) which models sand-swimming using empirically determined granular drag laws. The simulation reveals that the forward speed of the center of mass (CoM) oscillates about its average speed in antiphase with head drag. The coupling between overall body motion and body deformation results in a non-trivial pattern in the magnitude of lateral displacement of the segments along the body. The actuator torque and segment power are maximal near the center of the body and decrease to zero toward the head and the tail. Approximately 30% of the net swimming power is dissipated in head drag. The power consumption is proportional to the frequency in the biologically relevant range, which confirms that frictional forces dominate during sand-swimming by the sandfish. Comparison of the segmental forces measured in simulation with the force on a laterally oscillating rod reveals that a granular hysteresis effect causes the overestimation of the body thrust forces in the RFT. Our models provide detailed testable predictions for biological locomotion in a granular environment.

Texting during stair negotiation and implications for fall risk.

BACKGROUND/AIM: Walking requires the integration of the sensory and motor systems. Cognitive distractions have been shown to interfere with negotiation of complex walking environments, especially in populations at greater risk for falls (e.g. the elderly). With the pervasiveness of mobile messaging and the recent introduction of augmented reality mobile gaming, it is increasingly important to understand how distraction associated with the simultaneous use of a mobile device impacts navigation of the complex walking environments experienced in daily life. In this study, we investigated how gait kinematics were altered when participants performed a texting task during step negotiation. METHODS: Twenty participants (13 female, 7 males) performed a series of walking trials involving a step-deck obstacle, consisting of at least 3 texting trials and 3 non-texting trials. RESULTS: When texting, participants ascended more slowly and demonstrated reduced dual-step foot toe clearance. Participants similarly descended more slowly when texting and demonstrated reduced single-step foot heel clearance as well as reduced dual-step foot fore-aft heel clearance. CONCLUSION: These data support the conclusion that texting during stair negotiation results in changes to gait kinematics that may increase the potential for gait disruptions, falls, and injury. Further research should examine the effect texting has on performing other common complex locomotor tasks, actual fall risk, and the patterns of resulting injury rate and severity when negotiating complex environments.