A Comparative Survey of the Function, Mechanism and Control of Cellular Oscillators

Abstract
This review attempts to survey in a uniform manner the available evidence concerning the generation and behaviour of several well-investigated cellular oscillators. Members of two broad classifications are contrasted: (i) cytoplasmic oscillations, where the periodic phenomena is generated by an instability in a metabolic pathway and (ii) membrane oscillators in which a membrane potential rhythm is generated at the membrane. Interactions between the cytoplasmic and membrane compartments are considered and the effects of these interactions on oscillatory behaviour is discussed. Because of their biological importance and the greater body of experimental results, particular attention is directed to a study of membrane potential oscillations. These systems can be approximately classified in two groups: (i) systems in which a periodic potential results from oscillatory changes in permeability and (ii) systems in which potential oscillations result from the periodic activity of an electrogenic pump. The examples considered include the glycolytic oscillator, oscillations in vein contraction in the slime mould Physarum polycephalum, rhythmic aggregation in Dictyostelium discoideum, neural oscillators, the periodic potential in Purkinje fibres and the sino-atrial node and rhythmic behaviour in smooth muscle. Questions considered include the generation of periodic activity, the modulation of the oscillation by drugs and other metabolic and membrane effectors and the question of the functional role of these oscillations.