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In thermodynamic equilibrium, the fluctuation-dissipation theorem links thermal fluctuations and dissipation. Biological systems, however, are driven out of equilibrium by internal processes that produce additional, active fluctuations.…

Soft Condensed Matter · Physics 2026-01-29 Ashot Matevosyan , Frank Jülicher , Ricard Alert

Biological assemblies such as chromosomes, membranes, and the cytoskeleton are driven out of equilibrium at the nanoscale by enzymatic activity and molecular motors. Similar non-equilibrium dynamics can be realized in synthetic systems,…

Biological Physics · Physics 2020-06-04 Federica Mura , Grzegorz Gradziuk , Chase P. Broedersz

Living systems break detailed balance at small scales, consuming energy and producing entropy in the environment in order to perform molecular and cellular functions. However, it remains unclear how broken detailed balance manifests at…

In systems with detailed balance, the stationary distribution and the equilibrium distribution are identical, creating a clear connection between energetic and entropic quantities. Many driven systems violate detailed balance and still pose…

Statistical Mechanics · Physics 2025-01-22 Markus Hofer , Jan Korbel , Rudolf Hanel , Stefan Thurner

We examine a biomolecular machine involving a driven, observable process coupled to a hidden process in a kinetically cooperative manner. A stochastic thermodynamics framework is employed to analyze a fluctuation theorem for the…

Statistical Mechanics · Physics 2025-11-14 D. Evan Piephoff , Jianshu Cao

Measuring and quantifying non-equilibrium dynamics in active biological systems is a major challenge, because of their intrinsic stochastic nature and the limited number of variables accessible in any real experiment. We investigate what…

Biological Physics · Physics 2018-07-25 Federica Mura , Grzegorz Gradziuk , Chase P. Broedersz

Stochastic thermodynamics has largely succeeded in characterizing both equilibrium and far-from-equilibrium phenomena. Yet many opportunities remain for application to mesoscopic complex systems -- especially biological ones -- whose…

Statistical Mechanics · Physics 2022-11-08 Mikhael T. Semaan , James P. Crutchfield

Living organisms are inherently out-of-equilibrium systems. We employ new developments in stochastic energetics and rely on a minimal microscopic model to predict the amount of mechanical energy dissipated by such dynamics. Our model…

Detecting and quantifying non-equilibrium activity is essential for studying internally driven assemblies, including synthetic active matter and complex living systems such as cells or tissue. We discuss a non-invasive approach of measuring…

Biological Physics · Physics 2019-05-22 Grzegorz Gradziuk , Federica Mura , Chase P. Broedersz

Active matter systems exhibit rich emergent behavior due to constant injection and dissipation of energy at the level of individual agents. Since these systems are far from equilibrium, their dynamics and energetics cannot be understood…

Soft Condensed Matter · Physics 2020-04-23 Sarah Eldeen , Ryan Muoio , Paris Blaisdell-Pijuan , Ngoc La , Mauricio Gomez , Alex Vidal , Wylie Ahmed

We explore the idea that non-equilibrium steady states breaking detailed balance are obtained by deforming trajectories (lines in space-time) that have been sampled in a reference system with stochastic dynamics obeying detailed balance,…

Statistical Mechanics · Physics 2018-04-17 Thomas Speck

Using equilibrium fluctuations to understand the response of a physical system to an externally imposed perturbation is the basis for linear response theory, which is widely used to interpret experiments and shed light on microscopic…

Statistical Mechanics · Physics 2024-06-24 Jérémie Klinger , Grant M. Rotskoff

Biologically driven non-equilibrium fluctuations are often characterized by their non-Gaussianity or by an "effective temperature", which is frequency dependent and higher than the ambient temperature. We address these two measures…

Biological Physics · Physics 2011-06-16 Eyal Ben-Isaac , YongKeun Park , Gabriel Popescu , Frank L. H. Brown , Nir S. Gov , Yair Shokef

We investigate proteins within heterogeneous cell membranes where non-equilibrium phenomena arises from spatial variations in concentration and temperature. We develop simulation methods building on non-equilibrium statistical mechanics to…

Soft Condensed Matter · Physics 2025-08-28 D. Jasuja , P. J. Atzberger

The mechanical properties within living cells play a critical role in the adaptive regulation of their biological functions upon environmental and internal stimuli. While these properties exhibit nonequilibrium dynamics due to the thermal…

Biological Physics · Physics 2024-08-14 Yuika Ueda , Shinji Deguchi

Small thermodynamic systems exhibit peculiar behavior different from that observed in long-scale systems. Non-equilibrium processes taking place in those systems are strongly influenced by the presence of fluctuations which can be large.…

Statistical Mechanics · Physics 2007-05-23 J. M. Rubi

Stochastic thermodynamics as reviewed here systematically provides a framework for extending the notions of classical thermodynamics like work, heat and entropy production to the level of individual trajectories of well-defined…

Statistical Mechanics · Physics 2015-06-05 Udo Seifert

Living cells exhibit an important out-of-equilibrium mechanical activity, mainly due to the forces generated by molecular motors. These motor proteins, acting individually or collectively on the cytoskeleton, contribute to the violation of…

Biological Physics · Physics 2009-01-21 Francois Gallet , Delphine Arcizet , Pierre Bohec , Alain Richert

Life is characterized by a myriad of complex dynamic processes allowing organisms to grow, reproduce, and evolve. Physical approaches for describing systems out of thermodynamic equilibrium have been increasingly applied to living systems,…

Biological Physics · Physics 2020-12-16 Xiaona Fang , Karsten Kruse , Ting Lu , Jin Wang

Time-lapse microscopy imaging provides direct access to the dynamics of soft and living systems. At mesoscopic scales, such microscopy experiments reveal intrinsic fluctuations, which may have both thermal and non-equilibrium origins. These…

Biological Physics · Physics 2020-12-02 F. S. Gnesotto , G. Gradziuk , P. Ronceray , C. P. Broedersz
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