Escaping Flatland: three-dimensional kinematics and hydrodynamics of median fins in fishes.

Fish swimming has often been simplified into the motions of a two-dimensional slice through the horizontal midline, as though fishes live in a flat world devoid of a third dimension. While fish bodies do undulate primarily horizontally, this motion has important three-dimensional components, and fish fins can move in a complex three-dimensional manner. Recent results suggest that an understanding of the three-dimensional body shape and fin motions is vital for explaining the mechanics of swimming, and that two-dimensional representations of fish locomotion are misleading. In this study, we first examine axial swimming from the two-dimensional viewpoint, detailing the limitations of this view. Then we present data on the kinematics and hydrodynamics of the dorsal fin, the anal fin and the caudal fin during steady swimming and maneuvering in brook trout, Salvelinus fontinalis, bluegill sunfish, Lepomis macrochirus, and yellow perch, Perca flavescens. These fishes actively move the dorsal and anal fins during swimming, resulting in curvature along both anterio-posterior and dorso-ventral axes. The momentum imparted to the fluid by these fins comprises a substantial portion of total swimming force, adding to thrust and contributing to roll stability. While swimming, the caudal fin also actively curves dorso-ventrally, producing vortices separately from both its upper and lower lobes. This functional separation of the lobes may allow additional control of three-dimensional orientation, but probably reduces swimming efficiency. In contrast, fish may boost the caudal fin's efficiency by taking advantage of the flow from the dorsal and anal fins as it interacts with the flow around the caudal fin itself. During maneuvering, fish readily use their fins outside of the normal planes of motion. For example, the dorsal fin can flick laterally, orienting its surface perpendicular to the body, to help in turning and braking. These data demonstrate that, while fish do move primarily in the horizontal plane, neither their bodies nor their motions can accurately be simplified in a two-dimensional representation. To begin to appreciate the functional consequences of the diversity of fish body shapes and locomotor strategies, one must escape Flatland to examine all three dimensions.