Availability of water controls Crassulacean acid metabolism in succulents of the Richtersveld (Namib desert, South Africa).

Features of Crassulacean acid metabolism (CAM) were studied in a variety of different succulents in response to climatic conditions between March 1977 and October 1983 in the southern Namib desert (Richtersveld). A screening in 1977 and 1978 revealed that nearly all investigated succulents performed a CAM, but overnight accumulation of malate declined gradually with decreasing soil water potential, tissue osmotic potential, and leaf water content. This was further substantiated by an extended period of insufficient rainfall in 1979 and 1980 which damaged the evergreen CAM succulents between 80 and 100%. In most of the species still living, neither CO2-gas exchange nor diurnal acid fluctuation, indicative of CAM, could be detected unless an abundant rainfall restored both CAM features. Plants persisted in a stage of latent life.Water supply is one necessary prerequisite for CAM in the Richtersveld. But even well-watered plants with CAM were sensitive to short-term water stress caused by high water-vapour partialpressure deficit (VPD) in the night, which reduced or prevented CO2 uptake and resulted in a linear relation between overnight accumulated malate and VPD. The results do not support the opinion that, for the Namib succulents, CAM is an adaptive mechanism to water stress since long-term and short-term water stress stopped nocturnal malate synthesis, but instead lead to the conclusion that nocuturnal CO2 fixation is only performed when the water status of the plant can be improved simultaneously.

Occurrence and changes of proline content in plants in the southern Namib Desert in relations to increasing and decreasing drought.

Over a period of seven years (1977-1983) the proline content and its responses to climatic changes were investigated in plants - especially Mesembryanthemaceae - in the southern Namib Desert (South Africa). Among 95 species in 26 families, 61 had detectable amounts of proline. In several of these species the proline content increased considerably in years with insufficient rainfall but decreased when the rainfall was abundant again. When individuals of the same species were grown at different sites, water availability in the soil determined their proline content. Many of the investigated species showed a clear diurnal fluctuation in their proline content with a remarkable proline accumulation during times of highest evaporative demand. In general, the higher the proline content the more pronounced were the changes, indicating that in these species-predominantly annual plants - proline was most probably involved in drought tolerance. The observation that proline accumulation and degradation reacted sensitively to changing climatic conditions over many years confirmed the correlation of proline synthesis to increasing water stress as postulated by the results of laboratory experiments with Mesembryanthemaceae.

Water loss and malate fluctuations during the day for plants in the southern Namib desert.

The daily course of transpiration for 12 different plants growing in the southern Namib desert was investigated. Sclerophyllous species with C3 photosynthesis were the most effective water savers followed by succulents exhibiting CAM, while C3 pathway-succulents lose as much water as malacophyllous species. The different species showed either one or two peaked patterns of both transpiration and leaf conductance. With C3 plants the most common pattern is a single morning peak in leaf conductance followed by decreases in conductance over the remainder on the day. With CAM succulents leaf conductance is high in the morning, shows a pronounced midday depression and increases in the early afternoon when the malate pool is depleted, but malate consumption did not start earlier than 4 h after dawn. Seven of nine investigated C3 plants had rather high contents of malate, up to 180 µmol·g-1 dry matter. In these plants the malate content increased with increasing leaf conductance and disappeared when leaf conductance declined in the evening, indicating that malate was synthesized during photosynthesis.

CO2 exchange of CAM exhibiting suceulents in the southern Namib desert in relation to microclimate and water stress.

The responses of CO2 exchange and overnight malate accumulation of leaf and stem succulent CAM-plants to water stress and the particular climatic conditiens of fog and föhn in the southern Namib desert have been investigated. In most of the investigated CAM plants a long term water stress gradually attenuated any uptake of external CO2 and led to CO2 release throughout day and night. No CAM-idling was observed. Rainfall or irrigation immediately restored daytime CO2 uptake while the recovery of the noctural CO2 uptake was delayed. Dawn peak of photosynthesis was only found in well watered plants but was markedly reduced by the short term water stress of a föhn-storm. Morning fog with its higher diffuse light intensity compared with clear days increased photosynthetic CO2 uptake considerably. Even in well watered plants noctural CO2 uptake and malate accumulation were strongly affected by föhn indicating that the water vapour pressure deficit during the night determines the degree of acidification.

CO2 exchange of CAM exhibiting succulents in the southern Namib desert in relation to microclimate and water stress.

The responses of CO2 exchange and overnight malate accumulation of leaf and stem succulent CAM-plants to water stress and the particular climatic conditions of fog and föhn in the southern Namib desert have been investigated. In most of the investigated CAM plants a long term water stress gradually attenuated any uptake of external CO2 and led to CO2 release throughout day and night. No CAM-idling was observed. Rainfall or irrigation immediately restored daytime CO2 uptake while the recovery of the nocturnal CO2 uptake was delayed. Dawn peak of photosynthesis was only found in well watered plants but was markedly reduced by the short term water stress of a föhn-storm. Morning fog with its higher diffuse light intensity compared with clear days increased photosynthetic CO2 uptake considerably. Even in well watered plants nocturnal CO2 uptake and malate accumulation were strongly affected by föhn indicating that the water vapour pressure deficit during the night determines the degree of acidification.

The activity and malate inhibition/stimulation of phosphoenolpyruvate-carboxylase in crassulacean-acid-metabolism plants in their natural environment.

The effect of environmental conditions, temperature, relative humidity, and light, together with the regulation of PEPC (phosphoenolpyruvate-carboxylase) activity by malate and pH on CAM (crassulacean acid metabolism), was studied in members of the Mesembryanthemaceae in their natural environment, the southern Namib desert. It was found that during a 24 h period the characteristics of PEPC change. Before sunrise the activity is higher when measured at pH 7 than 8. With bright sunlight the activity measured at pH 7 drops to 20% of its pre-sunrise value, the activity only recovers gradually after malate disappearance and stays constant throughout the night. When measured at pH 8, PEPC shows an opposite behavior, i.e., activity increases in bright sunlight and declines as the pH 7 activity increases. A day-night oscillation in the capacity of malate to stimulate or inhibit PEPC was found. During the day malate inhibits about 90% of the PEPC activity at both pH 7 and 8. After sunset there is a sudden decrease in this inhibition and, at pH 8, malate stimulates the activity by 50%. At pH 7 the stimulation was less.Both stomatal conductance and malate formation were found to increase only when the relative humidity at night rose to 80%. Changes in the properties of the PEPC coincided with the exposure to bright sunlight and changes in leaf temperature. The importance of these metabolic and environmental controls on the regulation of CAM in the Mesembryanthemaceae will be discussed.