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Conditions for metachronal coordination in arrays of model cilia

Source Date of Publication:Sep 20,2021

Fanlong Meng, Rachel R. Bennett, Nariya Uchida, and Ramin Golestanian



Motile cilia can coordinate with each other to beat in the form of a metachronal wave, which can facilitate the self-propulsion of microorganisms such as Paramecium and can also be used for fluid transport such as mucus removal in trachea. How can we predict the collective behavior of arrays of many cilia coordinated by hydrodynamic interactions, and in particular, the properties of the emerging metachronal waves, from the single-cilium characteristics? We address this question using a bottom-up coarse-graining approach and present results that contribute to understanding how the dynamical self-organization of ciliary arrays can be controlled, which can have significant biological, medical, and engineering implications.


On surfaces with many motile cilia, beats of the individual cilia coordinate to form metachronal waves. We present a theoretical framework that connects the dynamics of an individual cilium to the collective dynamics of a ciliary carpet via systematic coarse graining. We uncover the criteria that control the selection of frequency and wave vector of stable metachronal waves of the cilia and examine how they depend on the geometric and dynamical characteristics of a single cilium, as well as the geometric properties of the array. We perform agent-based numerical simulations of arrays of cilia with hydrodynamic interactions and find quantitative agreement with the predictions of the analytical framework. Our work sheds light on the question of how the collective properties of beating cilia can be determined using information about the individual units and, as such, exemplifies a bottom-up study of a rich active matter system.