The rheological response under simple shear of an active suspension of Escherichia coli is determined in a large range of shear rates and concentrations. The effective viscosity and the time scales characterizing the bacterial organization under shear are obtained. In the dilute regime, we bring evidences for a low shear Newtonian plateau characterized by a shear viscosity decreasing with concentration. In the semi-dilute regime, for particularly active bacteria, the suspension display a "super-fluid" like transition where the viscous resistance to shear vanishes, thus showing that macroscopically, the activity of pusher swimmers organized by shear, is able to fully overcome the dissipative effects due to viscous loss.
As a paradigmatic model of active fluids, bacterial suspensions show intriguing rheological responses drastically different from their counterpart colloidal suspensions. Although the flow of bulk bacterial suspensions has been extensively studied, the rheology of bacterial suspensions under confinement has not been experimentally explored. Here, using a microfluidic viscometer, we systematically measure the rheology of dilute E. coli suspensions under different degrees of confinement. Our study reveals a strong confinement effect: the viscosity of bacterial suspensions decreases substantially …
We demonstrate a novel shear-induced mechanism for growth of concentration fluctuations in a bacterial suspension. Using a linear stability analysis, a homogeneously sheared suspension is shown to support exponentially growing layering perturbations in the shear-rate and bacterial concentration. Non-linear simulations show that the instability eventually leads to gradient-banded velocity profiles, with a local depletion of bacteria at the interface between the bands. Our results show that long-ranged hydrodynamic interactions are sufficient to explain recent observations of shear-bands in bact…
Bacteria suspension exhibits a wide range of collective phenomena arising from interactions between individual cells. Here we show Serratia marcescens cells near an air-liquid interface spontaneously aggregate into dynamic clusters through surface-mediated hydrodynamic interactions. These long-lived clusters translate randomly and rotate in the counter-clockwise direction; they continuously evolve, merge with others and split into smaller ones. Measurements indicate that long-ranged hydrodynamic interactions have strong influences on cluster properties. Bacterial clusters change material and f…
We experimentally study density fluctuations and energy spectra of bulk \textit{E. coli} suspensions of different concentrations. Our results verify the predicted scaling law of giant number fluctuations in three-dimensional (3D) wet active fluids. We find that such a scaling behavior persists at small scales even in low concentration suspensions well below the transition concentration to active turbulence. Our experiments also support the prediction on the energy spectra of dilute pusher swimmers and illustrate the spectral properties of the active turbulence of dense bacterial suspensions in…
Swimming bacteria create long-range velocity fields that stir a large volume of fluid and move around passive particles dispersed in the fluid. Recent experiments and simulations have shown that long-time mean-squared displacement of passive particles in a bath of swimming bacteria exhibits diffusive behaviour with the effective diffusion coefficient significantly larger than its thermal counterpart. Comprehensive theoretical prediction of this effective diffusion coefficient and understanding of the enhancement mechanism remain a challenge. Here, we adapt the kinetic theory by Lin et al., J. …
Active matter is a new class of material, intrinsically out-of equilibrium with intriguing properties. So far, the recent upsurge of studies has mostly focused on the spontaneous behavior of these systems --in the absence of external constraints or driving--. Yet, many real life systems evolve under constraints, being both submitted to flow and various taxis. In the present work, we demonstrate a new experimental system which opens up the way for quantitative investigations, and discriminating examinations, of the challenging theoretical description of such systems. We explore the behavior of …
Near field hydrodynamic interactions are essential to determine many important emergent behaviors observed in active suspensions, but have not been successfully modeled so far. In this work we propose an effective model capable of efficiently capturing the essence of the near field hydrodynamic interactions, validated numerically by a pedagogic model system consisting of an E. coli and a spherical tracer. The proposed model effectively captures all the details of near field hydrodynamics through only a tensorial coefficient of resistance, which is fundamentally different from, and thus cannot …
