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Register a new userEnhanced diffusion of nonswimmers in a three-dimensional bath of motile bacteria
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We show, using differential dynamic microscopy, that the diffusivity of non-motile cells in a three-dimensional (3D) population of motile E. coli is enhanced by an amount proportional to the active cell flux. While non-motile mutants without flagella and mutants with paralysed flagella have quite different thermal diffusivities and therefore hydrodynamic radii, their diffusivities are enhanced to the same extent by swimmers in the regime of cell densities explored here. Integrating the advective motion of non-swimmers caused by swimmers with finite persistence-length trajectories predicts our observations to within 2%, indicating that fluid entrainment is not relevant for diffusion enhancement in 3D.
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Chemotaxis of enzymes in response to gradients in the concentration of their substrate has been widely reported in recent experiments, but a basic understanding of the process is still lacking. Here, we develop a microscopic theory for chemotaxis, valid for enzymes and other small molecules. Our theory includes both non-specific interactions between enzyme and substrate, as well as complex formation through specific binding between the enzyme and the substrate. We find that two distinct mechanisms contribute to enzyme chemotaxis: a diffusiophoretic mechanism due to the non-specific interactions, and a new type of mechanism due to binding-induced changes in the diffusion coefficient of the enzyme. The latter chemotactic mechanism points towards lower substrate concentration if the substrate enhances enzyme diffusion, and towards higher substrate concentration if the substrate inhibits enzyme diffusion. For a typical enzyme, attractive phoresis and binding-induced enhanced diffusion will compete against each other. We find that phoresis dominates above a critical substrate concentration, whereas binding-induced enhanced diffusion dominates for low substrate concentration. Our results resolve an apparent contradiction regarding the direction of urease chemotaxis observed in experiments, and in general clarify the relation between enhanced diffusion and chemotaxis of enzymes. Finally, we show that the competition between the two distinct chemotactic mechanisms may be used to engineer nanomachines that move towards or away from regions with a specific substrate concentration.
Enzymes have been recently proposed to have mechanical activity associated with their chemical activity. In a number of recent studies, it has been reported that enzymes undergo enhanced diffusion in the presence of their corresponding substrate, when this substrate is uniformly distributed in solution. Moreover, if the concentration of the substrate is non-uniform, enzymes and other small molecules have been reported to show chemotaxis -- biased stochastic movement in the direction of the substrate gradient -- typically towards higher concentrations of this substrate, with a few exceptions. The underlying physical mechanisms responsible for enhanced diffusion and chemotaxis at the nanoscale, however, are still not well understood. Understanding these processes is important both for fundamental biological research, e.g. in the context of spatial organization of enzymes in metabolic pathways (metabolon formation), as well as for engineering applications, such as in the design of new vehicles for targeted drug delivery. In this Account, we will review the available experimental observations of both enhanced diffusion and chemotaxis, and we will discuss critically the different theories that have been proposed to explain the two. We first focus on enhanced diffusion, beginning with an overview of the experimental results. We then discuss the two main types of mechanisms that have been proposed, namely active mechanisms relying on the catalytic step of the enzymatic reaction, and equilibrium mechanisms which consider the reversible binding and unbinding of the substrate to the enzyme. We put particular emphasis on an equilibrium model recently introduced by us, which describes how the diffusion of dumbbell-like modular enzymes can be enhanced in the presence of substrate, thanks to a binding-induced reduction of the internal fluctuations of the enzyme. We then turn to chemotaxis, [...]
In the growth of bacterial colonies, a great variety of complex patterns are observed in experiments, depending on external conditions and the bacterial species. Typically, existing models employ systems of reaction-diffusion equations or consist of growth processes based on rules, and are limited to a discrete lattice. In contrast, the two-dimensional model proposed here is an off-lattice simulation, where bacteria are modelled as rigid circles and nutrients are point-like, Brownian particles. Varying the nutrient diffusion and concentration, we simulate a wide range of morphologies compatible with experimental observations, from round and compact to extremely branched patterns. A scaling relationship is found between the number of cells in the interface and the total number of cells, with two characteristic regimes. These regimes correspond to the compact and branched patterns, which are exhibited for sufficiently small and large colonies, respectively. In addition, we characterise the screening effect observed in the structures by analysing the multifractal properties of the growth probability.
We characterize cell motion in experiments and show that the transition to collective motion in colonies of gliding bacterial cells confined to a monolayer appears through the organization of cells into larger moving clusters. Collective motion by non-equilibrium cluster formation is detected for a critical cell packing fraction around 17%. This transition is characterized by a scale-free power-law cluster size distribution, with an exponent $0.88pm0.07$, and the appearance of giant number fluctuations. Our findings are in quantitative agreement with simulations of self-propelled rods. This suggests that the interplay of self-propulsion of bacteria and the rod-shape of bacteria is sufficient to induce collective motion.
Peritrichous bacteria such as Escherichia coli swim in viscous fluids by forming a helical bundle of flagellar filaments. The filaments are spatially distributed around the cell body to which they are connected via a flexible hook. To understand how the swimming direction of the cell is determined, we theoretically investigate the elastohydrodynamic motility problem of a multi-flagellated bacterium. Specifically, we consider a spherical cell body with a number N of flagella which are initially symmetrically arranged in a plane in order to provide an equilibrium state. We analytically solve the linear stability problem and find that at most 6 modes can be unstable and that these correspond to the degrees of freedom for the rigid-body motion of the cell body. Although there exists a rotation-dominated mode that generates negligible locomotion, we show that for the typical morphological parameters of bacteria the most unstable mode results in linear swimming in one direction accompanied by rotation around the same axis, as observed experimentally.
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