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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

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

Fluctuation theorems are fundamental extensions of the second law of thermodynamics for small nonequilibrium systems. While work and heat are equally important forms of energy exchange, fluctuation relations have not been experimentally…

Statistical Mechanics · Physics 2022-02-24 Markus Rademacher , Michael Konopik , Maxime Debiossac , David Grass , Eric Lutz , Nikolai Kiesel

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

Evaluating the linear response of a driven system to a change in environment temperature(s) is essential for understanding thermal properties of nonequilibrium systems. The system is kept in weak contact with possibly different fast…

Statistical Mechanics · Physics 2014-12-01 Marco Baiesi , Urna Basu , Christian Maes

The glassy cytoplasm, crowded with bio-macromolecules, is fluidized in living cells by mechanical energy derived from metabolism. Characterizing the living cytoplasm as a non-equilibrium system is crucial in elucidating the intricate…

Biological Physics · Physics 2023-07-28 K. Umeda , K. Nishizawa , W. Nagao , S. Inokuchi , Y. Sugino , H. Ebata , D. Mizuno

We propose a model for the dynamics of a probe embedded in a living cell, where both thermal fluctuations and nonequilibrium activity coexist. The model is based on a confining harmonic potential describing the elastic cytoskeletal matrix,…

Biological Physics · Physics 2015-06-18 É. Fodor , M. Guo , N. S. Gov , P. Visco , D. A. Weitz , F. van Wijland

Biological activity gives rise to non-equilibrium fluctuations in the cytoplasm of cells; however, there are few methods to directly measure these fluctuations. Using a reconstituted actin cytoskeleton, we show that the bending dynamics of…

Soft Condensed Matter · Physics 2009-11-13 C. P. Brangwynne , G. H. Koenderink , F. C. MacKintosh , D. A. Weitz

Living systems operate far from thermodynamic equilibrium. Enzymatic activity can induce broken detailed balance at the molecular scale. This molecular scale breaking of detailed balance is crucial to achieve biological functions such as…

Biological Physics · Physics 2018-05-02 F. Gnesotto , F. Mura , J. Gladrow , C. P. Broedersz

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

The fundamental insight into Brownian motion by Einstein is that all substances exhibit continual fluctuations due to thermal agitation balancing with the frictional resistance. However, even at thermal equilibrium, biological activity can…

Soft Condensed Matter · Physics 2014-02-19 Soya Shinkai , Yuichi Togashi

The dynamics and structure of nonequilibrium liquids, driven by non-conservative forces which can be either external or internal, generically hold the signature of the net dissipation of energy in the thermostat. Yet, disentangling…

Statistical Mechanics · Physics 2019-11-06 Laura Tociu , Étienne Fodor , Takahiro Nemoto , Suriyanarayanan Vaikuntanathan

In this communication, we propose a model to study the non-equilibrium process by which actin stress fibers develop force in contractile cells. The emphasis here is on the non-equilibrium thermodynamics, which is necessary to address the…

Subcellular Processes · Quantitative Biology 2014-04-25 Mirko Maraldi , Krishna Garikipati

Diverse physical systems are characterized by their response to small perturbations. Near thermodynamic equilibrium, the fluctuation-dissipation theorem provides a powerful theoretical and experimental tool to determine the nature of…

Statistical Mechanics · Physics 2020-03-25 Jeremy A. Owen , Todd R. Gingrich , Jordan M. Horowitz

We analyze the non-equilibrium shape fluctuations of giant unilamellar vesicles encapsulating motile bacteria. Owing to bacteria--membrane collisions, we experimentally observe a significant increase in the magnitude of membrane…

Soft Condensed Matter · Physics 2020-04-22 Sho C. Takatori , Amaresh Sahu

We formulate theoretical modeling approaches and develop practical computational simulation methods for investigating the non-equilibrium statistical mechanics of fluid interfaces with passive and active immersed particles. Our approaches…

Soft Condensed Matter · Physics 2024-10-03 Dev Jasuja , Paul J. Atzberger

The mechanics of cells is strongly affected by molecular motors that generate forces in the cellular cytoskeleton. We develop a model for cytoskeletal networks driven out of equilibrium by molecular motors exerting transient contractile…

Soft Condensed Matter · Physics 2009-11-13 Fred C. MacKintosh , Alex J. Levine

In development and homeostasis, multi-cellular systems exhibit spatial and temporal heterogeneity in their biochemical and mechanical properties. Nevertheless, it remains unclear how spatiotemporally heterogeneous forces affect the…

Soft Condensed Matter · Physics 2022-02-18 Takaki Yamamoto , Daniel M. Sussman , Tatsuo Shibata , M. Lisa Manning

In striking contrast to equilibrium systems, inertia can profoundly alter the structure of active systems. Here, we demonstrate that driven systems can exhibit effective equilibrium-like states with increasing particle inertia, despite…

Soft Condensed Matter · Physics 2023-02-23 Ahmad K. Omar , Katherine Klymko , Trevor GrandPre , Phillip L. Geissler , John F. Brady

Nonequilibrium systems exchange the energy with an environment in the form of work and heat. The work done on a system obeys the fluctuation theorem, while the dissipated heat which differs from the work by the internal energy change does…

Statistical Mechanics · Physics 2014-02-04 Jae Dong Noh
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