期刊
REPORTS ON PROGRESS IN PHYSICS
卷 81, 期 6, 页码 -出版社
IOP Publishing Ltd
DOI: 10.1088/1361-6633/aab3ed
关键词
non-equilibrium; fluctuations; active living matter; fluctuation-dissipation theorem; detailed balance; cellular biophysics
资金
- German Excellence Initiative via the program NanoSystems Initiative Munich (NIM)
- German Research Council (DFG) [Transregio 174]
- DFG through the Graduate School of Quantitative Biosciences Munich (QBM)
- European Union's Horizon research and innovation programme under European Training Network (ETN) [674979-NANOTRANS]
- National Science Foundation [PHY-1607611]
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 high-fidelity transcription and translation, sensing, adaptation, biochemical patterning, and force generation. While biological systems such as motor enzymes violate detailed balance at the molecular scale, it remains unclear how non-equilibrium dynamics manifests at the mesoscale in systems that are driven through the collective activity of many motors. Indeed, in several cellular systems the presence of non-equilibrium dynamics is not always evident at large scales. For example, in the cytoskeleton or in chromosomes one can observe stationary stochastic processes that appear at first glance thermally driven. This raises the question how non-equilibrium fluctuations can be discerned from thermal noise. We discuss approaches that have recently been developed to address this question, including methods based on measuring the extent to which the system violates the fluctuation-dissipation theorem. We also review applications of this approach to reconstituted cytoskeletal networks, the cytoplasm of living cells, and cell membranes. Furthermore, we discuss a more recent approach to detect actively driven dynamics, which is based on inferring broken detailed balance. This constitutes a non-invasive method that uses time-lapse microscopy data, and can be applied to a broad range of systems in cells and tissue. We discuss the ideas underlying this method and its application to several examples including flagella, primary cilia, and cytoskeletal networks. Finally, we briefly discuss recent developments in stochastic thermodynamics and non-equilibrium statistical mechanics, which offer new perspectives to understand the physics of living systems.
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