arXiv subjects

Eric Clément

Publications and source records attributed to Eric Clément.

At least 19 recordsLinked to original sources

arXiv2025

Confinement controls bacterial spreading at all scales

Navigation of microorganisms is controlled by internal processes ultimately sensitive to mechanical or chemical signaling encountered along the path. In many natural environments, such as porous soils or physiological ducts, motile species alternate between bulk and surface motion displaying in each case, distinct kinematics. This inherent complexity is key to many practical biological and ecological issues involving spreading and contamination, essential for understanding the spatiotemporal structuring of populations in their environment. However grasping the interplay between geometrical confinement and kinematics driven by internal biological responses remains poorly understood from a physical and biological standpoint. Here, we address this question through experimental and theoretical analysis in the heuristic situation of two parallel confining surfaces. We track wild-type E. coli - a model peritrichous flagellated bacterium - in 3D over extended periods of time. We obtain the first experimental measurements of the emerging diffusivity and bulk/surface residence times as a function of confinement height and the specific chiral kinematics at surfaces. All experimental results are quantitatively reproduced, without parametric adjustment, by a non-Markovian stochastic (BV) model that incorporates the internal biochemical memory carried by a phosphorylated protein switching the motor rotation. By matching the results with a Markovian (memoryless) companion model, we derive an analytical expression for the diffusivity and demonstrate how confining walls influence microbial long-range dispersion. This approach also provides a general conceptual basis for understanding how microorganisms navigate complex environments, in which their movement alternates between bulk and surfaces.

physics.bio-ph↗

arXiv2025

Active bacterial baths in droplets

Suspensions of self-propelled objects represent a novel paradigm in colloidal science. In such active baths traditional concepts, such as Brownian motion, fluctuation-dissipation relations, and work extraction from heat reservoirs, must be extended beyond the conventional framework of thermal baths. Unlike thermal baths, which are characterized by a single parameter, the temperature, the fundamental descriptors of an active bath remain elusive, especially in confined environments. In this study, buoyant, passive tracers are employed as generalized probes to investigate an active bath comprising motile bacteria confined within a droplet. We demonstrate that momentum transfer from the bath to the tracer can be effectively described as colored noise, characterized by temporal memory and an enhanced effective diffusivity significantly larger compared to thermal Brownian motion values. Using a stochastic analytical framework, we extract the temporal memory and diffusivity parameters that define such an active bath. Notably, the diffusivity scales linearly with bacterial concentration, modulated by a factor representing the role of confinement, expressed as the ratio of the confining radius to the probe radius. This finding, while still awaiting a complete theoretical explanation, offers new insights into the transport properties of confined active baths and paves the way for a deeper understanding of active emulsions driven by confined active matter.

cond-mat.soft↗

arXiv2024

Evolving motility of active droplets is captured by a self-repelling random walk model

Swimming droplets are a class of active particles whose motility changes as a function of time due to shrinkage and self-avoidance of their trail. Here we combine experiments and theory to show that our non-Markovian droplet (NMD) model, akin to a true self-avoiding walk [1], quantitatively captures droplet motion. We thus estimate the effective temperature arising from hydrodynamic flows and the coupling strength of the propulsion force as a function of fuel concentration. This framework explains a broad range of phenomena, including memory effects, solute-mediated interactions, droplet hovering above the surface, and enhanced collective diffusion.

cond-mat.soft↗

arXiv2024

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.

cond-mat.soft↗

arXiv2022

Run-to-Tumble Variability Controls the Surface Residence Times of ${\it E.~coli}$ Bacteria

