This paper reports an investigation of dynamical behaviors of motile rod-shaped bacteria within anisotropic viscoelastic environments defined by lyotropic liquid crystals (LCs). In contrast to passive microparticles (including non-motile bacteria) that associate irreversibly in LCs via elasticity-mediated forces, we report that motile Proteus mirabilis bacteria form dynamic and reversible multi-cellular assemblies when dispersed in a lyotropic LC. By measuring the velocity of the bacteria through the LC (8.8 +/− 0.2 μm/s) and by characterizing the ordering of the LC about the rod-shaped bacteria (tangential anchoring), we conclude that the reversibility of the inter-bacterial interaction emerges from the interplay of forces generated by the flagella of the bacteria and the elasticity of the LC, both of which are comparable in magnitude (tens of pN) for motile Proteus mirabilis cells. We also measured the dissociation process, which occurs in a direction determined by the LC, to bias the size distribution of multi-cellular bacterial complexes in a population of motile Proteus mirabilis relative to a population of non-motile cells. Overall, these observations and others reported in this paper provide insight into the fundamental dynamical behaviors of bacteria in complex anisotropic environments and suggest that motile bacteria in LCs are an exciting model system for exploration of principles for the design of active materials.

We report on the organization and dynamics of bacteria (Proteus mirabilis) dispersed within lyotropic liquid crystal (LC) films confined by pairs of surfaces that induce homeotropic (perpendicular) or hybrid (homeotropic and parallel orientations at each surface) anchoring of the LC. By using motile vegetative bacteria (3 µm in length) and homeotropically aligned LC films with thicknesses that exceed the length of the rod-shaped cells, a key finding reported in this paper is that elastic torques generated by the LC are sufficiently large to overcome wall-induced hydrodynamic torques acting on the cells, thus leading to LC-guided bacterial motion near surfaces that orient LCs. This result extends to bacteria within LC films with hybrid anchoring, and leads to the observation that asymmetric strain within a hybrid aligned LC rectifies motions of motile cells. In contrast, when the LC film thickness is sufficiently small that confinement prevents alignment of the bacteria cells along a homeotropically aligned LC director (achieved using swarm cells of length 10-60 µm), the bacterial cells propel in directions orthogonal to the director, generating transient distortions in the LC that have striking "comet-like" optical signatures. In this limit, for hybrid LC films, we find LC elastic stresses deform the bodies of swarm cells into bent configurations that follow the LC director, thus unmasking a coupling between bacterial shape and LC strain. Overall, these results provide new insight into the influence of surface-oriented LCs on dynamical bacterial behaviors and hint at novel ways to manipulate bacteria using confined LC phases that are not possible in isotropic solutions.

Bacteria often inhabit and exhibit distinct dynamical behaviors at interfaces, but the physical mechanisms by which interfaces cue bacteria are still poorly understood. In this work, we use interfaces formed between coexisting isotropic and liquid crystal (LC) phases to provide insight into how mechanical anisotropy and defects in LC ordering influence fundamental bacterial behaviors. Specifically, we measure the anisotropic elasticity of the LC to change fundamental behaviors of motile, rod-shaped Proteus mirabilis cells (3 μm in length) adsorbed to the LC interface, including the orientation, speed, and direction of motion of the cells (the cells follow the director of the LC at the interface), transient multicellular self-association, and dynamical escape from the interface. In this latter context, we measure motile bacteria to escape from the interfaces preferentially into the isotropic phase, consistent with the predicted effects of an elastic penalty associated with strain of the LC about the bacteria when escape occurs into the nematic phase. We also observe boojums (surface topological defects) present at the interfaces of droplets of nematic LC (tactoids) to play a central role in mediating the escape of motile bacteria from the LC interface. Whereas the bacteria escape the interface of nematic droplets via a mechanism that involved nematic director-guided motion through one of the two boojums, for isotropic droplets in a continuous nematic phase, the elasticity of the LC generally prevented single bacteria from escaping. Instead, assemblies of bacteria piled up at boojums and escape occurred through a cooperative, multicellular phenomenon. Overall, our studies show that the dynamical behaviors of motile bacteria at anisotropic LC interfaces can be understood within a conceptual framework that reflects the interplay of LC elasticity, surface-induced order, and topological defects.

We describe the controlled transport and delivery of non-motile eukaryotic cells and polymer microparticles by swimming bacteria suspended in nematic liquid crystals. The bacteria push reversibly attached cargo in a stable, unidirectional path (or along a complex patterned director field) over exceptionally long distances. Numerical simulations and analytical predictions for swimming speeds provide a mechanistic insight into the hydrodynamics of the system. This study lays the foundation for using cargo-carrying bacteria in engineering applications and for understanding interspecies interactions in polymicrobial communities.