Suspensions of aerobic bacteria often develop flows from the interplay of chemotaxis and buoyancy. We find in sessile drops that flows related to those in the Boycott effect of sedimentation carry bioconvective plumes down the slanted meniscus and concentrate cells at the drop edge, while in pendant drops such self-concentration occurs at the bottom. On scales much larger than a cell, concentrated regions in both geometries exhibit transient, reconstituting, high-speed jets straddled by vortex streets. A mechanism for large-scale coherence is proposed based on hydrodynamic interactions between swimming cells.

Bacterial processes ranging from gene expression to motility and biofilm formation are constantly challenged by internal and external noise. While the importance of stochastic fluctuations has been appreciated for chemotaxis, it is currently believed that deterministic long-range fluid dynamical effects govern cell-cell and cellsurface scattering-the elementary events that lead to swarming and collective swimming in active suspensions and to the formation of biofilms. Here, we report direct measurements of the bacterial flow field generated by individual swimming Escherichia coli both far from and near to a solid surface. These experiments allowed us to examine the relative importance of fluid dynamics and rotational diffusion for bacteria. For cell-cell interactions it is shown that thermal and intrinsic stochasticity drown the effects of long-range fluid dynamics, implying that physical interactions between bacteria are determined by steric collisions and near-field lubrication forces. This dominance of short-range forces closely links collective motion in bacterial suspensions to self-organization in driven granular systems, assemblages of biofilaments, and animal flocks. For the scattering of bacteria with surfaces, long-range fluid dynamical interactions are also shown to be negligible before collisions; however, once the bacterium swims along the surface within a few microns after an aligning collision, hydrodynamic effects can contribute to the experimentally observed, long residence times. Because these results are based on purely mechanical properties, they apply to a wide range of microorganisms.low Reynolds number hydrodynamics | microswimmers C ollective behavior of bacteria, such as biofilm formation (1), swarming (2), and turbulence-like motion in concentrated suspensions (3, 4), has profound effects on foraging, signaling, and transport of metabolites (5, 6), and can be of great biomedical importance (7,8). Large-scale coherence in bacterial systems typically arises from a combination of biochemical signaling (9) and physical interactions. Recent theoretical models that focus on physical aspects of bacterial dynamics identify pairwise long-range hydrodynamic interactions (10-16) as a key ingredient for collective swimming. Such ''microscopic" approaches underpin continuum theories that aim to describe the phenomenology of microbial suspensions (17-23). An assumption underlying many of these theories is that a self-propelled bacterium can be modeled as a force dipole; its body exerts a drag force F on the fluid that is balanced by the rearward flagellar thrust −F. The leading-order fluid velocity field at distance r is therefore a dipolar ''pusher" flow of magnitude u ∝ Fℓ∕ηr 2 (see streamlines in Fig. 1B), where η is the viscosity, and ℓ the distance between the forces (24, 25). While higher order corrections may be due to force-quadrupole contributions (26), the hypothesis that the leading-order flow field around a bacterium is dipolar has not yet been verified experimentally.A closely related, c...

Aerobic bacteria often live in thin fluid layers near solid-air-water contact lines, in which the biology of chemotaxis, metabolism, and cell-cell signaling is intimately connected to the physics of buoyancy, diffusion, and mixing. Using the geometry of a sessile drop, we demonstrate in suspensions of Bacillus subtilis the self-organized generation of a persistent hydrodynamic vortex that traps cells near the contact line. Arising from upward oxygentaxis and downward gravitational forcing, these dynamics are related to the Boycott effect in sedimentation and are explained quantitatively by a mathematical model consisting of oxygen diffusion and consumption, chemotaxis, and viscous fluid dynamics. The vortex is shown to advectively enhance uptake of oxygen into the suspension, and the wedge geometry leads to a singularity in the chemotactic dynamics near the contact line.bioconvection ͉ chemotaxis ͉ singularity ͉ Bacillus subtilis T he interplay of chemotaxis and diffusion of nutrients or signaling chemicals in bacterial suspensions can produce a variety of structures with locally high concentrations of cells, including phyllotactic patterns (1), filaments (2), and concentrations in fabricated microstructures (3). Less well explored are situations in which concentrating hydrodynamic f lows actually arise from these ingredients. Here we report a detailed experimental and theoretical study of an intriguing mechanism termed the ''chemotactic Boycott effect.'' Described brief ly before (4), it is intimately associated with buoyancy-driven f lows, metabolite diffusion, and slanted air-water menisci. The ubiquity of contact lines and their transport singularities (5) suggest importance of these observations in biofilm formation (6). The large-scale stirring created by these f lows illustrate important advective contributions to intercellular signaling, as in quorum sensing (7).The chemotactic Boycott effect takes its name from a phenomenon in sedimentation (8) that occurs when the chamber containing a fluid with settling particles is tilted from vertical. Settling depletes the fluid near the upper wall, making it buoyant relative to nearby fluid, whereupon it rises. This boundary flow stirs up the entire medium, greatly accelerating the settling process. In the chemotactic version, negatively buoyant aerobic bacteria swim up to the free surface of a sessile drop and slide down the slanted meniscus, producing high concentrations of cells near the three-phase contact line. In earlier work where this was observed (4), the detailed nature of hydrodynamic flows near the contact line was unclear. Here, by direct visualization and particle-imaging velocimetry (PIV), we show that the sliding surface layer drives a circulating hydrodynamic vortex in the meniscus region that is central to the microecology. Although counterintuitive in viscous flows, persistent circulation driven by forcing at the free surface is consistent with the classic analysis for vortex generation in wedge geometry (9).The initial discussion of the chemota...