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Driving assembly far from equilibrium

Nonequilibrium assembly and disassembly underlie key biological functions, such as growth, movement, and the dramatic morphological transformations that occur during organismal development. These processes are driven and regulated by energy dissipation, e.g. through nucleotide triphosphate hydrolysis. Energy consumption frees the organism from the rigid constraints of thermodynamic equilibrium, but also poses a significant challenge for researchers hoping to understand biological assembly or design synthetic systems with similar capabilities. While statistical mechanics can, in principle, predict equilibrium distributions of self-assembly reactions from microscopic interactions and dynamics, there is no analogous theory for nonequilibrium steady-states and the fluctuations around them. Despite recent progress through techniques such as computer simulations, large deviation theory, and stochastic thermodynamics (e.g. [1–11]), we are far from achieving a ‘grand challenge’ of nonequilibrium statistical mechanics: predicting the emergent products and functionalities of nonequilibrium assembly, or more ambitiously designing an assembly process to achieve a desired function. This commentary will focus on a recent advance in nonequilibrium assembly described in two papers by Nguyen and Vaikuntanathan [7, 12]. In [7], by using stochastic thermodynamics to calculate the entropy change during assembly and invoking the second law of thermodynamics, they obtained a bound on the nonequilibrium driving force required to perturb the ensemble of assembly configurations from its equilibrium distribution: