Recovering the activity parameters of an active fluid confined in a sphere.

The properties of an active fluid, for example, a bacterial bath or a collection of microtubules and molecular motors, can be accessed through the dynamics of passive particle probes. Here, in the perspective of analyzing experimental situations of confinement in droplets, we consider the kinematics of a negatively buoyant probe particle in an active fluid, both confined within a spherical domain. The active bath generates a fluctuating flow that pushes the particle with a velocity that is modeled as a colored stochastic noise, characterized by two parameters, the intensity and memory time of the active flow. When the particle departs a little from the bottom of the spherical domain, the configuration is well approximated by a particle in a two-dimensional harmonic trap subjected to the colored noise, in which case an analytical solution exists, which is the base for quantitative analysis. We numerically simulate the dynamics of the particle and use the planar, two-dimensional mean square displacement to recover the activity parameters of the bath. This approach yields satisfactory results as long as the particle remains relatively confined; that is, as long as the intensity of the colored noise remains low.

Frustrated 'run and tumble' of swimming Escherichia coli bacteria in nematic liquid crystals.

In many situations, bacteria move in complex environments, as soils, oceans or the human gut-track, where carrier fluids show complex structures associated with non-Newtonian rheology. Many fundamental questions concerning the ability to navigate in such environments remain unsolved. Recently, it has been shown that the kinetics of bacterial motion in structured fluids as liquid crystals (LCs) is constrained by the orientational molecular order (or director field) and that novel spatio-temporal patterns arise. A question unaddressed so far is how bacteria change swimming direction in such an environment. In this work, we study the swimming mechanism of a single bacterium, Esherichia coli, constrained to move along the director field of a lyotropic chromonic liquid crystal confined to a planar cell. Here, the spontaneous 'run and tumble' motion of the bacterium gets frustrated: the elasticity of the LC prevents flagella from unbundling. Interestingly, to change direction, bacteria execute a reversal motion along the director field, driven by the relocation of a single flagellum, a 'frustrated tumble'. We characterize this phenomenon in detail experimentally, exploiting exceptional spatial and temporal resolution of bacterial and flagellar dynamics, using a two colour Lagrangian tracking technique. We suggest a possible mechanism accounting for these observations.

Run-to-Tumble Variability Controls the Surface Residence Times of E. coli Bacteria.

Motile bacteria are known to accumulate at surfaces, eventually leading to changes in bacterial motility and biofilm formation. We use a novel two-color, three-dimensional Lagrangian tracking technique to follow simultaneously the body and the flagella of a wild-type Escherichia coli. We observe long surface residence times and surface escape corresponding mostly to immediately antecedent tumbling. A motility model accounting for a large behavioral variability in run-time duration reproduces all experimental findings and gives new insights into surface trapping efficiency.

Single-trajectory characterization of active swimmers in a flow.

We develop a maximum likelihood method to infer relevant physical properties of elongated active particles. Using individual trajectories of advected swimmers as input, we are able to accurately determine their rotational diffusion coefficients and an effective measure of their aspect ratio, also providing reliable estimators for the uncertainties of such quantities. We validate our theoretical construction using numerically generated active trajectories upon no flow, simple shear, and Poiseuille flow, with excellent results. Being designed to rely on single-particle data, our method eases applications in experimental conditions where swimmers exhibit a strong morphological diversity. We briefly discuss some of such ongoing experimental applications, specifically, in the characterization of swimming E. coli in a flow.

Chirality-induced bacterial rheotaxis in bulk shear flows.

Interaction of swimming bacteria with flows controls their ability to explore complex environments, crucial to many societal and environmental challenges and relevant for microfluidic applications such as cell sorting. Combining experimental, numerical, and theoretical analysis, we present a comprehensive study of the transport of motile bacteria in shear flows. Experimentally, we obtain with high accuracy and, for a large range of flow rates, the spatially resolved velocity and orientation distributions. They are in excellent agreement with the simulations of a kinematic model accounting for stochastic and microhydrodynamic properties and, in particular, the flagella chirality. Theoretical analysis reveals the scaling laws behind the average rheotactic velocity at moderate shear rates using a chirality parameter and explains the reorientation dynamics leading to saturation at large shear rates from the marginal stability of a fixed point. Our findings constitute a full understanding of the physical mechanisms and relevant parameters of bacteria bulk rheotaxis.

E. coli "super-contaminates" narrow ducts fostered by broad run-time distribution.

One notable feature of bacterial motion is their ability to swim upstream along corners and crevices, by leveraging hydrodynamic interactions. This motion through anatomic ducts or medical devices might be at the origin of serious infections. However, it remains unclear how bacteria can maintain persistent upstream motion while exhibiting run-and-tumble dynamics. Here, we demonstrate that Escherichia coli can travel upstream in microfluidic devices over distances of 15 mm in times as short as 15 min. Using a stochastic model relating the run times to the time that bacteria spend on surfaces, we quantitatively reproduce the evolution of the contamination profiles when considering a broad distribution of run times. The experimental data cannot be reproduced using the usually accepted exponential distribution of run times. Our study demonstrates that the run-and-tumble statistics determine macroscopic bacterial transport properties. This effect, which we name "super-contamination," could explain the fast onset of some life-threatening medical emergencies.

