Experimental hydrodynamics of fish locomotion: functional insights from wake visualization.

Despite enormous progress during the last twenty years in understanding the mechanistic basis of aquatic animal propulsion-a task involving the construction of a substantial data base on patterns of fin and body kinematics and locomotor muscle function-there remains a key area in which biologists have little information: the relationship between propulsor activity and water movement in the wake. How is internal muscular force translated into external force exerted on the water? What is the pattern of fluid force production by different fish fins (e.g., pectoral, caudal, dorsal) and how does swimming force vary with speed and among species? These types of questions have received considerable attention in analyses of terrestrial locomotion where force output by limbs can be measured directly with force plates. But how can forces exerted by animals moving through fluid be measured? The advent of digital particle image velocimetry (DPIV) has provided an experimental hydrodynamic approach for quantifying the locomotor forces of freely moving animals in fluids, and has resulted in significant new insights into the mechanisms of fish propulsion. In this paper we present ten "lessons learned" from the application of DPIV to problems of fish locomotion over the last five years. (1) Three-dimensional DPIV analysis is critical for reconstructing wake geometry. (2) DPIV analysis reveals the orientation of locomotor reaction forces. (3) DPIV analysis allows calculation of the magnitude of locomotor forces. (4) Swimming speed can have a major impact on wake structure. (5) DPIV can reveal interspecific differences in vortex wake morphology. (6) DPIV analysis can provide new insights into the limits to locomotor performance. (7) DPIV demonstrates the functional versatility of fish fins. (8) DPIV reveals hydrodynamic force partitioning among fins. (9) DPIV shows that wake interaction among fins may enhance thrust production. (10) Experimental hydrodynamic analysis can provide insight into the functional significance of evolutionary variation in fin design.