We analyse the time dependent non-linear rheology of a dilute bacterial suspension (e.g. E. coli) for a pair of impulsively started linear flows - simple shear and uniaxial extension. The rheology is governed by the bacterium orientation distribution which satisfies a kinetic equation that includes rotation by the imposed flow, and relaxation to isotropy via rotary diffusion and tumbling. The relevant dimensionless parameters are the Peclet number $Pe\equiv \dotγτ$, which dictates the importance of flow-induced orientation anisotropy, and $τD_r$, which quantifies the relative importance of the…
We report a novel form of convection in suspensions of the bioluminiscent marine bacterium $Photobacterium~phosphoreum$. Suspensions of these bacteria placed in a chamber open to the air create persistent luminiscent plumes most easily visible when observed in the dark. These flows are strikingly similar to the classical bioconvection pattern of aerotactic swimming bacteria, which create an unstable stratification by swimming upwards to an air-water interface, but they are a puzzle since the strain of $P.~phosphoreum$ used does not express flagella and therefore cannot swim. Systematic experim…
In Stokesian Dynamics, particles are assumed to interact in two ways: through long-range mobility interactions and through short-range lubrication interactions. To speed up computations, in shear-driven concentrated suspensions, often found in rheometric contexts, it is common to consider only lubrication. We show that, although this approximation may provide acceptable results in shear-driven, periodic suspensions, for bidisperse suspensions where the particles are exposed to an external force, it can produce physically unreasonable results. We suggest that this problem could be mitigated by …
Sedimentation in active fluids has come into focus due to the ubiquity of swimming micro-organisms in natural and artificial environments. Here, we experimentally investigate sedimentation of passive particles in water containing various concentrations of the bacterium E. coli. Results show that the presence of living bacteria reduces the velocity of the sedimentation front even in the dilute regime, where the sedimentation velocity is expected to be independent of particle concentration. Bacteria increase the effective diffusion coefficient of the passive particles, which determines the width…
Swimming microorganisms rely on effective mixing strategies to achieve efficient nutrient influx. Recent experiments, probing the mixing capability of unicellular biflagellates, revealed that passive tracer particles exhibit anomalous non-Gaussian diffusion when immersed in a dilute suspension of self-motile Chlamydomonas reinhardtii algae. Qualitatively, this observation can be explained by the fact that the algae induce a fluid flow that may occasionally accelerate the colloidal tracers to relatively large velocities. A satisfactory quantitative theory of enhanced mixing in dilute active sus…
Active matter exhibits various forms of non-equilibrium states in the absence of external forcing, including macroscopic steady-state currents. Such states are often too complex to be modelled from first principles and our understanding of their physics relies heavily on minimal models. These have mostly been studied in the case of "dry" active matter, where particle dynamics are dominated by friction with their surroundings. Significantly less is known about systems with long-range hydrodynamic interactions that belong to "wet" active matter. Dilute suspensions of motile bacteria, modelled as…
Concentrated suspensions of swimming microorganisms and other forms of active matter are known to display complex, self-organized spatio-temporal patterns on scales large compared to those of the individual motile units. Despite intensive experimental and theoretical study, it has remained unclear the extent to which the hydrodynamic flows generated by swimming cells, rather than purely steric interactions between them, drive the self-organization. Here we utilize the recent discovery of a spiral-vortex state in confined suspensions of \textit{B. subtilis} to study this issue in detail. Those …
Dense suspensions of swimming bacteria are known to exhibit collective behaviour arising from the interplay of steric and hydrodynamic interactions. Unconfined suspensions exhibit transient, recurring vortices and jets, whereas those confinedin circular domains may exhibit order in the form of a spiral vortex. Here we show that confinement into a long and narrow macroscopic `racetrack' geometry stabilises bacterial motion to form a steady unidirectional circulation. This motion is reproduced in simulations of discrete swimmers that reveal the crucial role that bacteria-driven fluid flows play …
