Abstract
Motile cilia beat in an asymmetric fashion in order to propel the surrounding fluid. When many cilia are located on a surface, their beating can synchronize such that their phases form metachronal waves. Here, we computationally study a model where each cilium is represented as a spherical particle, moving along a tilted trajectory with a position-dependent active driving force and a position-dependent internal drag coefficient. The model thus takes into account all the essential broken symmetries of the ciliary beat. We show that taking into account the near-field hydrodynamic interactions, the effective coupling between cilia even over an entire beating cycle can become nonreciprocal: The phase of a cilium is more strongly affected by an adjacent cilium on one side than by a cilium at the same distance in the opposite direction. As a result, synchronization starts from a seed at the edge of a group of cilia and propagates rapidly across the system, leading to a synchronization time that scales proportionally to the linear dimension of the system. We show that a ciliary carpet is characterized by three different velocities: the velocity of fluid transport, the phase velocity of metachronal waves, and the group velocity of order propagation. Unlike in systems with reciprocal coupling, boundary effects are not detrimental for synchronization, but rather enable the formation of the initial seed.