Hydrodynamic Synchronization In Strong Confinement

Cellular appendages conferring motility, such as flagella and cilia, are known to synchronise their periodic beats. The origin of synchronization is a combination of long-range hydrodynamic interactions with physical mechanisms allowing the phases of these biological oscillators to evolve. Two of such mechanisms have been identified by previous work, the elastic compliance of the periodic orbit or oscillations driven by phase-dependent biological forcing, both of which can lead generically to stable phase locking. In order to help uncover the physical mechanism for hydrodynamic synchronization most essential overall in biology, we theoretically investigate in this paper the effect of strong confinement on the effectiveness of hydrodynamic synchronization. Following past work, we use minimal models of cilia where appendages are modeled as rigid spheres forced to move along circular trajectories near a rigid surface. Strong confinement is modeled by adding a second nearby surface, parallel to the first one, where the distance between the surfaces is much smaller than the typical distance between the cilia, which results in a qualitative change in the nature of hydrodynamic interactions. We calculate separately the impact of hydrodynamic confinement on the synchronization dynamics of the elastic compliance and the force modulation mechanisms and compare our results to the usual case with a single surface. Applying our results to the biologically relevant situation of nodal cilia, we show that force modulation is a mechanism that leads to phase-locked states under strong confinement that are very similar to those without confinement as a difference with the elastic compliance mechanism. Our results point therefore to the robustness of force modulation for synchronization, an important feature for biological dynamics that therefore suggests it could be the most essential physical mechanism overall in arrays of nodal cilia. We further examine the distinct biologically relevant situation of primary cilia and show in that case that the difference in robustness of the mechanisms is not as pronounced but still favors the force modulation.