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Microtubule-kinesin active fluids consume ATP to generate internal active stresses, driving spontaneous and complex flows. While numerous studies have explored the fluid's autonomous behavior, its response to external mechanical forces remains less understood. This study explores how moving boundaries affect the flow dynamics of this active fluid when confined in a thin cuboidal cavity. Our experiments demonstrate a transition from chaotic, disordered vortices to a single, coherent system-wide vortex as boundary speed increases, resembling the behavior of passive fluids like water. Furthermore, our confocal microscopy revealed that boundary motion altered the microtubule network structure near the moving boundary. In the absence of motion, the network exhibited a disordered, isotropic configuration. However, as the boundary moved, microtubule bundles aligned with the shear flow, resulting in a thicker, tilted nematic layer extending over a greater distance from the moving boundary. These findings highlight the competing influences of external shear stress and internal active stress on both flow kinematics and microtubule network structure. This work provides insight into the mechanical properties of active fluids, with potential applications in areas such as adaptive biomaterials that respond to mechanical stimuli in biological environments. 

Microtubule-kinesin active fluids consume ATP to generate internal active stresses, driving spontaneous and complex flows. While numerous studies have explored the fluid's autonomous behavior, its response to external mechanical forces remains less understood. This study explores how moving boundaries affect the flow dynamics of this active fluid when confined in a thin cuboidal cavity. Our experiments demonstrate a transition from chaotic, disordered vortices to a single, coherent system-wide vortex as boundary speed increases, resembling the behavior of passive fluids like water. Furthermore, our confocal microscopy revealed that boundary motion altered the microtubule network structure near the moving boundary. In the absence of motion, the network exhibited a disordered, isotropic configuration. However, as the boundary moved, microtubule bundles aligned with the shear flow, resulting in a thicker, tilted nematic layer extending over a greater distance from the moving boundary. These findings highlight the competing influences of external shear stress and internal active stress on both flow kinematics and microtubule network structure. This work provides insight into the mechanical properties of active fluids, with potential applications in areas such as adaptive biomaterials that respond to mechanical stimuli in biological environments. *We acknowledge support from the National Science Foundation (NSF-CBET-2045621). This research is performed with computational resources supported by the Academic & Research Computing Group at Worcester Polytechnic Institute. We acknowledge the Brandeis Materials Research Science and Engineering Center (NSF-MRSEC-DMR-2011846) for use of the Biological Materials Facility. 

Active fluids have potential applications in micromixing, but little is known about the mixing kinematics of such systems with spatiotemporally-varying activity. To investigate, UV-activated caged ATP was used to activate controlled regions of microtubule-kinesin active fluid inducing a propagating active-passive interface. The mixing process of the system from non-uniform to uniform activity as the interface advanced was observed with fluorescent tracers and molecular dyes. At low Péclet numbers (diffusive transport), the active-inactive interface progressed toward the inactive area in a diffusion-like manner and at high Péclet numbers (convective transport), the active-inactive interface progressed in a superdiffusion-like manner. The results show mixing in non-uniform active fluid systems evolve from a complex interplay between the spatial distribution of ATP and its active transport. This active transport may be diffusion-like or superdiffusion-like depending on Péclet number and couples the spatiotemporal distribution of ATP and the subsequent localized active stresses of active fluid. Our work will inform the design of future microfluidic mixing applications and provide insight into intracellular mixing processes. *T.E.B., E.H.T., J.H.D., and K.-T.W. acknowledge support from the National Science Foundation (NSF-CBET-2045621). C.-C. C. was supported through the National Science and Technology Council (NSTC), Taiwan (111-2221-E-006-102-MY3). M.M.N. was supported by the U.S. Department of Energy, Office of Science, Office of Basic Energy Sciences (DE-SC0022280). 

Active fluids with spatiotemporally varying activity have potential applications to micromixing; however previously existing active fluids models are not prepared to account for spatiotemporally-varying active stresses. Our experimental work used UV-activated caged ATP to activate controlled regions of microtubule-kinesin active fluid inducing a propagating active-passive interface. Here, we recapitulate our experimental results with two models. The first model redistributes an initial ATP distribution by Fick's law and translates the ATP distribution into a velocity profile by Michaelis-Menton kinetics. This model reproduces our experimental measurements for the low-Péclet number limit within 10% error without fitting parameters. However, as the model is diffusion based, it fails to capture the convective based superdiffusive-like behaviour at high Péclet numbers. Our second model introduces a spatiotemporally varying ATP field to an existing nematohydrodynamic active fluid model and then couples the active stresses to local ATP concentrations. This model is successful in qualitatively capturing the superdiffusive-like progression of the active-inactive interface for high Peclet number (convective transport) experimental cases. Our results show that new model frameworks are necessary for capturing the behaviour of active fluid with spatiotemporally varying activity. *T.E.B., E.H.T., J.H.D., and K.-T.W. acknowledge support from the National Science Foundation (NSF-CBET-2045621). C.-C. C. was supported through the National Science and Technology Council (NSTC), Taiwan (111-2221-E-006-102-MY3). M.M.N. was supported by the U.S. Department of Energy, Office of Science, Office of Basic Energy Sciences (DE-SC0022280). 

