No Arabic abstract
We investigate theoretically the collective dynamics of a suspension of low Reynolds number swimmers that are confined to two dimensions by a thin fluid film. Our model swimmer is characterized by internal degrees of freedom which locally exert active stresses (force dipoles or quadrupoles) on the fluid. We find that hydrodynamic interactions mediated by the film can give rise to spontaneous continuous symmetry breaking (swarming), to states with either polar or nematic homogeneous order. For dipolar swimmers, the stroke averaged dynamics are enough to determine the leading contributions to the collective behaviour. In contrast, for quadrupolar swimmers, our analysis shows that detailed features of the internal dynamics play an important role in determining the bulk behaviour. In the broken symmetry phases, we investigate fluctuations of hydrodynamic variables of the system and find that these destabilize order. Interestingly, this instability is not generic and depends on length-scale.
Cellular appendages conferring motility, such as flagella or 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. To help uncover the physical mechanism for hydrodynamic synchronisation most essential overall in biology, we theoretically investigate the effect of strong confinement on the effectiveness of hydrodynamic synchronisation. We use minimal models 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. We calculate separately the impact of confinement on the synchronisation dynamics of the elastic compliance and the force modulation mechanisms and compare our results to the case with no confinement. 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 suggests it could be the most essential physical mechanism overall in arrays of nodal cilia. We further examine the distinct 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.
Social hierarchy is central to decision-making in the coordinated movement of many swarming species. Here we propose a hierarchical swarm model in the spirit of the Vicsek model of self-propelled particles. We show that, as the hierarchy becomes important, the swarming transition changes from the weak first-order transition observed for egalitarian populations, to a stronger first-order transition for intermediately strong hierarchies, and finally the discontinuity reduces till vanish, where the order-disorder transition appears to be absent in the extremely despotic societies. Associated to this we observe that the spatial structure of the swarm, as measured by the correlation between the density and velocity fields, is strongly mediated by the hierarchy. A two-group model and vectorial noise are also studied for verification. Our results point out the particular relevance of the hierarchical structures to swarming transitions when doing specific case studies.
Liquid-liquid phase separation occurs not only in bulk liquid, but also on surfaces. In physiology, the nature and function of condensates on cellular structures remain unexplored. Here, we study how the condensed protein TPX2 behaves on microtubules to initiate branching microtubule nucleation, which is critical for spindle assembly in eukaryotic cells. Using fluorescence, electron, and atomic force microscopies and hydrodynamic theory, we show that TPX2 on a microtubule reorganizes according to the Rayleigh-Plateau instability, like dew droplets patterning a spider web. After uniformly coating microtubules, TPX2 forms regularly spaced droplets from which branches nucleate. Droplet spacing increases with greater TPX2 concentration. A stochastic model shows that droplets make branching nucleation more efficient by confining the space along the microtubule where multiple necessary factors colocalize to nucleate a branch.
Suitable asymmetric microstructures can be used to control the direction of motion in microorganism populations. This rectification process makes it possible to accumulate swimmers in a region of space or to sort different swimmers. Here we study numerically how the separation process depends on the specific motility strategies of the microorganisms involved. Crucial properties such as the separation efficiency and the separation time for two bacterial strains are precisely defined and evaluated. In particular, the sorting of two bacterial populations inoculated in a box consisting of a series of chambers separated by columns of asymmetric obstacles is investigated. We show how the sorting efficiency is enhanced by these obstacles and conclude that this kind of sorting can be efficiently used even when the involved populations differ only in one aspect of their swimming strategy.
We show that the usual linear analysis of QGP Weibel instabilities based on the Maxwell-Boltzmann equation may be reproduced in a purely hydrodynamic model. The latter is derived by the Entropy Production Variational Method from a transport equation including collisions, and can describe highly nonequilibrium flow. We find that, as expected, collisions slow down the growth of Weibel instabilities. Finally, we discuss the strong momentum anisotropy limit.