Wake Dynamics and Fluid Forces of Turning Maneuvers in Sunfish

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Abstract
While experimental analyses of steady rectilinear locomotion in fishes are common, unsteady movement involving time-dependent variation in heading, speed and acceleration probably accounts for the greatest portion of the locomotor time budget. Turning maneuvers, in particular, are key elements of the unsteady locomotor repertoire of fishes and, by many species, are accomplished by generating asymmetrical forces with the pectoral fins. The development of such left–right asymmetries in force production is a critical and as yet unstudied aspect of aquatic locomotor dynamics. In this paper, we measure the fluid forces exerted by the left and right pectoral fins of bluegill sunfish (Lepomis macrochirus) during turning using digital particle image velocimetry (DPIV). DPIV allowed quantification of water velocity fields, and hence momentum, in the wake of the pectoral fins as sunfish executed turns; forces exerted during turning were compared with those generated by the immediately preceding fin beats during steady swimming. Sunfish generate the forces required for turning by modulating two variables: wake momentum and pectoral fin stroke timing. Fins on opposite sides of the fish play functionally distinct roles during turning maneuvers. The fin nearer the stimulus inducing the turn (i.e. the strong side fin) generates a laterally oriented vortex ring with a strong central jet whose associated lateral force is four times greater than that produced during steady swimming. Little posterior (thrust) force is generated by the strong-side fin, and this fin therefore acts to rotate the body away from the source of the stimulus. The contralateral (weak-side) fin generates a posteriorly oriented vortex ring with a thrust force nine times that produced by the fin during steady swimming. Minimal lateral force is exerted by the weak-side fin, and this fin therefore acts primarily to translate the body linearly away from the stimulus. Turning with the paired fins is not simply steady swimming performed unilaterally. Instead, turning involves asymmetrical fin movements and fluid forces that are distinct in both direction and magnitude from those used to swim forward at constant speed. These data reflect the plasticity of the teleost pectoral fin in performing a wide range of steady and unsteady locomotor tasks.