Abstract

We formulate a Markovian response theory for the tumble rate of a bacterium moving in a chemical field and use it in the Smoluchowski equation. Based on a multipole expansion for the one-particle distribution function and a reaction-diffusion equation for the chemoattractant field, we derive a polarization extended model, which also includes the recently discovered angle bias. In the adiabatic limit we recover a generalized Keller–Segel equation with diffusion and chemotactic coefficients that depend on the microscopic swimming parameters. Requiring the tumble rate to be positive, our model introduces an upper bound for the chemotactic drift velocity, which is no longer singular as in the original Keller–Segel model. Solving the Keller–Segel equations numerically, we identify traveling bacterial concentration pulses, for which we do not need a second, signaling chemical field nor a singular chemotactic drift velocity as demanded in earlier publications. We present an extensive study of the traveling pulses and demonstrate how their speeds, widths, and heights depend on the microscopic parameters. Most importantly, we discover a maximum number of bacteria that the pulse can sustain—the maximum carrying capacity. Finally, by tuning our parameters, we are able to match the experimental realization of the traveling bacterial pulse.

Highlights

  • Collective motion of biological and artificial microswimmers shows a broad range of interesting phenomena as demonstrated in several review articles [1,2,3,4,5]

  • Requiring the tumble rate to be positive, our model introduces an upper bound for the chemotactic drift velocity, which is no longer singular as in the original Keller–Segel model

  • In the following we study in detail traveling bacterial pulses in an initally uniform density field of a chemoattractant by numerically solving both the polarization extended model (PE) of equations (12), (15) and (16) and the generalized Keller–Segel model (KS) of equations (18) and (19)

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Summary

Introduction

Collective motion of biological and artificial microswimmers shows a broad range of interesting phenomena as demonstrated in several review articles [1,2,3,4,5]. Our generalized Keller–Segel model is able to describe traveling bacterial pulses without the need neither for a singular chemotactic drift velocity nor for a second chemoattractant. V0 is the swimming velocity of the bacterium and χ0 is a unitless measure of the chemotactic strength It depends on integrals over the response function R and moments of the tumble angle distribution P(β). The linear dependence of the tumble rate λ on c (r) · e was already introduced by Schnitzer as the leading order for the angular variation of λ [61] It follows directly from equation (1) by choosing R (t - t¢) proportional to the time derivative of the δ function as demonstrated by Locsei in [63].

The Keller–Segel model as adiabatic limit
Bias of tumble angles
Rescaling the Keller–Segel equations
Details of numerical solution scheme
Traveling concentration pulses of bacteria
Findings
Discussion and conclusions
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