A combined rheometry and imaging study of viscosity reduction in bacterial suspensions.

Suspending self-propelled "pushers" in a liquid lowers its viscosity. We study how this phenomenon depends on system size in bacterial suspensions using bulk rheometry and particle-tracking rheoimaging. Above the critical bacterial volume fraction needed to decrease the viscosity to zero, [Formula: see text], large-scale collective motion emerges in the quiescent state, and the flow becomes nonlinear. We confirm a theoretical prediction that such instability should be suppressed by confinement. Our results also show that a recent application of active liquid-crystal theory to such systems is untenable.

Magnetotactic bacteria in a droplet self-assemble into a rotary motor.

From intracellular protein trafficking to large-scale motion of animal groups, the physical concepts driving the self-organization of living systems are still largely unraveled. Self-organization of active entities, leading to novel phases and emergent macroscopic properties, recently shed new light on these complex dynamical processes. Here we show that under the application of a constant magnetic field, motile magnetotactic bacteria confined in water-in-oil droplets self-assemble into a rotary motor exerting a torque on the external oil phase. A collective motion in the form of a large-scale vortex, reversable by inverting the field direction, builds up in the droplet with a vorticity perpendicular to the magnetic field. We study this collective organization at different concentrations, magnetic fields and droplet radii and reveal the formation of two torque-generating areas close to the droplet interface. We characterize quantitatively the mechanical energy extractable from this new biological and self-assembled motor.

Oscillatory surface rheotaxis of swimming E. coli bacteria.

Bacterial contamination of biological channels, catheters or water resources is a major threat to public health, which can be amplified by the ability of bacteria to swim upstream. The mechanisms of this 'rheotaxis', the reorientation with respect to flow gradients, are still poorly understood. Here, we follow individual E. coli bacteria swimming at surfaces under shear flow using 3D Lagrangian tracking and fluorescent flagellar labelling. Three transitions are identified with increasing shear rate: Above a first critical shear rate, bacteria shift to swimming upstream. After a second threshold, we report the discovery of an oscillatory rheotaxis. Beyond a third transition, we further observe coexistence of rheotaxis along the positive and negative vorticity directions. A theoretical analysis explains these rheotaxis regimes and predicts the corresponding critical shear rates. Our results shed light on bacterial transport and reveal strategies for contamination prevention, rheotactic cell sorting, and microswimmer navigation in complex flow environments.

Non-local rheology in dense granular flows: Revisiting the concept of fluidity.

The aim of this article is to discuss the concepts of non-local rheology and fluidity, recently introduced to describe dense granular flows. We review and compare various approaches based on different constitutive relations and choices for the fluidity parameter, focusing on the kinetic elasto-plastic model introduced by Bocquet et al. (Phys. Rev. Lett 103, 036001 (2009)) for soft matter, and adapted for granular matter by Kamrin et al. (Phys. Rev. Lett. 108, 178301 (2012)), and the gradient expansion of the local rheology µ(I) that we have proposed (Phys. Rev. Lett. 111, 238301 (2013)). We emphasise that, to discriminate between these approaches, one has to go beyond the predictions derived from linearisation around a uniform stress profile, such as that obtained in a simple shear cell. We argue that future tests can be based on the nature of the chosen fluidity parameter, and the related boundary conditions, as well as the hypothesis made to derive the models and the dynamical mechanisms underlying their dynamics.

Mechanical fluctuations suppress the threshold of soft-glassy solids: The secular drift scenario.

We propose a dynamical mechanism leading to the fluidization by external mechanical fluctuations of soft-glassy amorphous material driven below the yield stress. The model is based on the combination of memory effect and nonlinearity, leading to an accumulation of tiny effects over a long term. We test this scenario on a granular packing driven mechanically below the Coulomb threshold. We provide evidence for an effective viscous response directly related to small stress modulations in agreement with the theoretical prediction of a generic secular drift. We propose to extend this result more generally to a large class of glassy systems.

Turning Bacteria Suspensions into Superfluids.

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 evidence for a low-shear Newtonian plateau characterized by a shear viscosity decreasing with concentration. In the semidilute regime, for particularly active bacteria, the suspension displays a "superfluidlike" 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.

Living on the edge: transfer and traffic of E. coli in a confined flow.

We quantitatively study the transport of E. coli near the walls of confined microfluidic channels, and in more detail along the edges formed by the interception of two perpendicular walls. Our experiments establish the connection between bacterial motion at the flat surface and at the edges and demonstrate the robustness of the upstream motion at the edges. Upstream migration of E. coli at the edges is possible at much larger flow rates compared to motion at the flat surfaces. Interestingly, the speed of bacteria at the edges mainly results from collisions between bacteria moving along this single line. We show that upstream motion not only takes place at the edge but also in an "edge boundary layer" whose size varies with the applied flow rate. We quantify the bacterial fluxes along the bottom walls and the edges and show that they result from both the transport velocity of bacteria and the decrease of surface concentration with increasing flow rate due to erosion processes. We rationalize our findings as a function of local variations in the shear rate in the rectangular channels and hydrodynamic attractive forces between bacteria and walls.

