How zebrafish turn: analysis of pressure force dynamics and mechanical work

Abstract

Whereas many fish swim steadily, zebrafish regularly exhibit unsteady burst-and-coast swimming, which is characterized by repeated sequences of turns followed by gliding periods. Such a behavior offers the opportunity to investigate the hypothesis that negative mechanical work occurs in posterior regions of the body during early phases of the turn near the time of maximal body curvature. Here we use a modified Particle Image Velocimetry (PIV) technique to obtain high-resolution flow fields around the zebrafish body during turns. Using detailed swimming kinematics coupled with body surface pressure computations, we estimated fluid-structure interaction forces and the pattern of forces and torques along the body during turning. We then calculated the mechanical work done by each body segment. We used estimated patterns of positive and negative work along the body to evaluate the hypothesis (based on fish midline kinematics) that the posterior body region would experience predominantly negative work. Between 10 and 20% of the total mechanical work was done by the fluid on the body (negative work), and negative work was concentrated in the anterior and middle areas of the body, not along the caudal region. Energetic costs of turning were calculated by considering the sum of positive and negative work and were compared to previous metabolic estimates of turning energetics in fishes. The analytical workflow presented here provides a rigorous way to quantify hydrodynamic mechanisms of fish locomotion and facilitates the understanding how body kinematics generates locomotor forces in freely-swimming fishes.