Universal thermodynamic bounds on symmetry breaking in living systems: from error correction to pattern formation

Living systems are maintained out-of-equilibrium by external driving forces, thus continuously dissipating energy. Non-equilibrium steady states are characterized by emergent selection phenomena that break equilibrium symmetries dictated solely by energetic properties. In our recent work, we use the matrix-tree theorem to derive universal thermodynamic bounds on these symmetry-breaking features in biochemical systems. The presented bounds are independent of the kinetics and hold for both closed and open reaction networks, whether they are uni-molecular or catalytic. Our results can also be extended to the Master Equations in the chemical space. Using our framework, we recover the thermodynamic constraints in kinetic proofreading. Finally, we show that the contrast of reaction-diffusion patterns can be bounded only by the non-equilibrium driving force. The generality of these results paves the way to understanding the potentialities and limitations of non-equilibrium conditions in shaping steady-state properties of biochemical systems.