Far-from-equilibrium processes constantly dissipate energy while converting a free-energy source to another form of energy. Living systems, for example, rely on an orchestra of molecular motors that consume chemical fuel to produce mechanical work. In this talk, I will describe two features of life, namely, time-irreversibility, and nonequilibrium self-assembly. Time irreversibility is the hallmark of nonequilibrium dissipative processes. Detecting dissipation is essential for our basic understanding of the underlying physical mechanism, however, it remains a challenge in the absence of observable directed motion, flows, or fluxes. Additional difficulty arises in complex systems where many internal degrees of freedom are inaccessible to an external observer. I will introduce a novel approach to detect time irreversibility and estimate the entropy production from time-series measurements, even in the absence of observable currents. This method can be implemented in scenarios where only partial information is available and thus provides a new tool for studying nonequilibrium phenomena. Further, I will explore the added benefits achieved by nonequilibrium driving for self-assembly, identify distinctive collective phenomena that emerge in a nonequilibrium self-assembly setting, and demonstrate the interplay between the assembly speed, kinetic stability, and relative population of dynamical attractors.

The motion of passive or active agents in soft materials generates long ranged deformation fields with signatures informed by hydrodynamics and the properties of the soft matter host. These signatures are even more complex when the soft matter host itself is an active material. Measurement of these fields reveals mechanics of the soft materials and hydrodynamics central to understanding self-organization. In this talk, I first introduce a new method based on correlated displacement velocimetry, and use the method to measure flow fields around particles trapped at the interface between immiscible fluids. These flow fields, decomposed into interfacial hydrodynamic multipoles, including force monopole and dipole flows, provide key insights essential to understanding the interface’s mechanical response. I then extend this method to various actomyosin systems to measure local strain fields around myosin molecular motors. I show how active stresses propagate in 2d liquid crystalline structures and in disordered networks that are formed by the actin filaments. In particular, the response functions of contractile and stable gels are characterized. Through similar analysis, I also measure the retrograde flow fields of stress fibers in single cells to understand subcellular mechanochemical systems.

Living cells contain millions of enzymes and proteins, which carry out multiple reactions simultaneously. To optimize these processes, cells compartmentalize reactions in membraneless liquid condensates. Certain features of cellular condensates can be explained by principles of liquid-liquid phase separation studied in material science. However, biological condensates exist in the inherently out of equilibrium environment of a living cell, being driven by force-generating microscopic processes. These cellular conditions are fundamentally different than the equilibrium conditions of liquid-liquid phase separation studied in materials science and physics. How condensates function in the active riotous environment of a cell is essential for understanding of cellular functions, as well as to the onset of neurodegenerative diseases. Currently, we lack model systems that enable rigorous studies of these processes. Living cells are too complex for quantitative analysis, while reconstituted equilibrium condensates fail to capture the non-equilibrium environment of biological cells. To bridge this gap, we reconstituted a DNA based membraneless condensates in an active environment that mimics the conditions of a living cell. We combine condensates with a reconstituted network of cytoskeletal filaments and molecular motors, and study how the mechanical interactions change the phase behavior and dynamics of membraneless structures. Studying these composite materials elucidates the fundamental physics rules that govern the behavior of liquid-liquid phase separation away from equilibrium while providing insight into the mechanism of condensate phase separation in cellular environments.