In our article we present a computational model for the simulation of self-propelled anisotropic bacteria. To this end we use a self-propelled particle model and augment it with a statistical algorithm for the run-and-tumble motion. We derive an equation for the distribution of reorientations of the bacteria that we use to analyze the statistics of the random walk and that allows us to tune the behavior of our model to the characteristics of an E.coli bacterium. We validate our implementation in terms of a single swimmer and demonstrate that our model is capable of reproducing E.coli's run-and-tumble motion with excellent accuracy.
Suitable asymmetric microstructures can be used to control the direction of motion in microorganism populations. This rectification process makes it possible to accumulate swimmers in a region of space or to sort different swimmers. Here we study numerically how the separation process depends on the specific motility strategies of the microorganisms involved. Crucial properties such as the separation efficiency and the separation time for two bacterial strains are precisely defined and evaluated. In particular, the sorting of two bacterial populations inoculated in a box consisting of a series…
We solve the problem of first-passage time for run-and-tumble particles in one dimension. Exact expression is derived for the mean first-passage time in the general case, considering external force-fields and chemotactic-fields, giving rise to space dependent swim-speed and tumble rate. Agreement between theoretical formulae and numerical simulations is obtained in the analyzed case studies -- constant and sinusoidal force fields, constant gradient chemotactic field. Reported findings can be useful to get insights into very different phenomena involving active particles, such as bacterial moti…
Run-and-tumble dynamics is a wide-spread mechanism of swimming bacteria. The accumulation of run-and-tumble microswimmers near impermeable surfaces is studied theoretically and numerically in the low-density limit in two and three spatial dimensions. Both uni-modal and exponential distributions of the run lengths are considered. Constant run lengths lead to {peaks and depletions regions} in the density distribution of particles near the surface, in contrast to {exponentially-distributed run lengths}. Finally, we present a universal accumulation law for large channel widths, which applies not o…
Bacterial biofilms cost an enormous amount of resources in the health, medical, and industrial sectors. To understand early biofilm formation, beginning from planktonic states of active bacterial suspensions (such as Escherichia coli) to microcolonization, it is vital to study the mechanics of accumulation near surfaces and subsequent deposition. In this study, analytical expressions for the mean orientation, density and angular distributions, and deposition rates in such bacterial suspensions are derived, with and without the effects of external guiding or taxis fields. Simulations of confine…
Recent experiments on the green alga Chlamydomonas that swims using synchronized beating of a pair of flagella have revealed that it exhibits a run-and-tumble behavior similar to that of bacteria such as E. Coli. Using a simple purely hydrodynamic model that incorporates a stroke cycle and an intrinsic Gaussian white noise, we show that a stochastic run-and-tumble behavior could emerge, due to the nonlinearity of the combined synchronization-rotation-translation dynamics. This suggests the intriguing possibility that the alga might exploit nonlinear mechanics---as opposed to sophisticated bioc…
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…
{\it E. coli} bacteria swim in straight runs interrupted by sudden reorientation events called tumbles. The resulting random walks give rise to density fluctuations that can be derived analytically in the limit of non interacting particles or equivalently of very low concentrations. However, in situations of practical interest, the concentration of bacteria is always large enough to make interactions an important factor. Using molecular dynamics simulations, we study the dynamic structure factor of a model bacterial bath for increasing values of densities. We show that it is possible to reprod…
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.
The run-and-tumble walk, consisting in randomly reoriented ballistic excursions, models phenomena ranging from gas kinetics to bacteria motility. We evaluate the mean time required for this walk to find a fixed target within a 2D or 3D spherical confinement. We find that the mean search time admits a minimum as a function of the mean run duration for various types of boundary conditions and run duration distributions (exponential, power-law, deterministic). Our result stands in sharp contrast to the pure ballistic motion, which is predicted to be the optimal search strategy in the case of Pois…
We study the stochastic dynamics of a particle with two distinct motility states. Each one is characterized by two parameters: one represents the average speed and the other represents the persistence quantifying the tendency to maintain the current direction of motion. We consider a run-and-tumble process, which is a combination of an active fast motility mode (persistent motion) and a passive slow mode (diffusion). Assuming stochastic transitions between the two motility states, we derive an analytical expression for the time evolution of the mean square displacement. The interplay of the ke…
Run-and-Tumble particles, mimicking the behaviour of microorganisms like E. coli, are a paradigmatic model of active matter. Due to self-propulsion, their random and undirected motion can be rectified in a ratchet potential. Using perturbative field theory, we determine the shape of the potential that produces the maximum particle current as a function of the particles' parameters.
