Mechanisms underlying rhythmic locomotion: body–fluid interaction in undulatory swimming

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Swimming of fish and other animals results from interactions of rhythmic body movements with the surrounding fluid. This paper develops a model for the body-fluid interaction in undulatory swimming of leeches, where the body is represented by a chain of rigid links and the hydrodynamic force model is based on resistive and reactive force theories. The drag and added-mass coefficients for the fluid force model were determined from experimental data of kinematic variables during intact swimming, measured through video recording and image processing. Parameter optimizations to minimize errors in simulated model behaviors revealed that the resistive force is dominant, and a simple static function of relative velocity captures the essence of hydrodynamic forces acting on the body. The model thus developed, together with the experimental kinematic data, allows us to investigate temporal and spatial (along the body) distributions of muscle actuation, body curvature, hydrodynamic thrust and drag, muscle power supply and energy dissipation into the fluid. We have found that: (1) thrust is generated continuously along the body with increasing magnitude toward the tail, (2) drag is nearly constant along the body, (3) muscle actuation waves travel two or three times faster than the body curvature waves and (4) energy for swimming is supplied primarily by the mid-body muscles, transmitted through the body in the form of elastic energy, and dissipated into the water near the tail.