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

cond-mat.soft↗

arXiv2022

Dispersion of motile bacteria in a porous medium

Understanding flow and transport of bacteria in porous media is crucial to technologies such as bioremediation, biomineralization or enhanced oil recovery. While physicochemical bacteria filtration is well-documented, recent studies showed that bacterial motility plays a key role in the transport process. Flow and transport experiments performed in microfluidic chips containing randomly placed obstacles confirmed that the distributions of non-motile particles stays compact, whereas for the motile strains, the distributions are characterized by both significant retention as well as fast downstream motion. For motile bacteria, the detailed microscopic study of individual bacteria trajectories reveals two salient features: (i) the emergence of an active retention process triggered by motility, (ii) enhancement of dispersion due to the exchange between fast flow channels and low flow regions in the vicinity of the solid grains. We propose a physical model based on a continuous time random walk approach. This approach accounts for bacteria dispersion via variable pore-scale flow velocities through a Markov model for equidistant particle speeds. Motility of bacteria is modeled by a two-rate trapping process that accounts for the motion towards and active trapping at the obstacles. This approach captures the forward tails observed for the distribution of bacteria displacements, and quantifies an enhanced hydrodynamic dispersion effect that originates in the interaction between flow at the pore-scale and bacterial motility. The model reproduces the experimental observations, and predicts bacteria dispersion and transport at the macroscale.

cond-mat.soft↗

arXiv2021

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.

cond-mat.soft↗

arXiv2019

3D spatial exploration by E. coli echoes motor temporal variability

Unraveling bacterial strategies for spatial exploration is crucial for understanding the complexity in the organization of life. Bacterial motility determines the spatio-temporal structure of microbial communities, controls infection spreading and the microbiota organization in guts or in soils. Most theoretical approaches for modeling bacterial transport rely on their run-and-tumble motion. For Escherichia coli, the run time distribution was reported to follow a Poisson process with a single characteristic time related to the rotational switching of the flagellar motors. However, direct measurements on flagellar motors show heavy-tailed distributions of rotation times stemming from the intrinsic noise in the chemotactic mechanism. Currently, there is no direct experimental evidence that the stochasticity in the chemotactic machinery affect the macroscopic motility of bacteria. In stark contrast with the accepted vision of run-and-tumble, here we report a large behavioral variability of wild-type \emph{E. coli}, revealed in their three-dimensional trajectories. At short observation times, a large distribution of run times is measured on a population and attributed to the slow fluctuations of a signaling protein triggering the flagellar motor reversal. Over long times, individual bacteria undergo significant changes in motility. We demonstrate that such a large distribution of run times introduces measurement biases in most practical situations. Our results reconcile the notorious conundrum between run time observations and motor switching statistics. We finally propose that statistical modeling of transport properties currently undertaken in the emerging framework of active matter studies, should be reconsidered under the scope of this large variability of motility features.

physics.bio-ph↗

arXiv2019

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. Selforganization of active entities, leading to novel phases and emergent macroscopic properties, recently shed new lights 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 droplets 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.

cond-mat.soft↗

arXiv2019

Swimming bacteria in Poiseuille flow: the quest for active Bretherton-Jeffery trajectories

Using a 3D Lagrangian tracking technique, we determine experimentally the trajectories of non-tumbling E. coli mutants swimming in a Poiseuille flow. We identify a typology of trajectories in agreement with a kinematic "active Bretherton-Jeffery" model, featuring an axi-symmetric self-propelled ellipsoid. In particular, we recover the "swinging" and "shear tumbling" kinematics predicted theoretically by Zöttl et al. Moreover using this model, we derive analytically new features such as quasi-planar piece-wise trajectories, associated with the high aspect ratio of the bacteria, as well as the existence of a drift angle around which bacteria perform closed cyclic trajectories. However, the agreement between the model predictions and the experimental results remains local in time, due to the presence of Brownian rotational noise.

physics.flu-dyn↗

arXiv2018

Effect of motility on the transport of bacteria populations through a porous medium

The role of activity on the hydrodynamic dispersion of bacteria in a model porous medium is studied by tracking thousands of bacteria in a microfluidic chip containing randomly placed pillars. We first evaluate the spreading dynamics of two populations of motile and non-motile bacteria injected at different flow rates. In both cases, we observe that the mean and the variance of the distances covered by the bacteria vary linearly with time and flow velocity, a result qualitatively consistent with the standard geometric dispersion picture. However, quantitatively, the motiles bacteria display a systematic retardation effect when compared to the non-motile ones. Furthermore, the shape of the traveled distance distribution in the flow direction differs significantly for both the motile and the non-motile strain, hence probing a markedly different exploration process. For the non-motile bacteria, the distribution is Gaussian whereas for the motile ones, the distribution displays a positive skewness and spreads exponentially downstream akin to a Gamma distribution. The detailed microscopic study of the trajectories reveals two salient effects characterizing the exploration process of motile bacteria : (i) The emergence of an "active" retention effect due to an extended exploration of the pore surfaces, (ii) an enhanced spreading at the forefront due to the transport of bacteria along "fast-tracks" where they acquire a velocity larger than the local flow velocity. We finally discuss the practical applications of these effects on the large-scale macroscopic transfer and contamination processes caused by microbes in natural environments.

