Spontaneous phase coordination and fluid pumping in model ciliary carpets
arXiv:2208.10811 · doi:10.1073/pnas.2214413119
Abstract
Ciliated tissues such as in the mammalian lungs, brains, and reproductive tracts, are specialized to pump fluid. They generate flows by the collective activity of hundreds of thousands of individual cilia that beat in a striking metachronal wave pattern. Despite progress in analyzing cilia coordination, a general theory that links coordination and fluid pumping in the limit of large arrays of cilia remains lacking. Here, we conduct in-silico experiments with thousands of hydrodynamically-interacting cilia, and we develop a continuum theory in the limit of infinitely-many independently beating cilia by combining tools from active matter and classical Stokes flow. We find, in both simulations and theory, that isotropic and synchronized ciliary states are unstable. Traveling waves emerge regardless of initial conditions, but the characteristics of the wave and net flows depend on cilia and tissue properties. That is, metachronal phase coordination is a stable global attractor in large ciliary carpets, even under finite perturbations to cilia and tissue properties. These results support the notion that functional specificity of ciliated tissues is interlaced with the tissue architecture and cilia beat kinematics and open up the prospect of establishing structure-to-function maps from cilium-level beat to tissue-level coordination and fluid pumping.
arXiv comment11 pages, 5 figures, 1 SI document (pdf) and 2 video included, revision for publication 09/19/2022
References in corpus (16)
- Finding the ciliary beating pattern with optimal efficiency
- Coordinated Beating of Algal Flagella is Mediated by Basal Coupling
- Cell body rocking is a dominant mechanism for flagellar synchronization in a swimming alga
- Multiscale polar theory of microtubule and motor-protein assemblies
- Efficiency optimization and symmetry-breaking in a model of ciliary locomotion
- Mixing and transport by ciliary carpets: a numerical study
- Microfluidic pumping using artificial magnetic cilia
- A multiscale biophysical model gives quantized metachronal waves in a lattice of cilia
- Bistability in the synchronization of actuated microfilaments
- In-phase and anti-phase flagellar synchronization by basal coupling
- Multisynchrony in active microfilaments
- Transitions in synchronization states of model cilia through basal-connection coupling
- Fluid mechanics of mosaic ciliated tissues
- Synchronization in cilia carpets and the Kuramoto model with local coupling: breakup of global synchronization in the presence of noise
- The Phase Synchronized State of Oriented Active Fluids
- Self-organized flows in phase-synchronizing active fluids