Phosphorylation energy hypothesis: open chemical systems and their biological functions. Author: Qian H. Affiliation: Department of Applied Mathematics, University of Washington, Seattle, WA 98195, USA. qian@amath.washington.edu Conference/Journal: Annu Rev Phys Chem. Date published: 2007 Other: Volume ID: 58 , Pages: 113-42 , Word Count: 99 Biochemical systems and processes in living cells generally operate far from equilibrium. This review presents an overview of a statistical thermodynamic treatment for such systems, with examples from several key components in cellular signal transduction. Open-system nonequilibrium steady-state (NESS) models are introduced. The models account quantitatively for the energetics and thermodynamics in phosphorylation-dephosphorylation switches, GTPase timers, and specificity amplification through kinetic proofreading. The chemical energy derived from ATP and GTP hydrolysis establishes the NESS of a cell and makes the cell--a mesoscopic-biochemical reaction system that consists of a collection of thermally driven fluctuating macromolecules--a genetically programmed chemical machine. PMID: 17059360