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      Transport and mixing by metachronal waves in nonreciprocal soft robotic pneumatic artificial cilia at low Reynolds numbers

      1 , 2 , 1
      Physics of Fluids
      AIP Publishing

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          New neurons follow the flow of cerebrospinal fluid in the adult brain.

          In the adult brain, neuroblasts born in the subventricular zone migrate from the walls of the lateral ventricles to the olfactory bulb. How do these cells orient over such a long distance and through complex territories? Here we show that neuroblast migration parallels cerebrospinal fluid (CSF) flow. Beating of ependymal cilia is required for normal CSF flow, concentration gradient formation of CSF guidance molecules, and directional migration of neuroblasts. Results suggest that polarized epithelial cells contribute important vectorial information for guidance of young, migrating neurons.
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            Overview of structure and function of mammalian cilia.

            Cilia are membrane-bounded, centriole-derived projections from the cell surface that contain a microtubule cytoskeleton, the ciliary axoneme, surrounded by a ciliary membrane. Axonemes in multiciliated cells of mammalian epithelia are 9 + 2, possess dynein arms, and are motile. In contrast, single nonmotile 9 + 0 primary cilia are found on epithelial cells, such as those of the kidney tubule, but also on nonepithelial cells, such as chondrocytes, fibroblasts, and neurons. The ciliary membranes of all cilia contain specific receptors and ion channel proteins that initiate signaling pathways controlling motility and/or linking mechanical or chemical stimuli, including sonic hedgehog and growth factors, to intracellular transduction cascades regulating differentiation, migration, and cell growth during development and in adulthood. Unique motile 9 + 0 cilia, found during development at the embryonic node, determine left-right asymmetry of the body.
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              Emergence of metachronal waves in cilia arrays.

              Propulsion by cilia is a fascinating and universal mechanism in biological organisms to generate fluid motion on the cellular level. Cilia are hair-like organelles, which are found in many different tissues and many uni- and multicellular organisms. Assembled in large fields, cilia beat neither randomly nor completely synchronously--instead they display a striking self-organization in the form of metachronal waves (MCWs). It was speculated early on that hydrodynamic interactions provide the physical mechanism for the synchronization of cilia motion. Theory and simulations of physical model systems, ranging from arrays of highly simplified actuated particles to a few cilia or cilia chains, support this hypothesis. The main questions are how the individual cilia interact with the flow field generated by their neighbors and synchronize their beats for the metachronal wave to emerge and how the properties of the metachronal wave are determined by the geometrical arrangement of the cilia, like cilia spacing and beat direction. Here, we address these issues by large-scale computer simulations of a mesoscopic model of 2D cilia arrays in a 3D fluid medium. We show that hydrodynamic interactions are indeed sufficient to explain the self-organization of MCWs and study beat patterns, stability, energy expenditure, and transport properties. We find that the MCW can increase propulsion velocity more than 3-fold and efficiency almost 10-fold--compared with cilia all beating in phase. This can be a vital advantage for ciliated organisms and may be interesting to guide biological experiments as well as the design of efficient microfluidic devices and artificial microswimmers.
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                Author and article information

                Contributors
                (View ORCID Profile)
                (View ORCID Profile)
                Journal
                Physics of Fluids
                Physics of Fluids
                AIP Publishing
                1070-6631
                1089-7666
                September 2021
                September 2021
                : 33
                : 9
                : 092009
                Affiliations
                [1 ]Zernike Institute for Advanced Materials, University of Groningen, 9747AG Groningen, The Netherlands
                [2 ]Department of Mechanical Engineering, Eindhoven University of Technology, P.O. Box 513, 5600 MB Eindhoven, The Netherlands and Institute for Complex Molecular Systems, Eindhoven University of Technology, The Netherlands
                Article
                10.1063/5.0054929
                f2233d62-ceb6-47cf-80c5-6c9668b9b81e
                © 2021
                History

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