Enkeleida Lushi

Fluid flows created by swimming bacteria drive self-organization in confined suspensions

Significance The collective dynamics of swimming microorganisms exhibits a complex interplay with the surrounding fluid: the motile cells stir the fluid, which in turn can reorient and advect them. This feedback loop can result in long-range interactions between the cells, an effect whose significance remains controversial. We present a computational model that takes into account these cell–fluid interactions and cell–cell forces and that predicts counterintuitive cellular order driven by long-range flows. This prediction is confirmed with experimental studies that track the orientation of cells in a confined, dense bacterial suspension. Concentrated suspensions of swimming microorganisms and other forms of active matter are known to display complex, self-organized spatiotemporal patterns on scales that are large compared with those of the individual motile units. Despite intensive experimental and theoretical study, it has remained unclear the extent to which the hydrodynamic flows generated by swimming cells, rather than purely steric interactions between them, drive the self-organization. Here we use the recent discovery of a spiral-vortex state in confined suspensions of Bacillus subtilis to study this issue in detail. Those experiments showed that if the radius of confinement in a thin cylindrical chamber is below a critical value, the suspension will spontaneously form a steady single-vortex state encircled by a counter-rotating cell boundary layer, with spiral cell orientation within the vortex. Left unclear, however, was the flagellar orientation, and hence the cell swimming direction, within the spiral vortex. Here, using a fast simulation method that captures oriented cell–cell and cell–fluid interactions in a minimal model of discrete particle systems, we predict the striking, counterintuitive result that in the presence of collectively generated fluid motion, the cells within the spiral vortex actually swim upstream against those flows. This prediction is then confirmed by the experiments reported here, which include measurements of flagella bundle orientation and cell tracking in the self-organized state. These results highlight the complex interplay between cell orientation and hydrodynamic flows in concentrated suspensions of microorganisms.

Collective dynamics in a binary mixture of hydrodynamically coupled microrotors.

We study, numerically, the collective dynamics of self-rotating nonaligning particles by considering a monolayer of spheres driven by constant clockwise or counterclockwise torques. We show that hydrodynamic interactions alter the emergence of large-scale dynamical patterns compared to those observed in dry systems. In dilute suspensions, the flow stirred by the rotors induces clustering of opposite-spin rotors, while at higher densities same-spin rotors phase separate. Above a critical rotor density, dynamic hexagonal crystals form. Our findings underscore the importance of inclusion of the many-body, long-range hydrodynamic interactions in predicting the phase behavior of active particles.

Microalgae Scatter off Solid Surfaces by Hydrodynamic and Contact Forces

Interactions between microorganisms and solid boundaries play an important role in biological processes, such as egg fertilization, biofilm formation, and soil colonization, where microswimmers move within a structured environment. Despite recent efforts to understand their origin, it is not clear whether these interactions can be understood as being fundamentally of hydrodynamic origin or hinging on the swimmers direct contact with the obstacle. Using a combination of experiments and simulations, here we study in detail the interaction of the biflagellate green alga Chlamydomonas reinhardtii, widely used as a model puller microorganism, with convex obstacles, a geometry ideally suited to highlight the different roles of steric and hydrodynamic effects. Our results reveal that both kinds of forces are crucial for the correct description of the interaction of this class of flagellated microorganisms with boundaries.

Periodic and Chaotic Orbits of Plane-Confined Micro-rotors in Creeping Flows

We explore theoretically the complex dynamics and emergent behaviors of spinning spheres immersed in viscous fluid. The particles are coupled to each other via the fluid in which they are suspended: Each particle disturbs the surrounding fluid with a rotlet field and that fluid flow affects the motion of the other particles. We notice the emergence of intricate periodic or chaotic trajectories that depend on the rotors initial position and separation. The point-rotor motions confined to a plane bear similarities to the classic 2D point-vortex dynamics. Our analyses highlight the complexity of the interaction between just a few rotors and suggest richer behavior in denser populations. We discuss how the model gives insight into more complex systems and suggest possible extensions for future theoretical studies.

Scattering of biflagellate microswimmers from surfaces

We use a three-bead-spring model to investigate the dynamics of biflagellate microswimmers near a surface. While the primary dynamics and scattering are governed by geometric-dependent direct contact, the fluid flows generated by the swimmer locomotion are important in orienting it toward or away from the surface. Flagellar noise and in particular cell spinning about the main axis help a surface-trapped swimmer escape, whereas the time a swimmer spends at the surface depends on the incident angle. The dynamics results from a nuanced interplay of direct collisions, hydrodynamics, noise, and the swimmer geometry. We show that to correctly capture the dynamics of a biflagellate swimmer, minimal models need to resolve the shape asymmetry.

Dynamics of inert spheres in active suspensions of micro-rotors

Inert particles suspended in active fluids of self-propelled particles are known to often exhibit enhanced diffusion and novel coherent structures. Here we numerically investigate the dynamical behavior and self-organization in a system consisting of passive and actively rotating spheres of the same size. The particles interact through direct collisions and the fluid flows generated as they move. In the absence of passive particles, three states emerge in a binary mixture of spinning spheres depending on particle fraction: a dilute gas-like state where the rotors move chaotically, a phase-separated state where like-rotors move in lanes or vortices, and a jammed state where crystals continuously assemble, melt and move (K. Yeo, E. Lushi, and P. M. Vlahovska, Phys. Rev. Lett., 2015, 114, 188301). Passive particles added to the rotor suspension modify the system dynamics and pattern formation: while states identified in the pure active suspension still emerge, they occur at different densities and mixture proportions. The dynamical behavior of the inert particles is also non-trivially dependent on the system composition.

Stability and dynamics of anisotropically-tumbling chemotactic swimmers

Microswimmers such as bacteria perform random walks known as run-and-tumbles to move up chemoattractant gradients and as a result aggregate with others. It is also known that such micro-swimmers can self-organize into macroscopic patterns due to interactions with neighboring cells through the fluidic environment they live in. While the pattern formation resulting from chemotactic and hydrodynamic interactions separately and together have been previously investigated, the effect of the anisotropy in the tumbles of microswimmers has been unexplored. Here we show through linear analysis and full nonlinear simulations that the slight anisotropy in the individual swimmer tumbles can alter the collective pattern formation in nontrivial ways. We show that tumbling anisotropy diminishes the magnitude of the chemotactic aggregates but may result in more such aggregation peaks.

Scattering of biflagellate micro-swimmers from surfaces

This work was supported in part by an Established Career Fellowship from the Engineering and Physical Sciences Research Council (REG).