The spatial and orientational distribution in a dilute active suspension of non-Brownian run-and-tumble spherical swimmers confined between two planar hard walls is calculated theoretically. Using a kinetic model based on coupled bulk/surface probability density functions, we demonstrate the existence of a concentration wall boundary layer with thickness scaling with the run length, the absence of polarization throughout the channel, and the presence of sharp discontinuities in the bulk orientation distribution in the neighborhood of orientations parallel to the wall in the near-wall region. O…
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…
Absorption problems of run-and-tumble particles, described by the telegrapher's equation, are analyzed in one space dimension considering partially reflecting boundaries. Exact expressions for the probability distribution function in the Laplace domain and for the mean time to absorption are given, discussing some interesting limits (Brownian and wave limit, large volume limit) and different case studies (semi-infinite segment, equal and symmetric boundaries, totally/partially reflecting boundaries).
The effect of crowding on the run-and-tumble dynamics of swimmers such as bacteria is studied using a discrete lattice model of mutually excluding particles that move with constant velocity along a direction that is randomized at a rate $α$. In stationary state, the system is found to break into dense clusters in which particles are trapped or stopped from moving. The characteristic size of these clusters predominantly scales as $α^{-0.5}$ both in 1D and 2D. For a range of densities, due to cooperative effects, the stopping time scales as ${\cal T}_{1d}^{0.85}$ and as ${\cal T}_{2d}^{0.8}$, wh…
This work considers an extension of the Kuramoto model with run-and-tumble dynamics -- a type of self-propelled motion. The difference between the extended and the original model is that in the extended version angular velocity of individual particles is no longer fixed but can change sporadically with a new velocity drawn from a distribution g(w). Because the Kuramoto model undergoes a phase transition, it offers a simple case study for investigating phase transition for a system with self-propelled particles.
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…
The persistent character of the motion of active particles gives rise to accumulation at boundaries. I investigate the problem of run-and-tumble swimmers confined in a 1D box with hard walls, reporting expressions for the particles probability distribution and wall pressure. A crossover box length value is found below which the initial value of the pressure turns out to be higher than the asymptotic one, indicating a bounce effect of the active "wave" of swimmers. The case of attracting and repelling boundaries are also investigated using two different tumble rates for particles in the bulk an…
We use a first-passage time approach to study the statistics of the trapping times induced by persistent motion of active particles colliding with flat boundaries. The angular first-passage time distribution and mean first-passage time is calculated exactly for active Brownian and run-and-tumble particles and the results are compared. Theoretical predictions are in excellent agreement with Langevin simulations. Our results shed further light onto how active particles with different dynamics may be equivalent in the bulk, yet behave differently near boundaries or obstacles.
We study the transport of bacteria in a porous media modeled by a square channel containing one cylindrical obstacle via molecular dynamics simulations coupled to a lattice Boltzmann fluid. Our bacteria model is a rod-shaped rigid body which is propelled by a force-free mechanism. To account for the behavior of living bacteria, the model also incorporates a run-and-tumble process. The model bacteria are capable of hydrodynamically interacting with both of the channel walls and the obstacle. This enables the bacteria to get reoriented when experiencing a shear-flow. We demonstrate that this mod…
One of simplest examples of navigation found in nature is run-and-tumble chemotaxis. Tumbles reorient cells randomly, and cells can drift toward attractants or away from repellents by biasing the frequency of these events. The post-tumble swimming directions are typically correlated with those prior, as measured by the variance of the reorientation angle distribution. This variance can range from large, in the case of bacteria, to so small that tumble events are imperceptible, as observed in choanoflagellates. This raises the question of optimality: why is such a range of persistence observed …
We study the long-time asymptotic behavior of the position distribution of a run-and-tumble particle (RTP) in two dimensions and show that the distribution at a time $t$ can be expressed as a perturbative series in $(γt)^{-1}$, where $γ^{-1}$ is the persistence time of the RTP. We show that the higher order corrections to the leading order Gaussian distribution generically satisfy an inhomogeneous diffusion equation where the source term depends on the previous order solutions. The explicit solution of the inhomogeneous equation requires the position moments, and we develop a recursive formali…
We simulate by lattice Boltzmann the nonequilibrium steady states of run-and-tumble particles (inspired by a minimal model of bacteria), interacting by far-field hydrodynamics, subject to confinement. Under gravity, hydrodynamic interactions barely perturb the steady state found without them, but for particles in a harmonic trap such a state is quite changed if the run length is larger than the confinement length: a self-assembled pump is formed. Particles likewise confined in a narrow channel show a generic upstream flux in Poiseuille flow: chiral swimming is not required.
