Hydromechanical properties of metachronal swimming in polychaetes

Free-swimming polychaetes are common in marine habitats and exhibit a unique form of swimming whereby a metachronal wave occurs simultaneously with a bending body wave. This body wave is unusual among swimming animals because it travels in the same direction as the animal's swimming direction. However, we currently lack a mechanistic understanding of this unusual form of locomotion. In this study we use a combination of high-speed, high-resolution video and particle image velocimetry (PIV) to quantify kinematics and fluid dynamics for three species of swimming polychaetes, spanning two orders of magnitude in size. We find that in all species, flows generated by metachronal waves of parapodia dominate while typical flows associated with body bending is absent. However, the parapodia are less flexible than propulsive structures in other metachronal swimmers. This creates a localized, but substantial upstream flow during the recovery stroke. Using body bending, the recovery stroke can occur mostly beneath the bulk flow from the power strokes, resulting in minimal inference while the subsequent power stroke can benefit from the pressure field generated during recovery. These results may have implications for future vehicle designs that incorporate metachronal locomotion.