Motility is a fundamental survival strategy of bacteria to navigate porous environments. Swimming cells thrive in quiescent wetlands and sediments at the bottom of the marine water column, where they mediate many essential biogeochemical processes. While swimming motility in bulk fluid is now well established, a comprehensive understanding of the mechanisms regulating self-transport in the confined interstices of porous media is lacking, and determining the interactions between cells and surfaces of the solid matrix becomes paramount. Here, we precisely track the movement of bacteria (\emph{Magnetococcus marinus}) through a series of microfluidic porous media with broadly varying geometries and show that cell motility results in a succession of scattering events from the porous microstructure. Order or disorder can impact the cells' motility over short ranges, but we directly demonstrate that their large-scale transport properties are regulated by the cutoff of their persistent swimming, which is dictated primarily by the porosity and scale of the porous geometry. The effective mean free path is established as the key geometrical parameter controlling transport, and along with with minimal knowledge of cell swimming motility and surface scattering properties, we implement a theoretical model that universally predicts the effective diffusion of cells for the geometries studied here. These results are an important step toward predicting the physical ecology of swimming cells in quiescent porous media and understanding their role in environmental and health hazards in stagnant water.
In many situations bacteria move in complex environments, as for example in soils, oceans or the human gut-track microbiome. In these natural environments, carrier fluids such as mucus or reproductive fluids show complex structure associated with non-Newtonian rheology. Many fundamental questions concerning the the ability to navigate in such environments remain unsolved due to the inherent complexity of the natural surroundings. Recently, the interaction of swimming bacteria with nematic liquid crystals has attracted lot of attention. In these structured fluids, the kinetics of bacterial moti…
Active matter exhibits remarkable collective behavior in which flows, continuously generated by active particles, are intertwined with the orientational order of these particles. The relationship remains poorly understood as the activity and order are difficult to control independently. Here we demonstrate important facets of this interplay by exploring dynamics of swimming bacteria in a liquid crystalline environment with pre-designed periodic splay and bend in molecular orientation. The bacteria are expelled from the bend regions and condense into polar jets that propagate and transport carg…
Rigid body rotation and chiral reorientation combine in filamentous E. coli swimming in low-Re flows
Northeastern University（东北大学）；Rutgers Robert Wood Johnson Medical School（罗格斯大学罗伯特·伍德·约翰逊医学院）；Gyeongsang National University（庆尚国立大学）；The University of British Columbia（不列颠哥伦比亚大学）
Centrifugation is a widespread laboratory technique used to separate mixtures into fractions characterized by a specific size, weight or density. We demonstrate that centrifugation can be also used to separate swimming cells having different motility. To do this we study self-propelled bacteria under the influence of an external centrifugal field. Using dynamic image correlation spectroscopy we measure the spatially resolved motility of bacteria after centrifugation. A significant gradient in swimming-speeds is observed for increasing centrifugal speeds. Our results can be reproduced by a mode…
Burgeoning interest in the area of bacteria-powered micro robotic systems prompted us to study the dynamics of cargo transport by single bacteria. In this paper, we have studied the swimming behaviour of oil-droplets attached as a cargo to the cell bodies of single bacteria. The oil-droplet loaded bacteria exhibit super-diffusive motion which is characterized by high degree of directional persistence. Interestingly, bacteria could navigate even when loaded with oil-droplets as large as 8 microns with an effective increase in rotational drag by more than 2 orders when compared to free bacteria.…
We studied the swimming of \textit{Escherichia coli} bacteria in the vicinity of the critical point in a solution of the non-ionic surfactant \chem{C_{12}E_{5}} in buffer solution. In phase contrast microscopy, each swimming cell produces a transient trail behind itself lasting several seconds. Comparing quantitative image analysis with simulations show that these trails are due to local phase re-organisation triggered by differential adsorption. This contrasts with similar trails seen in bacteria swimming in liquid crystals, which are due to shear effects. We show how our trails are controlle…
It is well known that flagellated bacteria swim in circles near surfaces. However, recent experiments have shown that a sulfide-oxidizing bacterium named Thiovulum majus can transition from swimming in circles to a surface bound state where it stops swimming while remaining free to move laterally along the surface. In this bound state, the cell rotates perpendicular to the surface with its flagella pointing away from it. Using numerical simulations and theoretical analysis, we demonstrate the existence of a fluid-structure interaction instability that causes cells with relatively short flagell…
Bacteria in bulk fluids swim collectively and display fascinating emergent dynamics. Although bacterial collective swimming in three-dimensional (3D) geometries has been well studied, its counterpart in confined two-dimensional (2D) geometries relevant to natural habitats of bacteria is still poorly understood. Here, through carefully designed experiments on Escherichia coli in Hele-Shaw chambers, we show that a small change in the degree of confinement leads to a drastic change in bacterial collective swimming. While long-range nematic order emerges for bacteria that can cross during encounte…
