Lipid Membrane Domains Control Actin Network Viscoelasticity.

The mammalian cell membrane is embedded with biomolecular condensates of protein and lipid clusters, which interact with an underlying viscoelastic cytoskeleton network to organize the cell surface and mechanically interact with the extracellular environment. However, the mechanical and thermodynamic interplay between the viscoelastic network and liquid-liquid phase separation of 2-dimensional (2D) lipid condensates remains poorly understood. Here, we engineer materials composed of 2D lipid membrane condensates embedded within a thin viscoelastic actin network.

Model predictive control of non-interacting active Brownian particles.

Active matter systems are strongly driven to assume non-equilibrium distributions owing to their self-propulsion, , flocking and clustering. Controlling the active matter systems' spatiotemporal distributions offers exciting applications such as directed assembly, programmable materials, and microfluidic actuation. However, these applications involve environments with coupled dynamics and complex tasks, making intuitive control strategies insufficient.

Dynamic swarms regulate the morphology and distribution of soft membrane domains.

We study the dynamic structure of lipid domain inclusions embedded within a phase-separated reconstituted lipid bilayer in contact with a swarming flow of gliding filamentous actin. Passive circular domains transition into highly deformed morphologies that continuously elongate, rotate, and pinch off into smaller fragments, leading to a dynamic steady state with ≈23× speedup in the relaxation of the intermediate scattering function compared with passive membrane domains driven by purely thermal forces. To corroborate experimental results, we develop a phase-field model of the lipid domains with two-way coupling to the Toner-Tu equations.

Dynamic surfactants drive anisotropic colloidal assembly.

Colloidal building blocks with re-configurable shapes and dynamic interactions can exhibit unusual self-assembly behaviors and pathways. In this work, we consider the phase behavior of colloids coated with surface-mobile polymer brushes that behave as "dynamic surfactants." Unlike traditional polymer-grafted colloids, we show that colloids coated with dynamic surfactants can acquire anisotropic macroscopic assemblies, even for spherical colloids with isotropic attractive interactions.

Surface Topography Induces and Orients Nematic Swarms of Active Filaments: Considerations for Lab-On-A-Chip Devices.

Surface-bound molecular motors can drive the collective motion of cytoskeletal filaments in the form of nematic bands and polar flocks in reconstituted gliding assays. Although these "swarming transitions" are an emergent property of active filament collisions, they can be controlled and guided by tuning the surface chemistry or topography of the substrate. To date, the impact of surface topography on collective motion in active nematics is only partially understood, with most experimental studies focusing on the escape of a single filament from etched channels.

Soft confinement of self-propelled rods: simulation and theory.

We present an analytical framework for evolving the dynamics of active rods under any periodic external potential, including confining channels and arrays of harmonic traps. As a proof of concept, we analyze the structure and dispersion of self-propelled rods under a soft, periodic one-dimensional (1D) confinement potential and under a two-dimensional (2D) periodic radial harmonic trap. While passive rods and polymers nematically order under 1D confinement, their diffusive transport along the director is limited by thermal diffusion.

Colloidal transport phenomena in dynamic, pulsating porous materials.

We study the transport phenomena of colloidal particles embedded within a moving array of obstacles that mimics a dynamic, time-varying porous material. While colloidal transport in an array of stationary obstacles ("passive" porous media) has been well studied, we lack the fundamental understanding of colloidal diffusion in a nonequilibrium porous environment. We combine Taylor dispersion theory, Brownian dynamics simulations, and optical tweezer experiments to study the transport of tracer colloidal particles in an oscillating lattice of obstacles.

Nonequilibrium interactions between multi-scale colloids regulate the suspension microstructure and rheology.

Understanding nonequilibrium interactions of multi-component colloidal suspensions is critical for many dynamical settings such as self-assembly and material processing. A key question is how the nonequilibrium distributions of individual components influence the effective interparticle interactions and flow behavior. In this work, we develop a first-principle framework to study a bidisperse suspension of colloids and depletants using a Smoluchowski equation and corroborated by Brownian dynamics (BD) simulations.

Active Surface Flows Accelerate the Coarsening of Lipid Membrane Domains.

Phase separation of multicomponent lipid membranes is characterized by the nucleation and coarsening of circular membrane domains that grow slowly in time as ∼t^{1/3}, following classical theories of coalescence and Ostwald ripening. In this Letter, we study the coarsening kinetics of phase-separating lipid membranes subjected to nonequilibrium forces and flows transmitted by motor-driven gliding actin filaments. We experimentally observe that the activity-induced surface flows trigger rapid coarsening of noncircular membrane domains that grow as ∼t^{2/3}, a 2x acceleration in the growth exponent compared to passive coalescence and Ostwald ripening.

Dynamic interfaces for contact-time control of colloidal interactions.

Understanding pairwise interactions between colloidal particles out of equilibrium has a profound impact on dynamical processes such as colloidal self assembly. However, traditional colloidal interactions are effectively quasi-static on colloidal timescales and cannot be modulated out of equilibrium. A mechanism to dynamically tune the interactions during colloidal contacts can provide new avenues for self assembly and material design.

Bio-enabled Engineering of Multifunctional "Living" Surfaces.

Through the magic of "active matter"─matter that converts chemical energy into mechanical work to drive emergent properties─biology solves a myriad of seemingly enormous physical challenges. Using active matter surfaces, for example, our lungs clear an astronomically large number of particulate contaminants that accompany each of the 10,000 L of air we respire per day, thus ensuring that the lungs' gas exchange surfaces remain functional. In this Perspective, we describe our efforts to engineer artificial active surfaces that mimic active matter surfaces in biology.

