Bacteria can chemotactically migrate up attractant gradients by controlling run-and-tumble motility patterns. In addition to this well-known chemotactic behaviour, several soil and marine bacterial species perform chemokinesis: they adjust their swimming speed according to the local concentration of chemoeffector, with higher speed at higher concentration. A field of attractant then induces a spatially varying swimming speed, which results in a drift towards lower attractant concentrations -- contrary to the drift created by chemotaxis. Here, to explore the biological benefits of chemokinesis and investigate its impact on the chemotactic response, we extend a Keller-Segel-type model to include chemokinesis. We apply the model to predict the dynamics of bacterial populations capable of chemokinesis and chemotaxis in chemoeffector fields inspired by microfluidic and agar plate migration assays. We find that chemokinesis combined with chemotaxis not only may enhance the population response with respect to pure chemotaxis, but also modifies it qualitatively. We conclude presenting predictions for bacteria around dynamic finite-size nutrient sources, simulating, e.g., a marine particle or a root. We show that chemokinesis can reduce the measuring bias that is created by a decaying attractant gradient.
The signaling apparatus mediating bacterial chemotaxis can adapt to a wide range of persistent external stimuli. In many cases, the bacterial activity returns to its pre-stimulus level exactly and this "perfect adaptability" is robust against variations in various chemotaxis protein concentrations. We model the bacterial chemotaxis signaling pathway, from ligand binding to CheY phosphorylation. By solving the steady-state equations of the model analytically, we derive a full set of conditions for the system to achieve perfect adaptation. The conditions related to the phosphorylation part of th…
{\sl Escherichia coli} ({\sl E. coli}) bacteria govern their trajectories by switching between running and tumbling modes as a function of the nutrient concentration they experienced in the past. At short time one observes a drift of the bacterial population, while at long time one observes accumulation in high-nutrient regions. Recent work has viewed chemotaxis as a compromise between drift toward favorable regions and accumulation in favorable regions. A number of earlier studies assume that a bacterium resets its memory at tumbles -- a fact not borne out by experiment -- and make use of app…
We consider the chemotaxis problem for a one-dimensional system. To analyze the interaction of bacteria and attractant we use a modified Keller-Segel model which accounts attractant absorption. To describe the system we use the chemotaxis sensitivity function, which characterizes nonuniformity of bacteria distribution. In particular, we investigate how the chemotaxis sensitivity function depends on the concentration of attractant at the boundary of the system. It is known that in the system without absorption the chemotaxis sensitivity function has a bell shape maximum. Here we show that attra…
This paper introduces a reaction-diffusion-chemotaxis model for bacterial aggregation patterns on the surface of thin agar plates. It is based on the non-linear degenerate cross diffusion model proposed by Kawasaki et al. (J. of Theor. Biol. 188(2) 1997) and it includes a suitable nutrient chemotactic term compatible with such type of diffusion. High resolution numerical simulations using Graphic Processing Units (GPUs) of the new model are presented, showing that the chemotactic term enhances the velocity of propagation of the colony envelope for dense-branching morphologies. In addition, the…
Classic experiments on the distribution of ducks around separated food sources found consistency with the `ideal free' distribution in which the local population is proportional to the local supply rate. Motivated by this experiment and others, we examine the analogous problem in the microbial world: the distribution of chemotactic bacteria around multiple nearby food sources. In contrast to the optimization of uptake rate that may hold at the level of a single cell in a spatially varying nutrient field, nutrient consumption by a population of chemotactic cells will modify the nutrient field, …
Many chemotactic bacteria inhabit environments in which chemicals appear as localized pulses and evolve by processes such as diffusion and mixing. We show that, in such environments, physical limits on the accuracy of temporal gradient sensing govern when and where bacteria can accurately measure the cues they use to navigate. Chemical pulses are surrounded by a predictable dynamic region, outside which bacterial cells cannot resolve gradients above noise. The outer boundary of this region initially expands in proportion to $\sqrt{t}$, before rapidly contracting. Our analysis also reveals how …
A Monte Carlo simulation of chemotactic bacteria is developed on the basis of the kinetic model and is applied to a one-dimensional traveling population wave in a microchannel. In this simulation, the Monte Carlo method, which calculates the run-and-tumble motions of bacteria, is coupled with a finite volume method to calculate the macroscopic transport of the chemical cues in the environment. The simulation method can successfully reproduce the traveling population wave of bacteria that was observed experimentally and reveal the microscopic dynamics of bacterium coupled with the macroscopic t…
We study the chemotaxis of a population of genetically identical swimming bacteria undergoing run and tumble dynamics driven by stochastic switching between clockwise and counterclockwise rotation of the flagellar rotary system. Understanding chemotaxis quantitatively requires that one links the switching rate of the rotary system in a gradient of chemoattractant/repellant to experimental measures of the efficiency of a population of bacteria in moving up/down the gradient. Here we achieve this by using a probabilistic model and show that the response of a population to the gradient is complex…