Peritrichously-flagellated bacteria, such as Escherichia coli, self-propel in fluids by using specialised motors to rotate multiple helical filaments. The rotation of each motor is transmitted to a short flexible segment called the hook which in turn transmits it to a flagellar filament, enabling swimming of the whole cell. Since multiple motors are spatially distributed on the body of the organism, one would expect the propulsive forces from the filaments to push against each other leading to negligible swimming. We use a combination of computations and theory to show that the swimming of mul…
We show that the recent experimental observation of the rectification of swimming bacteria in a system with an array of asymmetric barriers occurs due to the ballistic component of the bacteria trajectories introduced by the bacterial "motor." Each bacteria selects a random direction for motion and then moves in this direction for a fixed period of time before randomly changing its orientation and moving in a new direction. In the limit where the bacteria undergo only Brownian motion, rectification by the barriers does not occur. We also examine the effects of steric interactions between the b…
We numerically study the dynamics of run-and-tumble particles confined in two chambers connected by thin channels. Two dominant dynamical behaviors emerge: (i) an oscillatory pumping state, in which particles periodically fill the two vessels and (ii) a circulating flow state, dynamically maintaining a near constant population level in the containers when connected by two channels. We demonstrate that the oscillatory behaviour arises from the combination of a narrow channel, preventing bacteria reorientation, and a density dependent motility inside the chambers.
Self-propelled bacteria are marvels of nature with a potential to power dynamic materials and microsystems of the future. The challenge is in commanding their chaotic behavior. By dispersing swimming Bacillus subtilis in a liquid-crystalline environment with spatially-varying orientation of the anisotropy axis, we demonstrate control over the distribution of bacteria, geometry and polarity of their trajectories. Bacteria recognize subtle differences in liquid crystal deformations, engaging in bipolar swimming in regions of pure splay and bend but switching to unipolar swimming in mixed splay-b…
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 e…
Bacterial contamination of biological conducts, catheters or water resources is a major threat to public health and can be amplified by the ability of bacteria to swim upstream. The mechanisms of this rheotaxis, the reorientation with respect to flow gradients, often in complex and confined environments, are still poorly understood. Here, we follow individual E. coli bacteria swimming at surfaces under shear flow with two complementary experimental assays, based on 3D Lagrangian tracking and fluorescent flagellar labelling and we develop a theoretical model for their rheotactic motion. Three t…
Small organisms (e.g., bacteria) and artificial microswimmers move due to a combination of active swimming and passive Brownian motion. Considering a simplified linear three-sphere swimmer, we study how the swimmer size regulates the interplay between self-driven and diffusive behavior at low Reynolds number. Starting from the Kirkwood-Smoluchowski equation and its corresponding Langevin equation, we derive formulas for the orientation correlation time, the mean velocity and the mean square displacement in three space dimensions. The validity of the analytical results is illustrated through nu…
Swimming-limited aggregation of bacteria in liquid crystals
Laboratoire de Physique et Mécanique des Milieux Hétérogènes (PMMH), ESPCI Paris, PSL University, CNRS, Sorbonne University, and Paris Cité University（异质介质物理与力学实验室，巴黎高等物理化工学院，PSL大学，法国国家科学研究中心，索邦大学，巴黎西岱大学）；Laboratoire Gulliver, ESPCI Paris, PSL University, CNRS（高卢实验室，巴黎高等物理化工学院，PSL大学，法国国家科学研究中心）；Institut Universitaire de France (IUF)（法国高等研究院）；Institute of Physics, University of Amsterdam（阿姆斯特丹大学物理研究所）
The behavior of flagellated bacteria swimming in non-Newtonian media remains an area with contradictory and conflicting results. We report on the behavior of wild-type and smooth-swimming E. coli in Newtonian, shear thinning and viscoelastic media, measuring their trajectories and swimming speed using a three dimensional real-time tracking microscope. We conclude that the speed enhancement in Methocel solution at higher concentration is due to shear-thinning and an analytical model is used to support our experimental result. We argue that shear-induced normal stresses reduce the wobbling behav…
We present a simple model for bacteria like \emph{Escherichia coli} swimming near solid surfaces. It consists of two spheres of different radii connected by a dragless rod. The effect of the flagella is taken into account by imposing a force on the tail sphere and opposite torques exerted by the rod over the spheres. The hydrodynamic forces and torques on the spheres are computed by considering separately the interaction of a single sphere with the surface and with the flow produced by the other sphere. Numerically, we solve the linear system which contains the geometrical constraints and the …
Near a solid boundary, E. coli swims in clockwise circular motion. We provide a hydrodynamic model for this behavior. We show that circular trajectories are natural consequences of force-free and torque-free swimming, and the hydrodynamic interactions with the boundary, which also leads to a hydrodynamic trapping of the cells close to the surface. We compare the results of the model with experimental data and obtain reasonable agreement. In particular, we show that the radius of curvature of the trajectory increases with the length of the bacterium body.