Antibody binding reports spatial heterogeneities in cell membrane organization.

The spatial organization of cell membrane glycoproteins and glycolipids is critical for mediating the binding of ligands, receptors, and macromolecules on the plasma membrane. However, we currently do not have the methods to quantify the spatial heterogeneities of macromolecular crowding on live cell surfaces. In this work, we combine experiment and simulation to report crowding heterogeneities on reconstituted membranes and live cell membranes with nanometer spatial resolution.

Engineered molecular sensors for quantifying cell surface crowding.

Cells mediate interactions with the extracellular environment through a crowded assembly of transmembrane proteins, glycoproteins and glycolipids on their plasma membrane. The extent to which surface crowding modulates the biophysical interactions of ligands, receptors, and other macromolecules is poorly understood due to the lack of methods to quantify surface crowding on native cell membranes. In this work, we demonstrate that physical crowding on reconstituted membranes and live cell surfaces attenuates the effective binding affinity of macromolecules such as IgG antibodies in a surface crowding-dependent manner.

Enhanced dispersion in an oscillating array of harmonic traps.

Experiment, theory, and simulation are employed to understand the dispersion of colloidal particles in a periodic array of oscillating harmonic traps generated by optical tweezers. In the presence of trap oscillation, a nonmonotonic and anisotropic dispersion is observed. Surprisingly, the stiffest traps produce the largest dispersion at a critical frequency, and the particles diffuse significantly faster in the direction of oscillation than those undergoing passive Stokes-Einstein-Sutherland diffusion.

Boundary design regulates the diffusion of active matter in heterogeneous environments.

Physical boundaries play a key role in governing the overall transport properties of nearby self-propelled particles. In this work, we develop dispersion theories and conduct Brownian dynamics simulations to predict the coupling between surface accumulation and effective diffusivity of active particles in boundary-rich media. We focus on three models that are well-understood for passive systems: particle transport in (i) an array of fixed volume-excluding obstacles; (ii) a pore with spatially heterogeneous width; and (iii) a tortuous path with kinks and corners.

Molecular height measurement by cell surface optical profilometry (CSOP).

The physical dimensions of proteins and glycans on cell surfaces can critically affect cell function, for example, by preventing close contact between cells and limiting receptor accessibility. However, high-resolution measurements of molecular heights on native cell membranes have been difficult to obtain. Here we present a simple and rapid method that achieves nanometer height resolution by localizing fluorophores at the tip and base of cell surface molecules and determining their separation by radially averaging across many molecules.

Active Contact Forces Drive Nonequilibrium Fluctuations in Membrane Vesicles.

We analyze the nonequilibrium shape fluctuations of giant unilamellar vesicles encapsulating motile bacteria. Owing to bacteria-membrane collisions, we experimentally observe a significant increase in the magnitude of membrane fluctuations at low wave numbers, compared to the well-known thermal fluctuation spectrum. We interrogate these results by numerically simulating membrane height fluctuations via a modified Langevin equation, which includes bacteria-membrane contact forces.

Acoustic trapping of active matter.

Confinement of living microorganisms and self-propelled particles by an external trap provides a means of analysing the motion and behaviour of active systems. Developing a tweezer with a trapping radius large compared with the swimmers' size and run length has been an experimental challenge, as standard optical traps are too weak. Here we report the novel use of an acoustic tweezer to confine self-propelled particles in two dimensions over distances large compared with the swimmers' run length.

A theory for the phase behavior of mixtures of active particles.

Systems at equilibrium like molecular or colloidal suspensions have a well-defined thermal energy kBT that quantifies the particles' kinetic energy and gauges how "hot" or "cold" the system is. For systems far from equilibrium, such as active matter, it is unclear whether the concept of a "temperature" exists and whether self-propelled entities are capable of thermally equilibrating like passive Brownian suspensions. Here we develop a simple mechanical theory to study the phase behavior and "temperature" of a mixture of self-propelled particles.

Swim stress, motion, and deformation of active matter: effect of an external field.

We analyze the stress, dispersion, and average swimming speed of self-propelled particles subjected to an external field that affects their orientation and speed. The swimming trajectory is governed by a competition between the orienting influence (i.e.

In vivo corneal oxygen uptake during soft-contact-lens wear.

Purpose: We develop a new method to compute in situ corneal oxygen uptake during soft-contact-lens (SCL) wear using a micro-polarographic Clark electrode.

Methods: After steady SCL wear and subsequent removal, a membrane-covered polarographic microelectrode is immediately placed onto the cornea. The resulting polarographic signal is related to the steady-state corneal oxygen uptake rate during soft-contact-lens wear.

In vivo oxygen uptake into the human cornea.

Purpose: We provide a new procedure to quantify in situ corneal oxygen uptake using the micropolarographic Clark electrode.

Methods: Traditionally, upon placing a membrane-covered Clark microelectrode onto a human cornea, the resulting polarographic signal is interpreted as the oxygen partial pressure at the anterior corneal surface. However, the Clark electrode operates at a limiting current.

A quasi-2-dimensional model for respiration of the cornea with soft contact lens wear.

Purpose: Because neither the human cornea nor a soft contact lens (SCL) is of constant thickness, corneal oxygenation varies locally. To quantify the importance of cornea/SCL thickness variations on oxygen demand, we develop a quasi-2-dimensional (2D) respiration model that accounts for aerobic and anaerobic metabolism and bicarbonate buffering.

Methods: Because metabolism is critical to oxygen demand, we extend the 1-dimensional (1D), 6-layer oxygen metabolic model of Chhabra et al.