Frustrated run and tumble of swimming E-coli bacteria in nematic liquid crystals

In many situations bacteria move in complex environments, as for example in soils, oceans or the human gut-track microbiome. In these natural environments, carrier fluids such as mucus or reproductive fluids show complex structure associated with non-Newtonian rheology. Many fundamental questions concerning the the ability to navigate in such environments remain unsolved due to the inherent complexity of the natural surroundings. Recently, the interaction of swimming bacteria with nematic liquid crystals has attracted lot of attention. In these structured fluids, the kinetics of bacterial motion is constrained by the orientational molecular order of the liquid crystal (or director field) and novel spatio-temporal patterns arise from this orientational constraint, as well as from the interactions with topological defects. A question unaddressed so far is how bacteria are able to change swimming direction in such an environment. In this work, we study the swimming mechanism of a single bacterium, E. coli, constrained to move along the director field of a lyotropic chromonic liquid crystal (LCLC) that is confined to a planar cell. In such an environment, the spontaneous run and tumble motion of the bacterium gets frustrated: the elasticity of the liquid crystal prevents flagella from unbundling. Interestingly, in order to change direction, bacteria execute a reversal motion along the director field, driven by the relocation of a single flagellum to the other side of the bacterial body, coined as a frustrated tumble. We present a detailed experimental characterization of this phenomenon, exploiting exceptional spatial and temporal resolution of bacteria and flagella dynamics during swimming, obtained using a two color Lagrangian tracking technique. We suggest a possible mechanism behind the frustrated run and tumble motion, accounting for these observations.