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Register a new userThe role of correlations in the collective behaviour of microswimmer suspensions
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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.
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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.
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.
We develop a statistical framework for the rheology of dense, non-Brownian suspensions, based on correlations in a space representing forces, which is dual to position space. Working with the ensemble of steady state configurations obtained from simulations of suspensions in two dimensions, we find that the anisotropy of the pair correlation function in force space changes with confining shear stress ($sigma_{xy}$) and packing fraction ($phi$). Using these microscopic correlations, we build a statistical theory for the macroscopic friction coefficient: the anisotropy of the stress tensor, $mu = sigma_{xy}/P$. We find that $mu$ decreases (i) as $phi$ is increased and (ii) as $sigma_{xy}$ is increased. Using a new constitutive relation between $mu$ and viscosity for dense suspensions that generalizes the rate-independent one, we show that our theory predicts a Discontinuous Shear Thickening (DST) flow diagram that is in good agreement with numerical simulations, and the qualitative features of $mu$ that lead to the generic flow diagram of a DST fluid observed in experiments.
Suspensions of rear- and front-actuated microswimmers immersed in a fluid, known respectively as ``pushers and ``pullers, display qualitatively different collective behaviours: beyond a characteristic density, pusher suspensions exhibit a hydrodynamic instability leading to collective motion known as active turbulence, a phenomenon which is absent for pullers. In this Letter, we describe the collective dynamics of a binary pusher--puller mixture using kinetic theory and large-scale particle-resolved simulations. We derive and verify an instability criterion, showing that the critical density for active turbulence moves to higher values as the fraction $chi$ of pullers is increased and disappears for $chi geq 0.5$. We then show analytically and numerically that the two-point hydrodynamic correlations of the 1:1 mixture are equal to those of a suspension of noninteracting swimmers. Strikingly, our numerical analysis furthermore shows that the full probability distribution of the fluid velocity fluctuations collapses onto the one of a noninteracting system at the same density, where swimmer--swimmer correlations are strictly absent. Our results thus indicate that the fluid velocity fluctuations in 1:1 pusher--puller mixtures are exactly equal to those of the corresponding noninteracting suspension at any density, a surprising cancellation with no counterpart in equilibrium long-range interacting systems.
Dense suspensions of particles are relevant to many applications and are a key platform for developing a fundamental physics of out-of-equilibrium systems. They present challenging flow properties, apparently turning from liquid to solid upon small changes in composition or, intriguingly, in the driving forces applied to them. The emergent physics close to the ubiquitous jamming transition (and to some extent the glass and gelation transitions) provides common principles with which to achieve a consistent interpretation of a vast set of phenomena reported in the literature. In light of this, we review the current state of understanding regarding the relation between the physics at the particle scale and the rheology at the macroscopic scale. We further show how this perspective opens new avenues for the development of continuum models for dense suspensions.
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