The run-and-tumble (RT) dynamics followed by bacterial swimmers gives rise first to a ballistic motion due to their persistence, and later, through consecutive tumbles, to a diffusive process. Here we investigate how long it takes for a dilute swimmer suspension to reach the diffusive regime as well as what is the amplitude of the deviations from the diffusive dynamics. A linear time dependence of the mean-squared displacement (MSD) is insufficient to characterize diffusion and thus we also focus on the excess kurtosis of the displacement distribution. Four swimming strategies are considered: (i) the conventional RT model with complete reorientation after tumbling, (ii) the case of partial reorientation, characterized by a distribution of tumbling angles, (iii) a run-and-reverse model with rotational diffusion, and (iv) a RT particle where the tumbling rate depends on the stochastic concentration of an internal protein. By analyzing the associated kinetic equations for the probability density function and simulating the models, we find that for models (ii), (iii), and (iv) the relaxation to diffusion can take much longer than the mean time between tumble events, evidencing the existence of large tails in the particle displacements. Moreover, the excess kurtosis can assume large positive values. In model (ii) it is possible for some distributions of tumbling angles that the MSD reaches a linear time dependence but, still, the dynamics remains non-Gaussian for long times. This is also the case in model (iii) for small rotational diffusivity. For all models, the long-time diffusion coefficients are also obtained. The theoretical approach, which relies on eigenvalue and angular Fourier expansions of the van Hove function, is in excellent agreement with the simulations.
{\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…
We consider self-propelled particles undergoing run-and-tumble dynamics (as exhibited by E. coli) in one dimension. Building on previous analyses at drift-diffusion level for the one-particle density, we add both interactions and noise, enabling discussion of domain formation by "self-trapping", and other collective phenomena. Mapping onto detailed-balance systems is possible in certain cases.
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…
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.
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 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…
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 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…
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…
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…
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…
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.
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.
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…
In the present work we simulate the basic two-dimensional dynamics of swarming E. coli bacteria on the surface of a moderately soft agar plate. Individual bacteria are modelled by self-propelled ridged bodies (agents), which interact with each other only through inelastic collision and with the highly viscous environment through damping forces. The motion of single agents is modelled closely corresponding to the behaviour of swimming bacteria. The dynamics of the model were adjusted to reproduce the experimental measurements of swimming E. coli K 12. Accordingly, simulations with loosely packe…
We consider the dynamics of self-propelled particles subject to external torques. Two models for the reorientation of self-propulsion are considered, run-and-tumble particles, and active Brownian particles. Using the standard tools of non-equilibrium statistical mechanics we show that the run and tumble particles have a more robust response to torques. This macroscopic signature of the underlying reorientation statistics can be used to differentiate between the two types of self propelled particles. Further this result might indicate that run and tumble motion is indeed the evolutionarily stab…
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…
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.
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 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…
Microbiology is the science of microbes, particularly bacteria. Many bacteria are motile: they are capable of self-propulsion. Among these, a significant class execute so-called run-and-tumble motion: they follow a fairly straight path for a certain distance, then abruptly change direction before repeating the process. This dynamics has something in common with Brownian motion (it is diffusive at large scales), and also something in contrast. Specifically, motility parameters such as the run speed and tumble rate depend on the local environment and hence can vary in space. When they do so, eve…
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 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…
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 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…
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…
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 …
E.coli serves as prototype for the study of peritrichous enteric bacteria that perform runs and tumbles alternately. Bacteria run forward as a result of the counterclockwise (CCW) rotation of their flagella bundle and perform tumbles when at least one of their flagella rotates clockwise (CW), moving away from the bundle. The flagella are hooked to molecular rotary motors of nanometric diameter able to make transitions between CCW and CW rotations that last up to one hundredth of a second. At the same time, flagella move or rotate the bacteria's body microscopically during lapses that range bet…
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 …
Micro-swimmers such as bacteria perform random walks known as run-and-tumbles to move up chemo-attractant gradients and as a result aggregate with others. It is also known that such micro-swimmers can self-organize into macroscopic patterns due to interactions with neighboring cells through the fluidic environment they live in. While the pattern formation resulting from chemotactic and hydrodynamic interactions separately and together have been previously investigated, the effect of the anisotropy in the tumbles of micro-swimmers has been unexplored. Here we show through linear analysis and fu…
We consider the tracer diffusion $D_{rr}$ that arises from the run-and-tumble motion of low Reynolds number swimmers, such as bacteria. Assuming a dilute suspension, where the bacteria move in uncorrelated runs of length $λ$, we obtain an exact expression for $D_{rr}$ for dipolar swimmers in three dimensions, hence explaining the surprising result that this is independent of $λ$. We compare $D_{rr}$ to the contribution to tracer diffusion from entrainment.
