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In a companion paper, we put forth a thermodynamic model for complex formation via a chemical reaction involving multiple macromolecular species, which may subsequently undergo liquid-liquid phase separation and a further transition into a…

Biological Physics · Physics 2024-07-26 Ruoyao Zhang , Sheng Mao , Mikko P. Haataja

Biological cells exhibit a hierarchical spatial organization, where various compartments harbor condensates that form by phase separation. Cells can control the emergence of these condensates by affecting compartment size, the amount of the…

Biological Physics · Physics 2025-04-03 Riccardo Rossetto , Gerrit Wellecke , David Zwicker

Phase transitions, such as the freezing of water and the magnetisation of a ferromagnet upon lowering the ambient temperature, are familiar physical phenomena. Interestingly, such a collective change of behaviour at a phase transition is…

Biological Physics · Physics 2018-10-26 Chiu Fan Lee , Jean David Wurtz

Biomolecular condensates are essential for cellular organization and result from phase separation in systems far from thermodynamic equilibrium. Among various models, chemically active droplets play a significant role, consisting of…

Soft Condensed Matter · Physics 2025-05-19 Jacques Fries , Roxanne Berthin , Chengjie Luo , Marie Jardat , David Zwicker , Vincent Dahirel , Pierre Illien

Proliferation and motility are ubiquitous drivers of activity in biological systems. Here, we study a dense binary mixture of motile and proliferating particles with exclusively repulsive interactions, where homeostasis in the proliferating…

Soft Condensed Matter · Physics 2025-06-11 Lukas Hupe , Joanna M. Materska , David Zwicker , Ramin Golestanian , Bartlomiej Waclaw , Philip Bittihn

Phase separation in passive systems leads to uncontrolled droplet growth, limiting structural control in soft materials and cells. We identify a generic mechanism to arrest coarsening based on chemical interconversion between molecular…

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

If a given behavior of a multi-agent system restricts the phase variable to a invariant manifold, then we define a phase transition as change of physical characteristics such as speed, coordination, and structure. We define such a phase…

Dynamical Systems · Mathematics 2017-07-21 Kelum Gajamannage , Erik M. Bollt

Differences in activities in colloidal particles are sufficient to drive phase separation between active and passive (or less active) particles, even if they have only excluded volume interactions. In this paper, we study the phase…

Soft Condensed Matter · Physics 2020-05-27 Efe Ilker , Jean-François Joanny

Mixtures of several macromolecular species can lead to the formation of higher-order structures that often display non-ideal mixing behavior. In this work, we propose a minimal model of a quaternary system which considers the formation of a…

Soft Condensed Matter · Physics 2022-12-14 Ruoyao Zhang , Sheng Mao , Mikko Haataja

Unlike fluids at thermal equilibrium, biomolecular mixtures in living systems can sustain nonequilibrium steady states, in which active processes modify the conformational states of the constituent molecules. Despite qualitative…

Soft Condensed Matter · Physics 2023-03-31 Yongick Cho , William M. Jacobs

We study the thermodynamics of binary mixtures with the volume fraction of the minority component less than the amount required to form a flat interface and show that the surface tension dominated equilibrium phase of the mixture forms a…

Soft Condensed Matter · Physics 2023-03-07 Subhadip Biswas , Biswaroop Mukherjee , Buddhapriya Chakrabarti

Phase separation of two phase separating solutes in a common solvent can result in mesoscale (micron-sized) droplets with complex topologies of the domains of each solute within each droplet. Such topologies have been observed in-vitro in…

Soft Condensed Matter · Physics 2025-07-31 Amit Kumar , Gary H. Karpen , Samuel A. Safran

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

Liquids with quasi - chemical bonding between molecules are described in terms of vertex model. It is shown that this bonding results in liquid - liquid phase transition, which occurs between phases with different mean density of…

Statistical Mechanics · Physics 2007-05-23 L. Son , R. Ryltcev

Demixing of binary liquids is a ubiquitous transition, which is explained using a well-established thermodynamic formalism that requires equality of intensive thermodynamics parameters across the phase boundaries. Demixing transitions also…

During overflow metabolism, cells excrete glycolytic byproducts when growing under aerobic conditions in a seemingly wasteful fashion. While potentially advantageous for microbes with finite oxidative capacity, its role in higher organisms…

Living and engineered systems rely on the stable coexistence of two interspersed liquid phases. Yet surface tension drives their complete separation. Here we show that stable droplets of uniform and tuneable size can be produced through…

We study a mechanism for reliable switching in biomolecular signal-transduction cascades. Steady bistable states are created by system-size cooperative effects in populations of proteins, in spite of the fact that the phosphorylation-state…

Subcellular Processes · Quantitative Biology 2015-05-30 Eric Smith , Supriya Krishnamurthy , Walter Fontana , David Krakauer

Biomolecular condensates play a crucial role in the spatial organization of living matter. These membrane-less organelles, resulting from liquid-liquid phase separation, operate far from thermodynamic equilibrium, with their size and…

Soft Condensed Matter · Physics 2025-05-12 Jacques Fries , Roxanne Berthin , Marie Jardat , Pierre Illien , Vincent Dahirel

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