We study the collective dynamics of elongated swimmers in a very thin fluid layer by devising long, filamentous, non-tumbling bacteria. The strong confinement induces weak nematic alignment upon collision, which, for large enough density of cells, gives rise to global nematic order. This homogeneous but fluctuating phase, observed on the largest experimentally-accessible scale of millimeters, exhibits the properties predicted by standard models for flocking such as the Vicsek-style model of polar particles with nematic alignment: true long-range nematic order and non-trivial giant number fluct…
We study catalytic Janus swimmers and Escherichia coli bacteria swimming in a two-dimensional colloidal crystal. The Janus swimmers orbit individual colloids and hop between colloids stochastically, with a hopping rate that varies inversely with fuel (hydrogen peroxide) concentration. At high fuel concentration, these orbits are stable for 100s of revolutions, and the orbital speed oscillates periodically as a result of hydrodynamic, and possibly also phoretic, interactions between the swimmer and the six neighbouring colloids. Motile E.~coli bacteria behave very differently in the same colloi…
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 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. …
The sedimentation process in a suspension of bacteria is the result of the competition between gravity and the intrinsic motion of the microorganisms. We perform simulations of run-and-tumble "squirmers" that move in a fluid medium, focusing on the dependence of the non-equilibrium steady state on the bacterial swimming properties. We find that for high enough activity, the density profiles are no longer simple exponentials; we recover the numerical results via the introduction of a local effective temperature, suggesting that the breakdown of the Perrin-like exponential form is a collective e…
Flagellated bacteria exploiting helical propulsion are known to swim along circular trajectories near surfaces. Fluid dynamics predicts this circular motion to be clockwise (CW) above a rigid surface (when viewed from inside the fluid) and counter-clockwise (CCW) below a free surface. Recent experimental investigations showed that complex physicochemical processes at the nearby surface could lead to a change in the direction of rotation, both at solid surfaces absorbing slip-inducing polymers and interfaces covered with surfactants. Motivated by these results, we use a far-field hydrodynamic m…
In bacterial chemotaxis, E. coli cells drift up chemical gradients by a series of runs and tumbles. Runs are periods of directed swimming, and tumbles are abrupt changes in swimming direction. Near the beginning of each run, the rotating helical flagellar filaments which propel the cell form a bundle. Using resistive-force theory, we show that the counter-rotation of the cell body necessary for torque balance is sufficient to wrap the filaments into a bundle, even in the absence of the swirling flows produced by each individual filament.
Active matter comprised of self-propelled interacting units holds a major promise for extraction of useful work from its seemingly chaotic out-of-equilibrium dynamics. Streamlining active matter to produce work is especially important at microscale, where the viscous forces prevail over inertia and the useful modes of transport require very specific non-reciprocal type of motion. Here we report that microscopic active droplets representing aqueous dispersions of swimming bacteria Bacillus subtilis show unidirectional propulsion when placed in an inactive nematic medium. Random motion of bacter…
In this work we introduce a stochastic model to describe directional changes in the movement of swimming bacteria. We use the probability density function (PDF) of turn angles, measured on tumbling wild-type {\it E. coli}, to build a Langevin equation for the deflection of the bacterial body swimming in isotropic media. We have solved this equation analytically by means of the Green function method and shown that three parameters are sufficient to describe the movement: the characteristic time, the steady-state solution and the control parameter. We conclude that the tumble motion, which is ma…
The run and tumble motions of a swimming bacterium are well characterized by two stochastic variables: the speed $v(t)$ and the change of direction or deflection \mbox{$x(t)=\cosφ(t)$}, where $φ(t)$ is the turning angle at time $t$. Recently, we have introduced [Soft Matter {\bf 13}, 3385 (2017)] a single stochastic model for the deflection $x(t)$ of an {\sl E. coli} bacterium performing both types of movement in isotropic media without taxis, based on available experimental data. In this work we introduce Langevin equations for the variables $(v,x)$, which for particular values of a control p…
The natural habitats of microorganisms in the human microbiome and ocean and soil ecosystems are full of colloids and macromolecules, which impart non-Newtonian flow properties drastically affecting the locomotion of swimming microorganisms. Although the low-Reynolds-number hydrodynamics of the swimming of flagellated bacteria in simple Newtonian fluids has been well developed, our understanding of bacterial motility in complex non-Newtonian fluids is still primitive. Even after six decades of research, fundamental questions about the nature and origin of bacterial motility enhancement in poly…
The swimming behavior of bacteria and other microorganisms is sensitive to the physical properties of the fluid in which they swim. Mucus, biofilms, and artificial liquid-crystalline solutions are all examples of fluids with some degree of anisotropy that are also commonly encountered by bacteria. In this article, we study how liquid-crystalline order affects the swimming behavior of a model swimmer. The swimmer is a one-dimensional version of G. I. Taylor's swimming sheet: an infinite line undulating with small-amplitude transverse or longitudinal traveling waves. The fluid is a two-dimension…
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 …
Magnetotactic swimmers tend to align along magnetic field lines against stochastic reorientations. We show that the swimming strategy, e.g. active Brownian motion versus run-and-tumble dynamics, strongly affects the orientation statistics. The latter can exhibit a velocity condensation whereby the alignment probability density diverges. As a consequence, we find that the swimming strategy affects the nature of the phase transition to collective motion, indicating that Lévy run-and-tumble walks can outperform active Brownian processes as strategies to trigger collective behavior.