physics.flu-dyn↗

arXiv2018

Actuated rheology of magnetic micro-swimmers suspensions : emergence of motor and brake states

We study the effect of magnetic field on the rheology of magnetic micro-swimmers suspensions. We use a model of a dilute suspension under simple shear and subjected to a constant magnetic field. Particle shear stress is obtained for both pusher and puller types of micro-swimmers. In the limit of low shear rate, the rheology exhibits a constant shear stress, called actuated stress, which only depends on the swimming activity of the particles. This stress is induced by the magnetic field and can be positive (brake state) or negative (motor state). In the limit of low magnetic fields, a scaling relation of the motor-brake effect is derived as a function of the dimensionless parameters of the model. In this case, the shear stress is an affine function of the shear rate. The possibilities offered by such an active system to control the rheological response of a uid are finally discussed.

physics.bio-ph↗

arXiv2015

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

We propose a dynamical mechanism leading to the fluidization of soft-glassy amorphous mate-rial driven below the yield-stress by external mechanical fluctuations. The model is based on the combination of memory effect and non-linearity, 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 bring evidences for an effective viscous response directly related to small stress modulations in agreement with the theoretical prediction of a generic secular drift.

cond-mat.soft↗

arXiv2015

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 bacteria 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 bacteria speed 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 bacteria 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 the local variations of the shear rate in the rectangular channels and hydrodynamic attractive forces between bacteria and walls.

cond-mat.soft↗

arXiv2015

Turning bacteria suspensions into a "superfluid"

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.

physics.flu-dyn↗

arXiv2015

Microrheology to probe non-local effects in dense granular flows

A granular material is observed to flow under the Coulomb yield criterion as soon as this criterion is satisfied in a remote but contiguous region of space. We investigate this non-local effect using discrete element simulations, in a geometry similar, in spirit, to the experiment of Reddy et al. (Phys. Rev. Lett., 106 (2011) 108301): a micro-rheometer is introduced to determine the influence of a distant shear band on the local rheological behaviour. The numerical simulations recover the dominant features of this experiment: the local shear rate is proportional to that in the shear band and decreases (roughly) exponentially with the distance to the yield conditions. The numerical results are in quantitative agreement with the predictions of the non-local rheology proposed by (Phys. Rev. Lett., 111 (2013) 238301) and derived from a gradient expansion of the rheology $μ[I]$. The consequences of these findings for the dynamical mechanisms controlling non-locality are finally discussed.

cond-mat.soft↗

arXiv2013

A non-local rheology for 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 non-local 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.

cond-mat.soft↗

arXiv2013

Particles accelerate the detachment of viscous liquids

During detachment of a viscous fluid extruded from a nozzle a filament linking the droplet to the latter is formed. Under the effect of surface tension the filament thins until pinch off and final detachment of the droplet. In this paper we study the effect of the presence of individual particles trapped in the filament on the detachment dynamics using granular suspensions of small volume fractions (ϕ < 6%). We show that even a single particle strongly modifies the detachment dynamics. The particle perturbs the thinning of the thread and a large droplet of fluid around the particle is formed. This perturbation leads to an acceleration of the detachment of the droplet compared to the detachment ob- served for a pure fluid. We quantify this acceleration for single particles of different sizes and link it to similar ob- servations for suspensions of small volume fractions. Our study also gives more insight into particulate effects on de- tachment of more dense suspensions and allows to explain the accelerated detachment close to final pinch off observed previously (Bonnoit et al 2012)

cond-mat.soft↗