Crassulacean acid metabolism: plastic, fantastic

The occurrence, activity and plasticity of the CAM pathway is described from an introductory viewpoint, framed by the use of the four "Phases" of CAM as comparative indicators of the interplay between environmental constraints and internal molecular and biochemical regulation. Having described a number of "rules" which seem to govern the CAM cycle and apply uniformly to most species, a number of key regulatory points can then be identified. These include temporal separation of carboxylases, based on the circadian expression of key genes and their control by metabolites. The role of a circadian oscillator and interplay between tonoplast and nuclear control are central to maintaining the CAM cycle. Control of reserve carbohydrates is often neglected, but the importance of daily partitioning (for growth and the subsequent night-time CAM activity) and use at night is shown to drive the CAM cycle. Finally, it is shown that the genotypic and phenotypic plasticity in patterns of CAM expression is mediated partly by environmental conditions and molecular signalling, but also by diffusive constraints in succulent tissues. A transformation system is now required to allow these key areas of control to be elucidated.