Nonequilibrium physics in biology

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, which often exhibit phenomena unknown from those traditionally studied in physics. Spectacular advances in experimentation during the last decade or two, for example, in microscopy, single cell dynamics, in the reconstruction of sub- and multicellular systems outside of living organisms, or in high throughput data acquisition have yielded an unprecedented wealth of data about cell dynamics, genetic regulation, and organismal development. These data have motivated the development and refinement of concepts and tools to dissect the physical mechanisms underlying biological processes. Notably, the landscape and flux theory as well as active hydrodynamic gel theory have proven very useful in this endeavour. Together with concepts and tools developed in other areas of nonequilibrium physics, significant progresses have been made in unraveling the principles underlying efficient energy transport in photosynthesis, cellular regulatory networks, cellular movements and organization, embryonic development and cancer, neural network dynamics, population dynamics and ecology, as well as ageing, immune responses and evolution. Here, we review recent advances in nonequilibrium physics and survey their application to biological systems. We expect many of these results to be important cornerstones as the field continues to build our understanding of life.