Abstract Active fluids have applications in micromixing, but little is known about the mixing kinematics of systems with spatiotemporally-varying activity. To investigate, UV-activated caged ATP is used to activate controlled regions of microtubule-kinesin active fluid and the mixing process is observed with fluorescent tracers and molecular dyes. At low Péclet numbers (diffusive transport), the active-inactive interface progresses toward the inactive area in a diffusion-like manner that is described by a simple model combining diffusion with Michaelis-Menten kinetics. At high Péclet numbers (convective transport), the active-inactive interface progresses in a superdiffusion-like manner that is qualitatively captured by an active-fluid hydrodynamic model coupled to ATP transport. Results show that active fluid mixing involves complex coupling between distribution of active stress and active transport of ATP and reduces mixing time for suspended components with decreased impact of initial component distribution. This work will inform application of active fluids to promote micromixing in microfluidic devices. 

Fluid mixing is driven by the passive process of diffusion and the active process of stretching and folding, which homogenize the system's constituents. Conventionally, the active process is applied via external shearing machines such as a kitchen stand mixer. However, applying external shearing becomes more challenging in mesoscopic fluid systems due to the increasing difficulty of controlling the injection of energy on the micron scale. To overcome this challenge, we introduced microtubule-kinesin active fluid to power the active mixing process. To demonstrate its mixing capability, we created a multi-fluid system where active fluid is adjacent to an inactivated, passive fluid and allowed the active fluid to blend with the passive fluid until the system reaches a homogeneous state. We found that the mixing dynamics of such active-passive fluid mixing was dominated by the passive process of diffusion, until the activity of active fluid was tuned to be sufficiently high and the active processes of active fluid began to dominate the mixing process. Our work will stimulate the development of utilizing active fluid to accomplish mesoscale mixing tasks in multi-fluid systems at the micron scale. *We acknowledge support from the National Science Foundation (NSF-CBET-2045621). 

Abstract Active fluid droplets surrounded by oil can spontaneously develop circulatory flows. However, the dynamics of the surrounding oil and their influence on the active fluid remain poorly understood. To investigate interactions between the active fluid and the passive oil across their interface, kinesin-driven microtubule-based active fluid droplets were immersed in oil and compressed into a cylinder-like shape. The droplet geometry supported intradroplet circulatory flows, but the circulation was suppressed when the thickness of the oil layer surrounding the droplet decreased. Experiments with tracers and network structure analyses and continuum models based on the dynamics of self-elongating rods demonstrated that the flow transition resulted from flow coupling across the interface between active fluid and oil, with a millimeter–scale coupling length. In addition, two novel millifluidic devices were developed that could trigger and suppress intradroplet circulatory flows in real time: one by changing the thickness of the surrounding oil layer and the other by locally deforming the droplet. This work highlights the role of interfacial dynamics in the active fluid droplet system and shows that circulatory flows within droplets can be affected by millimeter–scale flow coupling across the interface between the active fluid and the oil. 

We study the influence of solid boundaries on dynamics and structure of kinesin-driven microtubule active fluids as the height of the container, H, increases from hundreds of micrometers to several millimeters. By three-dimensional tracking of passive tracers dispersed in the active fluid, we observe that the activity level, characterized by velocity fluctuations, increases as system size increases and retains a small-scale isotropy. Concomitantly, as the confinement level decreases, the velocity-velocity temporal correlation develops a strong positive correlation at longer times, suggesting the establishment of a "memory". We estimate the characteristic size of the flow structures from the spatial correlation function and find that, as the confinement becomes weaker, the correlation length, l_c, saturates at approximately 400 microns. This saturation suggests an intrinsic length scale which, along with the small-scale isotropy, demonstrates the multiscale nature of this kinesin-driven bundled microtubule active system. 

Boundary conditions influence the outcome of fluid dynamics in conventional passive fluid systems. Such an influence also extends to active fluid systems where fluid can flow by itself without an external driving force. For example, an active fluid that is confined in a thin cylinder can self-organize into a circulation along the central axis of the cylinder but thinning the cylinder to a disk-like geometry suppresses the formation of circulation. These phenomena demonstrated the role of confinement geometry on flow patterns of active fluid. Here, we demonstrate two flow patterns induced by confinement. First, we will show that active fluid can convect within a trapezoidal confinement. Such convection was in a temperature-uniform system, in contrast to Rayleigh-Bénard convection which is induced by a temperature gradient. This result suggested the feasibility of developing convection in a temperature-homogeneous system. Second, we demonstrate a confinement-induced stationary vortex near a corner of confinement whose corner angle is below a critical value. This is similar to conventional Moffatt eddies, except the fluid is internally driven. Our work paves the path to controlling self-organization of active fluid using confinement. 

Microtubule-based active matter provides insight into the self-organization of motile interacting constituents. We describe several formulations of microtubule-based 3D active isotropic fluids. Dynamics of these fluids is powered by three types of kinesin motors: a processive motor, a non-processive motor, and a motor which is permanently linked to a microtubule backbone. Another modification uses a specific microtubule crosslinker to induce bundle formation instead of a non-specific polymer depletant. In comparison to the already established system, each formulation exhibits distinct properties. These developments reveal the temporal stability of microtubule-based active fluids while extending their reach and the applicability.