Active particles such as swimming bacteria or self-propelled colloids are known to spontaneously organize into fascinating large-scale dynamic structures. The emergence of these collective states from the motility pattern of the individual particles, typically a random walk, is yet to be probed in a well-defined synthetic system. Here, we report the experimental realization of intermittent colloidal motion that reproduces the run-and-tumble and Levy trajectories common to many swimming and swarming bacteria. Our strategy enables to tailor the sequence of repeated "runs" (nearly constant-speed …
Biofilms are ubiquitous macro-colonies of bacteria that develop at various interfaces (solid-liquid, solid-gas or liquid-gas). The formation of biofilms starts with the attachment of individual bacteria to an interface, where they proliferate and produce a slimy polymeric matrix - two processes that result in colony growth and spreading. Recent experiments on the growth of biofilms on agar substrates under air have shown that for certain bacterial strains, the production of the extracellular matrix and the resulting osmotic influx of nutrient-rich water from the agar into the biofilm are more …
A simple model for simulating flows of active suspensions is investigated. The approach is based on dissipative particle dynamics. While the model is potentially applicable to a wide range of self-propelled particle systems, the specific class of self-motile bacterial suspensions is considered as a modeling scenario. To mimic the rod-like geometry of a bacterium, two dissipative particle dynamics particles are connected by a stiff harmonic spring to form an aggregate dissipative particle dynamics molecule. Bacterial motility is modeled through a constant self-propulsion force applied along the…
Adding a non-adsorbing polymer to passive colloids induces an attraction between the particles via the `depletion' mechanism. High enough polymer concentrations lead to phase separation. We combine experiments, theory and simulations to demonstrate that using active colloids (such as motile bacteria) dramatically changes the physics of such mixtures. First, significantly stronger inter-particle attraction is needed to cause phase separation. Secondly, the finite size aggregates formed at lower inter-particle attraction show unidirectional rotation. These micro-rotors demonstrate the self assem…
This fluid dynamics video presents three-dimensional kinetic simulations of the dynamics in suspensions of oxytactic bacteria confined in thin liquid film surrounded by air. At the initial time, the bacterial concentration is uniform and isotropic, and there is no oxygen inside the film. The spatio-temporal dynamics of the oxygen and bacterial concentration are analyzed. For small film thicknesses, there is a weak migration of bacteria to the boundaries, and the oxygen concentration is high inside the film as a result of diffusion; both bacterial and oxygen concentrations quickly reach steady …
Suspensions of self-motile, elongated particles are a topic of significant current interest, exemplifying a form of `active matter'. Examples include self-propelling bacteria, algae and sperm, and artificial swimmers. Ericksen's model of a transversely-isotropic fluid [J. L. Ericksen, Colloid Polym. Sci. 173(2):117-122 (1960)] treats suspensions of non-motile particles as a continuum with an evolving preferred direction; this model describes fibrous materials as diverse as extracellular matrix, textile tufts and plant cell walls. Director-dependent effects are incorporated through a modified s…
Collective behaviour in suspensions of microswimmers is often dominated by the impact of long-ranged hydrodynamic interactions. These phenomena include active turbulence, where suspensions of pusher bacteria at sufficient densities exhibit large-scale, chaotic flows. To study this collective phenomenon, we use large-scale (up to $N=3\times 10^6$) particle-resolved lattice Boltzmann simulations of model microswimmers described by extended stresslets. Such system sizes enable us to obtain quantitative information about both the transition to active turbulence and characteristic features of the t…
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 conserva…
Cellular suspensions such as dense bacterial flows exhibit a turbulence-like phase under certain conditions. We study this phenomenon of "active turbulence" statistically by using numerical tools. Following Wensink et al. [Proc. Natl. Acad. Sci. U.S.A. 109, 14308 (2012)], we model active turbulence by means of a generalized Navier-Stokes equation. Two-point velocity statistics of active turbulence, both in the Eulerian and the Lagrangian frame, is explored. We characterize the scale-dependent features of two-point statistics in this system. Furthermore, we extend this statistical study with me…
