Derivation of the bacterial run-and-tumble kinetic equation from a model with biochemical pathway

Kinetic-transport equations are, by now, standard models to describe the dynamics of populations of bacteria moving by run-and-tumble. Experimental observations show that bacteria increase their run duration when encountering an increasing gradient of chemotactic molecules. This led to a first class of models which heuristically include tumbling frequencies depending on the path-wise gradient of chemotactic signal. More recently, the biochemical pathways regulating the flagellar motors were uncovered. This knowledge gave rise to a second class of kinetic-transport equations, that takes into account an intra-cellular molecular content and which relates the tumbling frequency to this information. It turns out that the tumbling frequency depends on the chemotactic signal, and not on its gradient. For these two classes of models, macroscopic equations of Keller-Segel type, have been derived using diffusion or hyperbolic rescaling. We complete this program by showing how the first class of equations can be derived from the second class with molecular content after appropriate rescaling. The main difficulty is to explain why the path-wise gradient of chemotactic signal can arise in this asymptotic process. Randomness of receptor methylation events can be included, and our approach can be used to compute the tumbling frequency in presence of such a noise

A pathway-based mean-field model for E. coli chemotaxis: Mathematical derivation and Keller-Segel limit

A pathway-based mean-field theory (PBMFT) was recently proposed for E. coli chemotaxis in [G. Si, T. Wu, Q. Quyang and Y. Tu, Phys. Rev. Lett., 109 (2012), 048101]. In this paper, we derived a new moment system of PBMFT by using the moment closure technique in kinetic theory under the assumption that the methylation level is locally concentrated. The new system is hyperbolic with linear convection terms. Under certain assumptions, the new system can recover the original model. Especially the assumption on the methylation difference made there can be understood explicitly in this new moment system. We obtain the Keller-Segel limit by taking into account the different physical time scales of tumbling, adaptation and the experimental observations. We also present numerical evidence to show the quantitative agreement of the moment system with the individual based E. coli chemotaxis simulator.Comment: 21 pages, 3 figure

Overview of mathematical approaches used to model bacterial chemotaxis II: bacterial populations

We review the application of mathematical modeling to understanding the behavior of populations of chemotactic bacteria. The application of continuum mathematical models, in particular generalized Keller–Segel models, is discussed along with attempts to incorporate the microscale (individual) behavior on the macroscale, modeling the interaction between different species of bacteria, the interaction of bacteria with their environment, and methods used to obtain experimentally verified parameter values. We allude briefly to the role of modeling pattern formation in understanding collective behavior within bacterial populations. Various aspects of each model are discussed and areas for possible future research are postulated

Volcano effect in chemotactic aggregation and an extended Keller-Segel mode

Aggregation of chemotactic bacteria under a unimodal distribution of chemical cues was investigated by Monte Carlo simulation and asymptotic analysis based on a kinetic transport equation, which considers an internal adaptation dynamics as well as a finite tumbling duration. It was found that there exist two different regimes of the adaptation time, between which the effect of the adaptation time on the aggregation behavior is reversed; that is, when the adaptation time is as small as the running duration, the aggregation becomes increasingly steeper as the adaptation time increases, while, when the adaptation time is as large as the diffusion time of the population density, the aggregation becomes more diffusive as the adaptation time increases. Moreover, notably, the aggregation profile becomes bimodal (volcano) at the large adaptation-time regime while it is always unimodal at the small adaptation-time regime. The volcano effect occurs in such a way that the population of tumbling cells considerably decreases in a diffusion layer which is created near the peak of the external chemical cues due to the coupling of diffusion and internal adaptation of the bacteria. Two different continuum-limit models are derived by the asymptotic analysis according to the scaling of the adaptation time; that is, at the small adaptation-time regime, the Keller-Segel model while, at the large adaptation-time regime, an extension of KS model, which involves both the internal variable and the tumbling duration. Importantly, either of the models can accurately reproduce the MC results at each adaptation-time regime, involving the volcano effect. Thus, we conclude that the coupling of diffusion, adaptation, and finite tumbling duration is crucial for the volcano effect

Pattern formation of a pathway-based diffusion model: linear stability analysis and an asymptotic preserving method

We investigate the linear stability analysis of a pathway-based diffusion model (PBDM), which characterizes the dynamics of the engineered Escherichia coli populations [X. Xue and C. Xue and M. Tang, P LoS Computational Biology, 14 (2018), pp. e1006178]. This stability analysis considers small perturbations of the density and chemical concentration around two non-trivial steady states, and the linearized equations are transformed into a generalized eigenvalue problem. By formal analysis, when the internal variable responds to the outside signal fast enough, the PBDM converges to an anisotropic diffusion model, for which the probability density distribution in the internal variable becomes a delta function. We introduce an asymptotic preserving (AP) scheme for the PBDM that converges to a stable limit scheme consistent with the anisotropic diffusion model. Further numerical simulations demonstrate the theoretical results of linear stability analysis, i.e., the pattern formation, and the convergence of the AP scheme