Locomotor forces on a swimming fish: three-dimensional vortex wake dynamics quantified using digital particle image velocimetry.

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Quantifying the locomotor forces experienced by swimming fishes represents a significant challenge because direct measurements of force applied to the aquatic medium are not feasible. However, using the technique of digital particle image velocimetry (DPIV), it is possible to quantify the effect of fish fins on water movement and hence to estimate momentum transfer from the animal to the fluid. We used DPIV to visualize water flow in the wake of the pectoral fins of bluegill sunfish (Lepomis macrochirus) swimming at speeds of 0.5-1.5 L s(-)(1), where L is total body length. Velocity fields quantified in three perpendicular planes in the wake of the fins allowed three-dimensional reconstruction of downstream vortex structures. At low swimming speed (0.5 L s(-)(1)), vorticity is shed by each fin during the downstroke and stroke reversal to generate discrete, roughly symmetrical, vortex rings of near-uniform circulation with a central jet of high-velocity flow. At and above the maximum sustainable labriform swimming speed of 1.0 L s(-)(1), additional vorticity appears on the upstroke, indicating the production of linked pairs of rings by each fin. Fluid velocity measured in the vicinity of the fin indicates that substantial spanwise flow during the downstroke may occur as vortex rings are formed. The forces exerted by the fins on the water in three dimensions were calculated from vortex ring orientation and momentum. Mean wake-derived thrust (11.1 mN) and lift (3.2 mN) forces produced by both fins per stride at 0.5 L s(-)(1) were found to match closely empirically determined counter-forces of body drag and weight. Medially directed reaction forces were unexpectedly large, averaging 125 % of the thrust force for each fin. Such large inward forces and a deep body that isolates left- and right-side vortex rings are predicted to aid maneuverability. The observed force balance indicates that DPIV can be used to measure accurately large-scale vorticity in the wake of swimming fishes and is therefore a valuable means of studying unsteady flows produced by animals moving through fluids.