We study methylation dynamics of the chemoreceptors as an {\sl E.coli} cell moves around in a spatially varying chemo-attractant environment. We consider attractant concentration with strong and weak spatial gradient. During the uphill and downhill motion of the cell along the gradient, we measure the temporal variation of average methylation level of the receptor clusters. Our numerical simulations show that the methylation dynamics depends sensitively on the size of the receptor clusters and also on the strength of the gradient. At short times after the beginning of a run, the methylation dy…
Ephemeral aggregations of bacteria are ubiquitous in the environment, where they serve as hotbeds of metabolic activity, nutrient cycling, and horizontal gene transfer. In many cases, these regions of high bacterial concentration are thought to form when motile cells use chemotaxis to navigate to chemical hotspots. However, what governs the dynamics of bacterial aggregations is unclear. Here, we use a novel experimental platform to create realistic sub-millimeter scale nutrient pulses with controlled nutrient concentrations. By combining experiments, mathematical theory and agent-based simulat…
Bacterial chemotaxis is controlled by the signalling of a cluster of receptors. A cooperative model is presented, in which coupling between neighbouring receptor dimers enhances the sensitivity with which stimuli can be detected, without diminishing the range of chemoeffector concentration over which chemotaxis can operate. Individual receptor dimers have two stable conformational states: one active, one inactive. Noise gives rise to a distribution between these states, with the probability influenced by ligand binding, and also by the conformational states of adjacent receptor dimers. The two…
We study the migration of chemotactic wild-type Escherichia coli populations in semisolid (soft) agar in the concentration range C = 0.15-0.5% (w/v). For C < 0.35%, expanding bacterial colonies display characteristic chemotactic rings. At C = 0.35%, however, bacteria migrate as broad circular bands rather than sharp rings. These are growth/diffusion waves arising because of suppression of chemotaxis by the agar and have not been previously reported experimentally to our knowledge. For C = 0.4-0.5%, expanding colonies do not span the depth of the agar and develop pronounced front instabilities.…
We provide a detailed stochastic description of the swimming motion of an E.coli bacterium in two dimension, where we resolve tumble events in time. For this purpose, we set up two Langevin equations for the orientation angle and speed dynamics. Calculating moments, distribution and autocorrelation functions from both Langevin equations and matching them to the same quantities determined from data recorded in experiments, we infer the swimming parameters of E.coli . They are the tumble rate $λ$, the tumble time $r^{-1}$ , the swimming speed $v_0$ , the strength of speed fluctuations $σ$, the r…
Motivated by the observation of non-exponential run-time distributions of bacterial swimmers, we propose a minimal phenomenological model for taxis of active particles whose motion is controlled by an internal clock. The ticking of the clock depends on an external concentration field, e.g. a chemical substance. We demonstrate that these particles can detect concentration gradients and respond to them by moving up- or down-gradient depending on the clock design, albeit measurements of these fields are purely local in space and instantaneous in time. Altogether, our results open a new route in t…
Many motile biological cells navigate along concentration gradients of signaling molecules: This chemotaxis guides for instance sperm cells from marine invertebrates, which have to find egg cells in the ocean. While chemotaxis has been intensively studied for idealized conditions of rotationally symmetric gradients in still water, natural gradients are usually distorted, e.g., by turbulent flows in the ocean. Recent experiments and direct numerical simulations with sperm cells and bacteria surprisingly suggest the existence of an optimal flow strength at which chemotaxis is more effective than…
A generically observed mechanism that drives the self-organization of living systems is interaction via chemical signals among the individual elements -- which may represent cells, bacteria, or even enzymes. Here we propose a novel mechanism for such interactions, in the context of chemotaxis, which originates from the polarity of the particles and which generalizes the well-known Keller--Segel interaction term. We study the resulting large-scale dynamical properties of a system of such chemotactic particles using the exact stochastic formulation of Dean and Kawasaki along with dynamical renor…
We study chemotaxis of a single {\sl E.coli} bacterium in a medium where the nutrient chemical is also undergoing diffusion and its concentration has the form of a Gaussian whose width increases with time. We measure the average first passage time of the bacterium at a region of high nutrient concentration. In the limit of very slow nutrient diffusion, the bacterium effectively experiences a Gaussian concentration profile with a fixed width. In this case we find that there exists an optimum width of the Gaussian when the average first passage time is minimum, {\sl i.e.}, the search process is …
We study a generic model for the polarisation and motility of self-propelled soft objects, biological cells or biomimetic systems, interacting with a viscous substrate. The active forces generated by the cell on the substrate are modelled by means of oscillating force multipoles at the cell-substrate interface. Symmetry breaking and cell polarisation for a range of cell sizes naturally `emerge' from long range mechanical interactions between oscillating units, mediated both by the intracellular medium and the substrate. However, the harnessing of cell polarisation for motility requires substra…
Bacteria such as Escherichia coli move about in a series of runs and tumbles: while a run state (straight motion) entails all the flagellar motors spinning in counterclockwise mode, a tumble is caused by a shift in the state of one or more motors to clockwise spinning mode. In the presence of an attractant gradient in the environment, runs in the favourable direction are extended, and this results in a net drift of the organism in the direction of the gradient. The underlying signal transduction mechanism produces directed motion through a bi-lobed response function which relates the clockwise…
