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Kinetic proofreading is an error correction mechanism present in theprocesses of the central dogma and beyond, and typically requires the freeenergy of nucleotide hydrolysis for its operation. Though the molecular playersof many biological proofreading schemes are known, our understanding of howenergy consumption is managed to promote fidelity remains incomplete. In ourwork, we introduce an alternative conceptual scheme called 'the piston model ofproofreading' where enzyme activation through hydrolysis is replaced withallosteric activation achieved through mechanical work performed by a piston onregulatory ligands. Inspired by Feynman's ratchet and pawl mechanism, weconsider a mechanical engine designed to drive the piston actions powered by alowering weight, whose function is analogous to that of ATP synthase in cells.Thanks to its mechanical design, the piston model allows us to tune the 'knobs'of the driving engine and probe the graded changes and trade-offs betweenspeed, fidelity and energy dissipation. It provides an intuitive explanation ofthe conditions necessary for optimal proofreading and reveals the unexpectedcapability of allosteric molecules to beat the Hopfield limit of fidelity byleveraging the diversity of states available to them. The framework that webuilt for the piston model can also serve as a basis for additional studies ofdriven biochemical systems.

Allostery and Kinetic Proofreading