Design principles of biochemical oscillators

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Abstract
Many physiological behaviours of cells are repeated periodically in time, such as hormone secretion, second messenger signalling, cell-division cycles and circadian rhythms. Oscillatory behaviour is a systems-level property of the interactions of genes, proteins and metabolites in the cell. All biochemical oscillators are characterized by negative feedback with time delay. Time delay can be due to a chain of intermediates between the 'cause' and 'effect' of the negative-feedback loop or to a positive-feedback loop in the network. For a biochemical reaction network to oscillate, the kinetics of the reactions must be sufficiently nonlinear and their timescales must be properly balanced, in quantitative terms that are predicted by theoretical analysis. These conditions are satisfied by various specific network topologies that provide the basis for a classification of biochemical oscillators.