Simulation of snakes using vertical body bending to traverse terrain with large height variation

Snake moves across various terrains by bending its elongated body. Recent studies discovered that snakes can use vertical bending to traverse terrain of large height variation, such as horizontally oriented cylinders, a wedge (Jurestovsky, Usher, Astley, 2021, J. Exp. Biol.), and uneven terrain (Fu & Li, 2020, Roy. Soc. Open Sci.; Fu, Astley, Li, J. Exp. Biol., in review). Here, to understand how vertical bending generates propulsion, we developed a dynamic simulation of a snake traversing a wedge (height ≈ 0.05 body length, slope = 27°) and a half cylindrical obstacle (height ≈ 0.1 body length). By propagating down the body an internal torque profile with a maximum around the obstacle, the simulated snake moved forward as observed in the animal. Remarkably, even when frictional drag is low (snake-terrain kinetic friction coefficient of 0.20), the body must push against the wedge with a pressure 5 times that from body weight to generate sufficient forward propulsion to move forward. This indicated that snakes are highly capable of bending vertically to push against the environment to generate propulsion. Testing different controllers revealed that contact force feedback further helps generate and maintain propulsion effectively under unknown terrain perturbations.