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Cells operate in part by compartmentalizing chemical reactions. For example, recent work has shown that chromatin, the material that contains the cell's genome, can auto-regulate its structure by utilizing reaction products (proteins, RNA)…

Soft Condensed Matter · Physics 2024-10-30 Sam Wilken , Gabrielle R. Abraham , Omar A. Saleh

Phase separation of various materials has been studied for one and a half centuries. In the last two decades, phase separation of proteins and nucleic acids has received enormous attention, due its relevance to cellular functions. However,…

Soft Condensed Matter · Physics 2026-05-28 Huan-Xiang Zhou

We present a generic mechanism by which reproducing microorganisms, with a diffusivity that depends on the local population density, can form stable patterns. It is known that a decrease of swimming speed with density can promote separation…

Populations and Evolution · Quantitative Biology 2010-07-13 M. E. Cates , D. Marenduzzo , I. Pagonabarraga , J. Tailleur

Biomolecular phase separation is typically attributed to the polymer physics of long, disordered chains. However, the underlying chemical grammar, i.e. the specific interactions between protein and RNA building blocks, remains poorly…

Biological Physics · Physics 2026-04-21 Greta Grassmann , Giancarlo Ruocco , Mattia Miotto

Living systems operate out of equilibrium, continuously consuming energy to sustain organised, functional states. Their emergent behaviour usually relies on a set of interconnected chemical reaction networks (CRNs) driven by external fluxes…

Statistical Mechanics · Physics 2026-02-03 Shiling Liang , Paolo De Los Rios , Daniel Maria Busiello

Biomolecular condensates in cells are often rich in catalytically-active enzymes. This is particularly true in the case of the large enzymatic complexes known as metabolons, which contain different enzymes that participate in the same…

Subcellular Processes · Quantitative Biology 2021-10-20 Vincent Ouazan-Reboul , Jaime Agudo-Canalejo , Ramin Golestanian

We address a fundamental question: under which conditions do the dynamics and thermodynamics of open chemical reaction networks (CRNs), grounded on the notion of idealized chemostats that exchange selected species, emerge from underlying…

Statistical Mechanics · Physics 2026-04-17 Benedikt Remlein , Massimiliano Esposito , Francesco Avanzini

Living cells exhibit a complex organization comprising numerous compartments, among which are RNA- and protein-rich membraneless, liquid-like organelles known as biomolecular condensates. Energy-consuming processes regulate their formation…

Evidence accumulated over the past decade provides support for liquid-liquid phase separation as the mechanism underlying the formation of biomolecular condensates, which include not only membraneless organelles such as nucleoli and RNA…

Biomolecules · Quantitative Biology 2019-09-12 Ivan Peran , Tanja Mittag

Biomolecular condensates form by phase separation of biological polymers and have important functions in the cell $-$ functions that are inherently connected to their physical properties. A remarkable aspect of such condensates is that…

In active systems, whose constituents have non-equilibrium dynamics at local level, fluid-fluid phase separation is widely observed. Examples include the formation of membraneless organelles within cells; the clustering of self-propelled…

Soft Condensed Matter · Physics 2025-07-11 M. E. Cates , C. Nardini

We identify the thermodynamic conditions necessary to observe indefinite growth in homogeneous open chemical reaction networks (CRNs) satisfying mass action kinetics. We also characterize the thermodynamic efficiency of growth by…

Molecular Networks · Quantitative Biology 2024-12-05 Shesha Gopal Marehalli Srinivas , Francesco Avanzini , Massimiliano Esposito

The liquid-liquid phase separation of biomolecules is an important process for intracellular organization. Biomolecular sequence combinatorics leads to a large variety of proteins and nucleic acids which can interact to form a diversity of…

Soft Condensed Matter · Physics 2025-08-27 Aria S. Chaderjian , Sam Wilken , Omar A. Saleh

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

Microphase separation is common in active biological systems as exemplified by the separation of RNA and DNA-rich phases in the cell nucleus driven by the transcriptional activity of polymerase enzymes acting similarly to amphiphiles in a…

Soft Condensed Matter · Physics 2025-07-22 Rakesh Chatterjee , Hui-Shun Kuan , Frank Julicher , Vasily Zaburdaev

The separation of substances into different phases is ubiquitous in nature and important scientifically and technologically. This phenomenon may become drastically different if the species involved, whether molecules or supramolecular…

The dynamics of a chemical reaction network (CRN) is often modelled under the assumption of mass action kinetics by a system of ordinary differential equations (ODEs) with polynomial right-hand sides that describe the time evolution of…

Dynamical Systems · Mathematics 2023-01-05 Radek Erban , Hye-Won Kang

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

Coupling between chemical fuel consumption and phase separation can lead to condensation at a nonequilibrium steady state, resulting in phase behaviors that are not described by equilibrium thermodynamics. Theoretical models of such…

Soft Condensed Matter · Physics 2025-05-23 Yongick Cho , William M. Jacobs

Chemical reaction networks (CRNs) are foundational models for describing complex biochemical processes. We study noncompetitive CRNs, a class of networks whose static states are rate-independent, and that can implement ReLU neural networks.…

Molecular Networks · Quantitative Biology 2025-12-22 Louis Faul , Xavier Richard , Mary Betrisey , Christian Mazza