We study a generic model for quorum-sensing bacteria in circular confinement. Every bacterium produces signaling molecules, the local concentration of which triggers a response when a certain threshold is reached. If this response lowers the motility then an aggregation of bacteria occurs, which differs fundamentally from standard motility-induced phase separation due to the long-ranged nature of the concentration of signal molecules. We analyze this phenomenon analytically and by numerical simulations employing two different protocols leading to stationary cluster and ring morphologies, respe…
We study the large scale behavior of a collection of hard core run and tumble particles on a one dimensional lattice with periodic boundary conditions. Each particle has persistent motion in one direction decided by an associated spin variable until the direction of spin is reversed. We map the run and tumble model to a mass transfer model with fluctuating directed bonds. We calculate the steady state single site mass distribution in the mass model within a mean field approximation for larger spin-flip rates and by analyzing an appropriate coalescence fragmentation model for small spin-flip ra…
The random energy landscapes developed by speckle fields can be used to confine and manipulate a large number of micro-particles with a single laser beam. By means of molecular dynamics simulations, we investigate the static and dynamic properties of an active suspension of swimming bacteria embedded into speckle patterns. Looking at the correlation of the density fluctuations and the equilibrium density profiles, we observe a crossover phenomenon when the forces exerted by the speckles are equal to the bacteria's propulsion.
Understanding how bacteria move in porous media is critical to applications in healthcare, agriculture, environmental remediation, and chemical sensing. Recent work has demonstrated that E. coli, which moves by run-and-tumble dynamics in a homogeneous medium, exhibits a new form of motility when confined in a disordered porous medium: hopping-and-trapping motility, in which cells perform rapid, directed hops punctuated by intervals of slow, undirected trapping. Here, we use direct visualization to shed light on how these processes depend on pore-scale confinement and cellular activity. We find…
We consider an overdamped run-and-tumble particle in two dimensions, with self propulsion in an orientation that stochastically rotates by 90 degrees at a constant rate, clockwise or counter-clockwise with equal probabilities. In addition, the particle is confined by an external harmonic potential of stiffness $μ$, and possibly diffuses. We find the exact time-dependent distribution $P\left(x,y,t\right)$ of the particle's position, and in particular, the steady-state distribution $P_{\text{st}}\left(x,y\right)$ that is reached in the long-time limit. We also find $P\left(x,y,t\right)$ for a "f…
Many organisms navigate gradients by alternating straight motions (runs) with random reorientations (tumbles), transiently suppressing tumbles whenever attractant signal increases. This induces a functional coupling between movement and sensation, since tumbling probability is controlled by the internal state of the organism which, in turn, depends on previous signal levels. Although a negative feedback tends to maintain this internal state close to adapted levels, positive feedback can arise when motion up the gradient reduces tumbling probability, further boosting drift up the gradient. Impo…
One striking 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 E. coli can travel upstream in microfluidic devices over distances of 15 millimeters in times as short as 15 minutes. Using a stochastic model relating the run times to the time bacteria spend on surfaces, we…
We consider an active run-and-tumble particle (RTP) in $d$ dimensions and compute exactly the probability $S(t)$ that the $x$-component of the position of the RTP does not change sign up to time $t$. When the tumblings occur at a constant rate, we show that $S(t)$ is independent of $d$ for any finite time $t$ (and not just for large $t$), as a consequence of the celebrated Sparre Andersen theorem for discrete-time random walks in one dimension. Moreover, we show that this universal result holds for a much wider class of RTP models in which the speed $v$ of the particle after each tumbling is r…
We study a one-dimensional three-state run-and-tumble model motivated by the bacterium Caulobacter crescentus which displays a cell cycle between two non-proliferation mobile phase and a proliferating sedentary phase. Our model implements kinetic transitions between the two mobile and one sedentary states described in terms of their number densities, where mobility is allowed with different running speeds in forward and backward direction. We start by analyzing the stationary states of the system and compute the mean and squared-displacements for the distribution of all cells, as well as for t…
In this paper we develop a field-theoretic description for run and tumble chemotaxis, based on a density functional description of crystalline materials modified to capture orientational ordering. We show that this framework, with its in-built multi-particle interactions, soft-core repulsion and elasticity is ideal for describing continuum collective phases with particle resolution, but on diffusive timescales. We show that our model exhibits particle aggregation in an externally imposed constant attractant field, as is observed for phototactic or thermotactic agents. We also show that this mo…
