Organisms use hair-like cilia that beat in a metachronal fashion to actively transport fluid and suspended particles. Metachronal motion emerges due to a phase difference between beating cycles of neighboring cilia and appears as traveling waves propagating along ciliary carpet. In this work, we demonstrate biomimetic artificial cilia capable of metachronal motion. The cilia are micromachined magnetic thin filaments attached at one end to a substrate and actuated by a uniform rotating magnetic field. We show that the difference in magnetic cilium length controls the phase of the beating motion. We use this property to induce metachronal waves within a ciliary array and explore the effect of operation parameters on the wave motion. The metachronal motion in our artificial system is shown to depend on the magnetic and elastic properties of the filaments, unlike natural cilia, where metachronal motion arises due to fluid coupling. Our approach enables an easy integration of metachronal magnetic cilia in lab-on-a-chip devices for enhanced fluid and particle manipulations.
Motile cilia are slender, hair-like cellular appendages that spontaneously oscillate under the action of internal molecular motors and are typically found in dense arrays. These active filaments coordinate their beating to generate metachronal waves that drive long-range fluid transport and locomotion. Until now, our understanding of their collective behavior largely comes from the study of minimal models that coarse-grain the relevant biophysics and the hydrodynamics of slender structures. Here we build on a detailed biophysical model to elucidate the emergence of metachronal waves on millime…
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 wavevector of stable metchacronal waves of the cilia and examine how they depend on the geometric and dynamical characteristics of single cilia, as well as the geometric properties of the array. We perform agent-based numerical simulations of arrays of cilia with…
Surfing on metachronal waves: ciliary transport by inertial coasting
University of Pennsylvania（宾夕法尼亚大学）；The Zeeman Institute for Systems Biology and Infectious Disease Epidemiology Research（泽曼系统生物学与传染病流行病学研究所）；University of Warsaw（华沙大学）；University of Warwick（华威大学）
Microorganisms develop coordinated beating patterns on surfaces lined with cilia known as metachronal waves. For a chain of cilia attached to a flat ciliate, it has been shown that hydrodynamic interactions alone can lead the system to synchronize. However, several microorganisms possess a curve shaped ciliate body and so to understand the effect of this geometry on the formation of metachronal waves, we evaluate the hydrodynamic interactions of cilia near a large spherical body. Using a minimal model, we show that for a chain of cilia around the sphere, the natural periodicity in the geometry…
When tiny soft ferromagnetic particles are placed along a liquid interface and exposed to a vertical magnetic field, the balance between capillary attraction and magnetic repulsion leads to self-organization into well-defined patterns. Here, we demonstrate experimentally that precessing magnetic fields induce metachronal waves on the periphery of these assemblies, similar to the ones observed in ciliates and some arthropods. The outermost layer of particles behaves like an array of cilia or legs whose sequential movement causes a net and controllable locomotion. This bioinspired many-particle …
By taking into account the hydrodynamic interactions in a one dimensional array of model cilia attached to a no-slip cylinderical surface, we investigate their synchronized motion. We show, how does the emergence of metachronal waves depend on the initial state of the system and investigate the conditions under which, the formation of symplectic and antiplectic waves are possible.
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 …
Coordinated cilia are used throughout the natural world for micronscale fluid transport. They are often modelled with regular filament arrays on fixed, planar surfaces. Here, we simulate hundreds of interacting active filaments on spherical surfaces, where defects in the cilia displacement field must be present. We see synchronised beating towards or about two defects for spheres held fixed. Defects alter filament beating which causes the sphere to move once released. This motion feeds back to the filaments resulting in a whirling state with metachronal behaviour along the equator.
Eukaryotic cilia and flagella are chemo-mechanical oscillators capable of generating long-range coordinated motions known as metachronal waves. Pair synchronization is a fundamental requirement for these collective dynamics, but it is generally not sufficient for collective phase-locking, chiefly due to the effect of long-range interactions. Here we explore experimentally and numerically a minimal model for a ciliated surface; hydrodynamically coupled oscillators rotating above a no-slip plane. Increasing their distance from the wall profoundly effects the global dynamics, due to variations in…
One of the vital functions of naturally occurring cilia is fluid transport. Biological cilia use spatially asymmetric strokes to generate a net fluid flow that can be utilized for feeding, swimming, and other functions. Biomimetic synthetic cilia with similar asymmetric beating can be useful for fluid manipulations in lab-on-chip devices. In this paper, we demonstrate the microfluidic pumping by magnetically actuated synthetic cilia arranged in multi-row arrays. We use a microchannel loop to visualize flow created by the ciliary array and to examine pumping for a range of cilia and microchanne…
One of the major challenges in modern robotics is controlling micromanipulation by active and adaptive materials. In the respiratory system, such actuation enables pathogen clearance by means of motile cilia. While various types of artificial cilia have been engineered recently, they often involve complex manufacturing protocols and focus on transporting liquids only. Here, we create soft magnetic carpets via an easy self-assembly route based on the Rosensweig instability. These carpets can transport liquids but also solid objects that are larger and heavier than the artificial cilia, using a …
Small - but finite - fluid inertia can be leveraged to generate steady flows out of liquid vibrations around an immersed interface. In engineering, external high-frequency drivers (10$^2$-10$^5$ Hz) allow this inertial rectification phenomenon, known as viscous streaming, to be employed in micron-scale devices for precise flow control, particle manipulation and spatially controlled chemistry. However, beyond artificial settings, streaming may also be accessed by larger-scale biological systems pertaining to lower frequencies. Then, millimeter-size bacteria or larvae that oscillate cilia and ap…