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 measu…
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…
Experiments have recently shown the feasibility of utilising bacteria as micro-scale robotic devices, with special attention paid to the development of bacteria-driven micro-swimmers taking advantage of built-in actuation and sensing mechanisms of cells. Here we propose a stochastic fluid dynamic model to describe analytically and computationally the dynamics of microscopic particles driven by the motion of surface-attached bacteria undergoing run-and-tumble motion. We compute analytical expressions for the rotational diffusion coefficient, the swimming speed and the effective diffusion coeffi…
Genetically engineered bacteria to increase the tumbling frequency of the run-and-tumble motion for the higher local bacterial density form visible stripe pattern composed of successive high and low density regions on an agar plate. We propose a model that includes a simplified regulatory dynamics of the tumbling frequency in individual cells to clarify the role of finite response time. We show that the time-delay due to the response dynamics results in the instability in a homogeneous steady state allowing a pattern formation. For further understanding, we propose a simplified two-state model…
We consider the tracer diffusion $D_{rr}$ that arises from the run-and-tumble motion of low Reynolds number swimmers, such as bacteria. In unbounded dilute suspensions, where the dipole swimmers move in uncorrelated runs of length $λ$, an exact solution showed that $D_{rr}$ is independent of $λ$. Here we verify this result in numerical simulations for a particular model swimmer, the spherical squirmer. We also note that in confined microenvironments, such as microscopic droplets, microfluidic devices and bacterial microzones in marine ecosystems, the size of the system can be comparable to $λ$…
The green alga {\it Chlamydomonas} swims with synchronized beating of its two flagella, and is experimentally observed to exhibit run-and-tumble behaviour similar to bacteria. Recently we studied a simple hydrodynamic three-sphere model of {\it Chlamydomonas} with a phase dependent driving force which can produce run-and-tumble behaviour when intrinsic noise is added, due to the non-linear mechanics of the system. Here, we consider the noiseless case and explore numerically the parameter space in the driving force profiles, which determine whether or not the synchronized state evolves from a g…
Microswimmers such as E. Coli bacteria accumulate and exhibit an intriguing dynamics near walls, governed by hydrodynamic and steric interactions. Insight into the underlying mechanisms and predominant interactions demand a detailed characterization of the entrapment process. We employ a mesoscale hydrodynamics simulation approach to study entrapment of a E. coli-type cell at a no-slip wall. The cell is modeled by a spherocylindrical body with several explicit helical flagella. Three stages of the entrapment process can be distinguished: the approaching regime, where a cell swims toward the wa…
We present a fast, high-throughput method for characterizing the motility of microorganisms in 3D based on standard imaging microscopy. Instead of tracking individual cells, we analyse the spatio-temporal fluctuations of the intensity in the sample from time-lapse images and obtain the intermediate scattering function (ISF) of the system. We demonstrate our method on two different types of microorganisms: bacteria, both smooth swimming (run only) and wild type (run and tumble) Escherichia coli, and the bi-flagellate alga Chlamydomonas reinhardtii. We validate the methodology using computer sim…
