The hydrodynamics of locomotion at intermediate Reynolds numbers:undulatory swimming in ascidian larvae (Botrylloidessp.)

Understanding how the shape and motion of an aquatic animal affects the performance of swimming requires knowledge of the fluid forces that generate thrust and drag. These forces are poorly understood for the large diversity of animals that swim at Reynolds numbers (Re) between 10(0) and 10(2). We experimentally tested quasi-steady and unsteady blade-element models of the hydrodynamics of undulatory swimming in the larvae of the ascidian Botrylloides sp. by comparing the forces predicted by these models with measured forces generated by tethered larvae and by comparing the swimming speeds predicted with measurements of the speed of freely swimming larvae. Although both models predicted mean forces that were statistically indistinguishable from measurements, the quasi-steady model predicted the timing of force production and mean swimming speed more accurately than the unsteady model. This suggests that unsteady force (i.e. the acceleration reaction) does not play a role in the dynamics of steady undulatory swimming at Re approximately 10(2). We explored the relative contribution of viscous and inertial force to the generation of thrust and drag at 10(0)<Re<10(2) by running a series of mathematical simulations with the quasi-steady model. These simulations predicted that thrust and drag are dominated by viscous force (i.e. skin friction) at Re approximately 10(0) and that inertial force (i.e. form force) generates a greater proportion of thrust and drag at higher Re than at lower Re. However, thrust was predicted to be generated primarily by inertial force, while drag was predicted to be generated more by viscous than inertial force at Re approximately 10(2). Unlike swimming at high (>10(2)) and low (<10(0)) Re, the fluid forces that generate thrust cannot be assumed to be the same as those that generate drag at intermediate Re.