The near-surface swimming patterns of bacteria are strongly determined by the hydrodynamic interactions between bacteria and the surface, which trap bacteria in smooth circular trajectories that lead to inefficient surface exploration. Here, we show by combining experiments and a data-driven mathematical model that surface exploration of enterohemorrhagic Escherichia coli (EHEC) -- a pathogenic strain of E. coli causing serious illnesses such as bloody diarrhea -- results from a complex interplay between motility and transient surface adhesion events. These events allow EHEC to break the smoot…
A growing body of work aims at designing and testing micron-scale synthetic swimmers. One method, inspired by the locomotion of flagellated bacteria, consists of applying a rotating magnetic field to a rigid, helically-shaped, propeller attached to a magnetic head. When the resulting device, termed an artificial bacteria flagellum, is aligned perpendicularly to the applied field, the helix rotates and the swimmer moves forward. Experimental investigation of artificial bacteria flagella shows that at low frequency of the applied field, the axis of the helix does not align perpendicularly to the…
External control of the swimming speed of `active particles' can be used to self assemble designer structures in situ on the micrometer to millimeter scale. We demonstrate such reconfigurable templated active self assembly in a fluid environment using light powered strains of Escherichia coli. The physics and biology controlling the sharpness and formation speed of patterns is investigated using a bespoke fast-responding strain.
Suspensions of motile cells are model systems for understanding the unique mechanical properties of living materials which often consist of ensembles of self-propelled particles. We present here a quantitative comparison of theory against experiment for the rheology of such suspensions. The influence of motility on viscosities of cell suspensions is studied using a novel acoustically-driven microfluidic capillary-breakup extensional rheometer. Motility increases the extensional viscosity of suspensions of algal pullers, but decreases it in the case of bacterial or sperm pushers. A recent model…
Peritrichous bacteria such as Escherichia coli swim in viscous fluids by forming a helical bundle of flagellar filaments. The filaments are spatially distributed around the cell body to which they are connected via a flexible hook. To understand how the swimming direction of the cell is determined, we theoretically investigate the elastohydrodynamic motility problem of a multi-flagellated bacterium. Specifically, we consider a spherical cell body with a number N of flagella which are initially symmetrically arranged in a plane in order to provide an equilibrium state. We analytically solve the…
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 …
E. coli bacteria swim following a run and tumble pattern. In the run state all flagella join in a single helical bundle that propels the cell body along approximately straight paths. When one or more flagellar motors reverse direction the bundle unwinds and the cell randomizes its orientation. This basic picture represents an idealization of a much more complex dynamical problem. Although it has been shown that bundle formation can occur at either pole of the cell, it is still unclear whether this two run states correspond to asymmetric propulsion features. Using holographic microscopy we reco…
Bacterial chemotaxis is one of the most extensively studied adaptive responses in cells. Many bacteria are able to bias their apparently random motion to produce a drift in the direction of the increasing chemoattractant concentration. It has been recognized that the particular motility pattern employed by moving bacteria has a direct impact on the efficiency of chemotaxis. The linear theory of chemotaxis pioneered by de Gennes allows for calculation of the drift velocity in small gradients for bacteria with basic motility patterns. However, recent experimental data on several bacterial specie…
Cells swimming in confined environments are attracted by surfaces. We measure the steady-state distribution of smooth-swimming bacteria (Escherichia coli) between two glass plates. In agreement with earlier studies, we find a strong increase of the cell concentration at the boundaries. We demonstrate theoretically that hydrodynamic interactions of the swimming cells with solid surfaces lead to their re-orientation in the direction parallel to the surfaces, as well as their attraction by the closest wall. A model is derived for the steady-state distribution of swimming cells, which compares fav…
