Endogenous rhythms and chaos in crassulacean acid metabolism.

Endogenous free-running regular circadian oscillations of net CO2 exchange in the crassulacean-acidmetabolism (CAM) plant Kalanchoë daigremontiana Hamet et Perrier de la Bâthie under constant external conditions in continuous light have been shown to change to irregular non-predictable (chaotic) time behaviour as irradiance or temperature are raised above a critical level. A model of CAM has been constructed with pools of major metabolites of varying concentrations, flows of metabolites leading to exchange between pools, metabolite transformations determined by chemical reactions, and feedback regulations. The model is described by a system of coupled non-linear differential equations. It shows stable rhythmicity in normal dark-light cycles and in continuous light and, like the K. daigremontiana leaves in the experiments, a change to chaos as irradiance is increased. The maintenance of endogenous oscillations in the model is brought about by a hysteresis switch or beat oscillator between two stable oscillation modes. In CAM these stable modes are vacuolar malate accumulation and remobilization. The model shows that the physical nature of the beat oscillator in the leaves can be explained by the balance between active and passive transport at the tonoplast.