Comparative Studies Reveal Principles of Movement on and within Granular Media

Terrestrial locomotion can take place on complex substrates such as leaf litter, debris, and soil that flow or solidify in response to stress. While principles of movement in air and water are revealed through study of the hydrodynamic equations of fluid motion, discovery of principles of movement in complex terrestrial environments is less advanced in part because describing the physics of limb and body interaction with such environments remains challenging. We report progress our group has made in discovering principles of movement of organisms and models of organisms (robots) on and within granular materials (GM) like sand. We review current understanding of localized intrusion in GM relevant to foot and body interactions. We discuss the limb-ground interactions of a desert lizard, a hatchling sea turtle, and various robots and reveal that control of granular solidification can generate effective movement. We describe the sensitivity of movement on GM to gait parameters and discuss how changes in material state can strongly affect locomotor performance. We examine subsurface movement, common in desert animals like the sandfish lizard. High speed x-ray imaging resolves subsurface kinematics, while electromyography (EMG) allows muscle activation patterns to be studied. Our resistive force theory, numerical, and robotic models of sand-swimming reveal that subsurface swimming occurs in a “frictional fluid” whose properties differ from Newtonian fluids.