Metachronal wave and hydrodynamic interaction for deterministic switching rowers

We employ a model system, called rowers, as a generic physical framework to define the problem of the coordinated motion of cilia (the metachronal wave) as a far from equilibrium process. Rowers are active (two-state) oscillators interacting solely through forces of hydrodynamic origin. In this work, we consider the case of fully deterministic dynamics, find analytical solutions of the equation of motion in the long wavelength (continuum) limit, and investigate numerically the short wavelength limit. We prove the existence of metachronal waves below a characteristic wavelength. Such waves are unstable and become stable only if the sign of the coupling is reversed. We also find that with normal hydrodynamic interaction the metachronal pattern has the form of stable trains of traveling wave packets sustained by the onset of anti-coordinated beating of consecutive rowers.Comment: 11 pages, 7 figure

Discrete cilia modelling with singularity distributions

We discuss in detail techniques for modelling flows due to finite and infinite arrays of beating cilia. An efficient technique, based on concepts from previous ‘singularity models’ is described, that is accurate in both near and far-fields. Cilia are modelled as curved slender ellipsoidal bodies by distributing Stokeslet and potential source dipole singularities along their centrelines, leading to an integral equation that can be solved using a simple and efficient discretisation. The computed velocity on the cilium surface is found to compare favourably with the boundary condition. We then present results for two topics of current interest in biology. 1) We present the first theoretical results showing the mechanism by which rotating embryonic nodal cilia produce a leftward flow by a ‘posterior tilt,’ and track particle motion in an array of three simulated nodal cilia. We find that, contrary to recent suggestions, there is no continuous layer of negative fluid transport close to the ciliated boundary. The mean leftward particle transport is found to be just over 1 μm/s, within experimentally measured ranges. We also discuss the accuracy of models that represent the action of cilia by steady rotlet arrays, in particular, confirming the importance of image systems in the boundary in establishing the far-field fluid transport. Future modelling may lead to understanding of the mechanisms by which morphogen gradients or mechanosensing cilia convert a directional flow to asymmetric gene expression. 2) We develop a more complex and detailed model of flow patterns in the periciliary layer of the airway surface liquid. Our results confirm that shear flow of the mucous layer drives a significant volume of periciliary liquid in the direction of mucus transport even during the recovery stroke of the cilia. Finally, we discuss the advantages and disadvantages of the singularity technique and outline future theoretical and experimental developments required to apply this technique to various other biological problems, particularly in the reproductive system

Sperm Chemotaxis: A Primer

Sperm become attracted by chemical substances that are released from the outer coating of the egg, a process called chemotaxis. In this paper the cellular pathway and the motor response during chemotaxis of sperm from sea urchin and starfish are briefly outlined