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Register a new userActive turbulence in microswimmer suspensions -- the role of active hydrodynamic stress and volume exclusion
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Microswimmers exhibit an intriguing, highly-dynamic collective motion with large-scale swirling and streaming patterns, denoted as active turbulence -- reminiscent of classical high-Reynolds-number hydrodynamic turbulence. Various experimental, numerical, and theoretical approaches have been applied to elucidate similarities and differences to inertial hydrodynamic and active turbulence. These studies reveal a wide spectrum of possible structural and dynamical behaviors of active mesoscale systems, not necessarily consistent with the predictions of the Kolmogorov-Kraichnan theory of turbulence. We use squirmers embedded in a mesoscale fluid, modeled by the multiparticle collision dynamics (MPC) approach, to explore the collective behavior of bacteria-type microswimmers. Our model includes the active hydrodynamic stress generated by propulsion, and a rotlet dipole characteristic for flagellated bacteria. We find emergent clusters, activity-induced phase separation, and swarming, depending on density, active stress, and the rotlet dipole strength. The analysis of the squirmer dynamics in the swarming phase yields Kolomogorov-Kraichnan-type hydrodynamic turbulence and energy spectra for sufficiently high concentrations and strong rotlet dipoles. This emphasizes the paramount importance of the hydrodynamic flow field for swarming and bacterial turbulence.
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Tracer particles immersed in suspensions of biological microswimmers such as E. coli or Chlamydomonas display phenomena unseen in conventional equilibrium systems, including strongly enhanced diffusivity relative to the Brownian value and non-Gaussian displacement statistics. In dilute, 3-dimensional suspensions, these phenomena have typically been explained by the hydrodynamic advection of point tracers by isolated microswimmers, while, at higher concentrations, correlations between pusher microswimmers such as E. coli can increase the effective diffusivity even further. Anisotropic tracers in active suspensions can be expected to exhibit even more complex behaviour than spherical ones, due to the presence of a nontrivial translation-rotation coupling. Using large-scale lattice Boltzmann simulations of model microswimmers described by extended force dipoles, we study the motion of ellipsoidal point tracers immersed in 3-dimensional microswimmer suspensions. We find that the rotational diffusivity of tracers is much less affected by swimmer-swimmer correlations than the translational diffusivity. We furthermore study the anisotropic translational diffusion in the particle frame and find that, in pusher suspensions, the diffusivity along the ellipsoid major axis is higher than in the direction perpendicular to it, albeit with a smaller ratio than for Brownian diffusion. Thus, we find that far field hydrodynamics cannot account for the anomalous coupling between translation and rotation observed in experiments, as was recently proposed. Finally, we study the probability distributions (PDFs) of translational and rotational displacements. In accordance with experimental observations, for short observation times we observe strongly non-Gaussian PDFs that collapse when rescaled with their variance, which we attribute to the ballistic nature of tracer motion at short times.
In this Letter, we study the collective behaviour of a large number of self-propelled microswimmers immersed in a fluid. Using unprecedently large-scale lattice Boltzmann simulations, we reproduce the transition to bacterial turbulence. We show that, even well below the transition, swimmers move in a correlated fashion that cannot be described by a mean-field approach. We develop a novel kinetic theory that captures these correlations and is non-perturbative in the swimmer density. To provide an experimentally accessible measure of correlations, we calculate the diffusivity of passive tracers and reveal its non-trivial density dependence. The theory is in quantitative agreement with the lattice Boltzmann simulations and captures the asymmetry between pusher and puller swimmers below the transition to turbulence.
Large scale simulations and analytical theory have been combined to obtain the non-equilibrium velocity distribution, $f(v)$, of randomly accelerated particles in suspension. The simulations are based on an event-driven algorithm, generalised to include friction. They reveal strongly anomalous but largely universal distributions which are independent of volume fraction and collision processes, which suggests a one-particle model should capture all the essential features. We have formulated this one-particle model and solved it analytically in the limit of strong damping, where we find that $f(v)$ decays as $1/v$ for multiple decades, eventually crossing over to a Gaussian decay for the largest velocities. Many particle simulations and numerical solution of the one-particle model agree for all values of the damping.
Mechanical stress plays an intricate role in gene expression in individual cells and sculpting of developing tissues. However, systematic methods of studying how mechanical stress and feedback help to harmonize cellular activities within a tissue have yet to be developed. Motivated by our observation of the cellular constriction chains (CCCs) during the initial phase of ventral furrow formation in the Drosophila melanogaster embryo, we propose an active granular fluid (AGF) model that provides valuable insights into cellular coordination in the apical constriction process. In our model, cells are treated as circular particles connected by a predefined force network, and they undergo a random constriction process in which the particle constriction probability P is a function of the stress exerted on the particle by its neighbors. We find that when P favors tensile stress, constricted particles tend to form chain-like structures. In contrast, constricted particles tend to form compact clusters when P favors compression. A remarkable similarity of constricted-particle chains and CCCs observed in vivo provides indirect evidence that tensile-stress feedback coordinates the apical constriction activity. We expect that our particle-based AGF model will be useful in analyzing mechanical feedback effects in a wide variety of morphogenesis and organogenesis phenomena.
Active fluids exhibit spontaneous flows with complex spatiotemporal structure, which have been observed in bacterial suspensions, sperm cells, cytoskeletal suspensions, self-propelled colloids, and cell tissues. Despite occurring in the absence of inertia, chaotic active flows are reminiscent of inertial turbulence, and hence they are known as active turbulence. Here, we survey the field, providing a unified perspective over different classes of active turbulence. To this end, we divide our review in sections for systems with either polar or nematic order, and with or without momentum conservation (wet/dry). Comparing to inertial turbulence, we highlight the emergence of power-law scaling with either universal or non-universal exponents. We also contrast scenarios for the transition from steady to chaotic flows, and we discuss the absence of energy cascades. We link this feature to both the existence of intrinsic length scales and the self-organized nature of energy injection in active turbulence, which are fundamental differences with inertial turbulence. We close by outlining the emerging picture, remaining challenges, and future directions.
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