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Phase separating systems that are maintained away from thermodynamic equilibrium via molecular processes represent a class of active systems, which we call active emulsions. These systems are driven by external energy input for example…

Soft Condensed Matter · Physics 2019-06-07 Christoph A. Weber , David Zwicker , Frank Jülicher , Chiu Fan Lee

These lecture notes describe a basic theory of chemically active droplets, which are droplets kept away from equilibrium by driven chemical reactions. The notes assume a basic familiarity with equilibrium thermodynamics of phase separation,…

Soft Condensed Matter · Physics 2024-07-16 David Zwicker

Biomolecular condensates are small droplets forming spontaneously in biological cells via phase separation. They play a role in many cellular processes, but it is unclear how cells control them. Cellular regulation often relies on…

Soft Condensed Matter · Physics 2024-03-15 Jan Kirschbaum , David Zwicker

Droplet formation has emerged as an essential concept for the spatiotemporal organisation of biomolecules in cells. However, classical descriptions of droplet dynamics based on passive liquid-liquid phase separation cannot capture the…

Biological Physics · Physics 2025-11-14 David Zwicker , Oliver W. Paulin , Cathelijne ter Burg

Phase-separated liquid condensates can spatially organize and thereby regulate chemical processes. However, the physicochemical mechanisms underlying such regulation remain elusive as the intramolecular interactions responsible for phase…

Soft Condensed Matter · Physics 2024-03-11 Sudarshana Laha , Jonathan Bauermann , Frank Jülicher , Thomas C. T. Michaels , Christoph A. Weber

Phase separation and transitions among different molecular states are ubiquitous in living cells. Such transitions can be governed by local equilibrium thermodynamics or by active processes controlled by biological fuel. It remains largely…

Biological Physics · Physics 2021-11-24 G. Bartolucci , O. Adame-Arana , X. Zhao , C. A. Weber

Droplets are essential for spatially controlling biomolecules in cells. To work properly, cells need to control the emergence and morphology of droplets. On the one hand, driven chemical reactions can affect droplets profoundly. For…

Soft Condensed Matter · Physics 2024-03-14 Noah Ziethen , David Zwicker

It has been proposed that during the early steps in the origin of life, small droplets could have formed via the segregation of molecules from complex mixtures by phase separation. These droplets could have provided chemical reaction…

Biological Physics · Physics 2017-04-26 David Zwicker , Rabea Seyboldt , Christoph A. Weber , Anthony A. Hyman , Frank Jülicher

Chemically active droplets provide simple models for cell-like systems that can grow and divide. Such active droplet systems are driven away from thermodynamic equilibrium and turn over chemically, which corresponds to a simple metabolism.…

Biological Physics · Physics 2022-09-28 Jonathan Bauermann , Christoph A. Weber , Frank Jülicher

Phase separation, crucial for spatially segregating biomolecules in cells, is well-understood in the simple case of a few components with pairwise interactions. Yet, biological cells challenge the simple picture in at least two ways: First,…

Soft Condensed Matter · Physics 2024-03-12 Chengjie Luo , Yicheng Qiang , David Zwicker

We study the hydrodynamics and shape changes of chemically active droplets. In non-spherical droplets, surface tension generates hydrodynamic flows that drive liquid droplets into a spherical shape. Here we show that spherical droplets that…

Biological Physics · Physics 2018-11-12 Rabea Seyboldt , Frank Jülicher

Liquid-liquid phase separation is important across biology, physics, and materials science. Although usually studied at equilibrium, active components - such as motor proteins, enzymes, and synthetic microswimmers - are increasingly…

Soft Condensed Matter · Physics 2026-01-06 Kan Chang , Yulin Li , Ming Yuan , Masaki Sano , Zhihong You , H. P. Zhang

We use a continuum model to examine the effect of activity on a phase separating mixture of an extensile active nematic and a passive fluid. We highlight the distinct role of previously considered interfacial active stresses and bulk active…

Soft Condensed Matter · Physics 2022-12-26 Fernando Caballero , M. Cristina Marchetti

The formation of small droplets and bubbles in turbulent flows is a crucial process in geophysics and engineering, whose underlying physical mechanism remains a puzzle. In this letter, we address this problem by means of high-resolution…

Chemical activity is known to affect phase coexistence and coarsening in liquid mixtures, most commonly through reaction-induced changes of intermolecular interactions. Here, we analyze a scenario in which chemical reactions regulate…

Soft Condensed Matter · Physics 2026-03-19 Florian Raßhofer , Erwin Frey

Phase separation drives the formation of biomolecular condensates in cells, which comprise many components and sometimes possess multiple phases. The equilibrium physics of phase separation is well understood, but many components in…

Soft Condensed Matter · Physics 2025-08-14 Yicheng Qiang , Chengjie Luo , David Zwicker

Chemical processes in closed systems are poorly controllable since they always relax to equilibrium. Living systems avoid this fate and give rise to a much richer diversity of phenomena by operating under nonequilibrium conditions. Recent…

Chemical Physics · Physics 2019-09-11 Emanuele Penocchio , Riccardo Rao , Massimiliano Esposito

Liquid-liquid phase separation is key to understanding aqueous two-phase systems (ATPS) arising throughout cell biology, medical science, and the pharmaceutical industry. Controlling the detailed morphology of phase-separating compound…

Soft Condensed Matter · Physics 2023-10-27 Eric W. Hester , Sean P. Carney , Vishwesh Shah , Alyssa Arnheim , Bena Patel , Dino Di Carlo , Andrea L. Bertozzi

Droplets form a cornerstone of the spatiotemporal organization of biomolecules in cells. These droplets are controlled using physical processes like chemical reactions and imposed gradients, which are costly to simulate using traditional…

Soft Condensed Matter · Physics 2022-06-23 Ajinkya Kulkarni , Estefania Vidal-Henriquez , David Zwicker

Biomolecular condensates play a central role in the spatial organization of living matter. Their formation is now well understood as a form of liquid-liquid phase separation that occurs very far from equilibrium. For instance, they can be…

Soft Condensed Matter · Physics 2024-10-14 Roxanne Berthin , Jacques Fries , Marie Jardat , Vincent Dahirel , Pierre Illien
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