CARBON DIOXIDE AND WATER DEMAND: CRASSULACEAN ACID METABOLISM (CAM), A VERSATILE ECOLOGICAL ADAPTATION EXEMPLIFYING THE NEED FOR INTEGRATION IN ECOPHYSIOLOGICAL WORK

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Abstract
Summary: Plants having crassulacean acid metabolism (CAM) tend to occupy habitats where the prevailing environmental stress is scarcity of water. These are semi‐arid or arid regions, salinas or epiphytic sites. CAM plants manage the dilemma of desiccation or starvation by nocturnal malic acid accumulation in the vacuoles. Malic acid serves as a form of CO2 storage and as an osmoticum. In this way malic acid accumulation allows, firstly, separation of uptake and assimilation of atmospheric CO2 with water‐saving daytime stomatal closure and, secondly, osmotic acquisition of water. There is no very special trait which is specific for CAM. An array of biophysical and biochemical functional elements, which are also found in other plants, is integrated in CAM performance. This leads to a large diversity of behaviour which makes CAM plants highly versatile in their response to environmental variables. Besides CO2 dark fixation, transport of malic acid across the tonoplast is one of the key elements in CAM function. This is examined in detail at the level of membrane biophysics and biochemistry. The versatility of CAM is illustrated by examples from field work, with comparisons involving different species, seasons, modes of photosynthesis (CAM vs C3), kinds of stress and ways of stress imposition. Contents Summary 593 I. Studies of CAM: an example for the ecophysiological approach 594 II. Malic acid transport at the tonoplast 602 III. Regulation 605 IV. Desiccation or starvation 610 V. Comparative autecology 614 VI. Ecology: promise of integration 621 Acknowledgements 622 References 622