Functional morphology and hydrodynamics of backward swimming in bluegill sunfish, Lepomis macrochirus

Abstract

Most teleost fishes, like the bluegill sunfish Lepomis macrochirus, have multiple flexible fins that are used as modifiable control surfaces. This helps to make fish highly maneuverable, permitting behaviors like reversing direction of motion and swimming backwards without having to rotate body position. To answer the question of how fish swim backwards we used high-speed videography and electromyography to determine the kinematics and muscle activity necessary to produce reverse-direction propulsion in four bluegill sunfish. We found that, in contrast to slow forward swimming, low-speed backward swimming is a multi-fin behavior, utilizing the pectoral, dorsal, anal, and caudal fins. The pectoral fins alternate beats, each fin broadly flaring on the outstroke and feathered on the instroke. The dorsal fin and dorsal portion of the caudal fin move out of phase as do the anal fin and ventral portion of the caudal fin. Electromyography of muscles in the pectoral, dorsal, anal, and caudal fins demonstrated bilateral activation when these fins changed direction, suggesting that fins are stiffened at this time. In addition to backward propulsion by the pectoral fins, particle image velocimetry revealed that the dorsal and anal fins are capable of producing reverse momentum jets to propel the fish backward. Because teleost fishes are statically unstable, locomotion at slow speeds requires precise fin control to adequately balance torques produced about the center of mass. Therefore, the kinematics of backward swimming may be the result of compensation for rolling, pitching, and yawning instability. We suggest that asymmetric pectoral fin activity with feathering during adduction balances rolling instability. The ventral to dorsal undulatory wave on the caudal fin controls pitch instability and yaw instability encountered from pectoral-driven backward locomotion. Thrust generation from the dorsal and anal fins decreases the destabilizing effect of the long moment arm of the tail in backward swimming. Thus, backward locomotion at slow speed is not simply the reverse of slow forward swimming.

Keywords: Backward swimming; Fin; Hydrodynamics; Locomotion; Muscle activity.