Clogging transition of many-particle systems flowing through bottlenecks.

When a large set of discrete bodies passes through a bottleneck, the flow may become intermittent due to the development of clogs that obstruct the constriction. Clogging is observed, for instance, in colloidal suspensions, granular materials and crowd swarming, where consequences may be dramatic. Despite its ubiquity, a general framework embracing research in such a wide variety of scenarios is still lacking. We show that in systems of very different nature and scale -including sheep herds, pedestrian crowds, assemblies of grains, and colloids- the probability distribution of time lapses between the passages of consecutive bodies exhibits a power-law tail with an exponent that depends on the system condition. Consequently, we identify the transition to clogging in terms of the divergence of the average time lapse. Such a unified description allows us to put forward a qualitative clogging state diagram whose most conspicuous feature is the presence of a length scale qualitatively related to the presence of a finite size orifice. This approach helps to understand paradoxical phenomena, such as the faster-is-slower effect predicted for pedestrians evacuating a room and might become a starting point for researchers working in a wide variety of situations where clogging represents a hindrance.

Non-Newtonian viscosity of Escherichia coli suspensions.

The viscosity of an active suspension of E. coli bacteria is determined experimentally as a function of the shear rate using a Y-shaped microfluidic channel. From the relative suspension viscosity, we identify rheological thickening and thinning regimes as well as situations at low shear rate where the viscosity of the bacteria suspension can be lower than the viscosity of the suspending fluid. In addition, bacteria concentration and velocity profiles in the bulk are directly measured in the microchannel.

Nonlocal rheology of granular flows across yield conditions.

The rheology of dense granular flows is studied numerically in a shear cell controlled at constant pressure and shear stress, confined between two granular shear flows. We show that a liquid state can be achieved even far below the yield stress, whose flow can be described with the same rheology as above the yield stress. A nonlocal constitutive relation is derived from dimensional analysis through a gradient expansion and calibrated using the spatial relaxation of velocity profiles observed under homogeneous stresses. Both for frictional and frictionless grains, the relaxation length is found to diverge as the inverse square root of the distance to the yield point, on both sides of that point.

Hot spots in an athermal system.

We study experimentally the dynamical heterogeneities occurring at slow shear, in a model amorphous glassy material, i.e., a 3D granular packing. The deformation field is resolved spatially by using a diffusive wave spectroscopy technique. The heterogeneities show up as localized regions of strong deformations spanning a mesoscopic size of about 10 grains and called the "hot spots." The spatial clustering of hot spots is linked to the subsequent emergence of shear bands. Quantitatively, their appearance is associated with the macroscopic plastic deformation, and their rate of occurrence gives a physical meaning to the concept of "fluidity," recently used to describe the local and nonlocal rheology of soft glassy materials.

Creep and fluidity of a real granular packing near jamming.

We study the internal dynamical processes taking place in a granular packing below yield stress. At all packing fractions and down to vanishingly low applied shear, a logarithmic creep is observed. The experiments are analyzed using a viscoelastic model which introduces an internal, time-dependent, fluidity variable. For all experiments, the creep dynamics can be rescaled onto a unique curve which displays jamming at the random-close-packing limit. At each packing fraction, we measure a stress corresponding to the onset of internal granular reorganization and a slowing down of the creep dynamics before the final yield.

Enhanced diffusion due to active swimmers at a solid surface.

We consider two systems of active swimmers moving close to a solid surface, one being a living population of wild-type E. coli and the other being an assembly of self-propelled Au-Pt rods. In both situations, we have identified two different types of motion at the surface and evaluated the fraction of the population that displayed ballistic trajectories (active swimmers) with respect to those showing randomlike behavior. We studied the effect of this complex swimming activity on the diffusivity of passive tracers also present at the surface. We found that the tracer diffusivity is enhanced with respect to standard Brownian motion and increases linearly with the activity of the fluid, defined as the product of the fraction of active swimmers and their mean velocity. This result can be understood in terms of series of elementary encounters between the active swimmers and the tracers.

Mesoscopic length scale controls the rheology of dense suspensions.

From the flow properties of dense granular suspensions on an inclined plane, we identify a mesoscopic length scale strongly increasing with volume fraction. When the flowing layer height is larger than this length scale, a diverging Newtonian viscosity is determined. However, when the flowing layer height drops below this scale, we evidence a nonlocal effective viscosity, decreasing as a power law of the flow height. We establish a scaling relation between this mesoscopic length scale and the suspension viscosity. These results support recent theoretical and numerical results implying collective and clustered granular motion when the jamming point is approached from below.