Fluid circulation driven by collectively organized metachronal waves in swimming T. aceti nematodes

Recent experiments have shown that the nematode T. aceti can assemble into collectively undulating groups at the edge of fluid drops. This coordinated state consists of metachronal waves and drives fluid circulation inside the drop. We find that the circulation velocity is about 2 mm/s and nearly half the speed of the metachronal wave. We develop a quasi-two-dimensional hydrodynamics model using the Stokes flow approximation. The periodic motion of the nematodes constitute our moving boundary condition that drives the flow. Our model suggests that large-amplitude excursions of the nematode tails produce the fluid circulation. We discuss the constraints on containers that would enhance fluid motion, which could be used in the future design of on demand flow generating systems.