Collective motion of self-propelled organisms or synthetic particles, often termed “active fluid,” has attracted enormous attention in the broad scientific community because of its fundamentally nonequilibrium nature. Energy input and interactions among the moving units and the medium lead to complex dynamics. Here, we introduce a class of active matter––living liquid crystals (LLCs)––that combines living swimming bacteria with a lyotropic liquid crystal. The physical properties of LLCs can be controlled by the amount of oxygen available to bacteria, by concentration of ingredients, or by temperature. Our studies reveal a wealth of intriguing dynamic phenomena, caused by the coupling between the activity-triggered flow and long-range orientational order of the medium. Among these are (i) nonlinear trajectories of bacterial motion guided by nonuniform director, (ii) local melting of the liquid crystal caused by the bacteria-produced shear flows, (iii) activity-triggered transition from a nonflowing uniform state into a flowing one-dimensional periodic pattern and its evolution into a turbulent array of topological defects, and (iv) birefringence-enabled visualization of microflow generated by the nanometers-thick bacterial flagella. Unlike their isotropic counterpart, the LLCs show collective dynamic effects at very low volume fraction of bacteria, on the order of 0.2%. Our work suggests an unorthodox design concept to control and manipulate the dynamic behavior of soft active matter and opens the door for potential biosensing and biomedical applications. Significance We propose a class of active matter, the living liquid crystal (LLC), representing motile rod-shaped bacteria placed in water-based nontoxic liquid crystal. Long-range orientational order of the liquid crystal and the swimming activity of bacteria demonstrate a strong coupling that dramatically alters individual and collective bacterial dynamics. For example, swimming bacteria perturb the orientational order of the liquid crystal or even cause its local melting, making the flagella motion optically visible. Second, self-organized textures emerge from the initial uniform LLC alignment with a characteristic length controlled by a balance between bacteria activity and anisotropic viscoelasticity of liquid crystal. Third, the local liquid crystal orientation controls the direction of motion of bacteria. LLC can lead to valuable biosensoring and biomedical applications.

Using a magnetic Frederiks transition technique, we measure the temperature and concentration dependences of splay $K_1$, twist $K_2$, and bend $K_3$ elastic constants for the lyotropic chromonic liquid crystal sunset yellow formed through noncovalent reversible aggregation of organic molecules in water. $K_1$ and $K_3$ are comparable to each other and are an order of magnitude higher than $K_2$. At higher concentrations and lower temperatures, $K_1$ and the ratios $K_1/K_3$ and $K_1/K_2$ increase, which is attributed to elongation of self-assembled lyotropic chromonic liquid crystal aggregates, a feature not found in conventional thermotropic and lyotropic liquid crystals formed by covalently bound units of a fixed length.

Using dynamic light scattering, we study orientational fluctuation modes in the nematic phase of a self-assembled lyotropic chromonic liquid crystal (LCLC) disodium cromoglycate and measure the Frank elastic moduli and viscosity coefficients. The elastic moduli of splay (K₁) and bend (K₃) are in the order of 10 pN while the twist modulus (K₂) is an order of magnitude smaller. The splay constant K₁ and the ratio K₁/K₃ both increase substantially as the temperature T decreases, which we attribute to the elongation of the chromonic aggregates at lower temperatures. The bend viscosity is comparable to that of thermotropic liquid crystals, while the splay and twist viscosities are several orders of magnitude larger. The temperature dependence of bend viscosity is weak. The splay and twist viscosities change exponentially with the temperature. In addition to the director modes, the fluctuation spectrum reveals an additional mode that is attributed to diffusion of structural defects in the column-like aggregates.

Flagellated bacteria such as Escherichia coli and Bacillus subtilis exhibit effective mechanisms for swimming in fluids and exploring the surrounding environment. In isotropic fluids such as water, the bacteria change swimming direction through the run-and-tumble process. Lyotropic chromonic liquid crystals (LCLCs) have been introduced recently as an anisotropic environment in which the direction of preferred orientation, the director, guides the bacterial trajectories. In this work, we describe the behavior of bacteria B. subtilis in a homeotropic LCLC geometry, in which the director is perpendicular to the bounding plates of a shallow cell. We demonstrate that the bacteria are capable of overcoming the stabilizing elastic forces of the LCLC and swim perpendicularly to the imposed director (and parallel to the bounding plates). The effect is explained by a finite surface anchoring of the director at the bacterial body; the role of surface anchoring is analyzed by numerical simulations of a rod realigning in an otherwise uniform director field. Shear flows produced by a swimming bacterium cause director distortions around its body, as evidenced both by experiments and numerical simulations. These distortions contribute to a repulsive force that keeps the swimming bacterium at a distance of a few micrometers away from the bounding plates. The homeotropic alignment of the director imposes two different scenarios of bacterial tumbling: one with an 180° reversal of the horizontal velocity and the other with the realignment of the bacterium by two consecutive 90° turns. In the second case, the angle between the bacterial body and the imposed director changes from 90° to 0° and then back to 90°; the new direction of swimming does not correlate with the previous swimming direction.

We use hydrogels to drive liquid crystals, achieving stretchable electrooptics. The device performance was maintained under a biaxial stretch of 1.5.