Cellular appendages conferring motility, such as flagella and cilia, are known to synchronise their periodic beats. The origin of synchronisation is a combination of long-range hydrodynamic interactions with physical mechanisms allowing the phases of these biological oscillators to evolve. Two of such mechanisms have been identified by previous work, the elastic compliance of the periodic orbit or oscillations driven by phase-dependent biological forcing, both of which can lead generically to stable phase locking. In order to help uncover the physical mechanism for hydrodynamic synchronisation most essential overall in biology, we theoretically investigate in this paper the effect of strong confinement on the effectiveness of hydrodynamic synchronisation. Following past work, we use minimal models of cilia where appendages are modelled as rigid spheres forced to move along circular trajectories near a rigid surface. Strong confinement is modelled by adding a second nearby surface, parallel to the first one, where the distance between the surfaces is much smaller than the typical distance between the cilia, which results in a qualitative change in the nature of hydrodynamic interactions. We calculate separately the impact of hydrodynamic confinement on the synchronisation dynamics of the elastic compliance and the force modulation mechanisms and compare our results to the usual case with a single surface. Applying our results to the biologically relevant situation of nodal cilia, we show that force modulation is a mechanism that leads to phase-locked states under strong confinement that are very similar to those without confinement as a difference with the elastic compliance mechanism. Our results point therefore to the robustness of force modulation for synchronisation, an important feature for biological dynamics that therefore suggests it could be the most essential physical mechanism overall in arrays of nodal cilia. We further examine the distinct biologically-relevant situation of primary cilia and show in that case that the difference in robustness of the mechanisms is not as pronounced but still favours the force modulation.
We theoretically investigate the dynamics of model microswimmers in singular vortices, discover the existence of bounded orbits and use the model to successfully explain the previously observed depletion zone in bacterial suspensions.
We consider the dynamics of micro-sized, asymmetrically coated thermoresponsive hydrogel ribbons (microgels) under periodic heating and cooling in the confined space between two planar surfaces. As the result of the temperature changes, the volume and, thus, the shape of the slender microgel change, which leads to repeated cycles of bending and elastic relaxation, and to net locomotion. Small devices designed for biomimetic locomotion need to exploit flows that are not symmetric in time (non-reciprocal) to escape the constraints of the scallop theorem and undergo net motion. Unlike other biological slender swimmers, the non-reciprocal bending of the gel centerline is not sufficient here to explain for the overall swimming motion. We show instead that the swimming of the gel results from the flux of water periodically emanating from (or entering) the gel itself due to its shrinking (or swelling). The associated flows induce viscous stresses that lead to a net propulsive force on the gel. We derive a theoretical model for this hypothesis of jet-driven propulsion, which leads to excellent agreement with our experiments.
The Green's function of the incompressible Stokes equations, the stokeslet, represents the singular flow due to a point force. Its classical value in an unbounded fluid has been extended near surfaces of various shapes, including flat walls and spheres, and in most cases the presence of a surface leads to an advection flow induced at the location of the point force. In this paper, motivated by the biological transport of cargo along polymeric filaments inside eukaryotic cells, we investigate the reaction flow at the location of the point force due to a rigid slender filament located at a separation distance intermediate between the filament radius and its length (i.e. we compute the advection of the point force induced by the presence of the filament). An asymptotic analysis of the problem reveals that the leading-order approximation for the force distribution along the axis of the filament takes a form analogous to resistive-force theory but with drag coefficients that depend logarithmically on the distance between the point force and the filament. A comparison of our theoretical prediction with boundary element computations show good agreement. We finally briefly extend the model to the case of curved filaments.
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