A succulent is a plant with water storing tissue, but succulence clearly is a quality that can be possessed to a higher or lesser degree. This paper gives a definition of succulence and discusses problems concerning Dells measure of succulence. A new measure, Succulence Quotient, is proposed. It measures the amount of water that a plant can store at the expenditure of one gram of organic matter. We demonstrate this measure on a number of plants from the Namib desert, southern Africa, and compare it with the measured caloric values of the plant tissues.We also discuss life cycles of desert succulents in terms of utilizable and structural biomass and water. We contrast the concepts of growth form, life form and life strategy, and propose, on the basis of life cycle characteristics, an outline classification of life strategies of desert succulents. One detail of this outline classification is further worked out and illustrated for succulents from the Namib desert.
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.
The combination of a chamber for CO2 gas exchange with a potometric measuring arrangement allowed concomitant investigations into CO2 gas exchange, transpiration and water uptake by the roots of whole plants of Senecio medley-woodii, a species which exhibits Crassulacean acid metabolism. The water-uptake rate showed the same daily pattern as malate concentration and osmotic potential. The accumulation of organic acids resulting from nocturnal CO2 fixation enhanced the water-uptake rate from dusk to dawn. During the day the water-uptake rates decreased with decreasing organic-acid concentration. With gradually increasing water stress, CO2 dark fixation of S. medley-woodii was increased as long as water could be taken up by the roots. It was also shown that a reestablished water supply after drought caused a similar increase which in both cases ameliorated the water uptake in order to conserve a positive water balance for as long as possible. This water-uptake pattern shows that Crassulacean acid metabolism is not only a water-saving adaptation but also enhances water uptake and is directly correlated with the amelioration of the plant water status.
The spectral properties of juvenile and adult leaves of Tussilago farfara L. were measured for the wave range from 400 to 1350 nm. The significance of leaf pubescence for the absorption of global radiation was determined. The absorption is lowered by epidermal harrs only for the visible part of the spectrum (400-1350 nm). Absorptivities of the infrared do not vary whether hairs are present or not. The spectral properties of the upper and the lower surfaces without hairs are not equal. The hairs on the lower surface increase the amount of infrared radiation that is reflected by the upper surface. As shown by calculations of the absorbed radiation, pubescence has little influence on energy input. If leaves are put in an inverse position, the absorption of the global radiation energy is lowered by 15% compared with the same leaf in the regular position.
The diurnal course of CO gas exchange, CO incorporation, malate and citrate content, and traspiration of Welwitschia mirabilis were measured in one of its natural habitats, the Welwitschia-Vlakte in the central Namib desert (Namibia), in order to decide which CO fixation pathway is used by this gymnosperm.The CO gas exchange of Welwitschia is that of a C plant under arid conditions. Younger leaf parts show a two-peaked pattern of photosynthetic CO uptake whereas in older parts the morning peak is followed by net CO release during the rest of the day. The maximum rates of net photosynthesis decrease from 3.4 μmol m s in 1-year-old parts to 1 μmol m s in 7-year-old parts. No net CO uptake was detected during the night. The diurnal CO balance indicates that the old leaf parts live at the expense of the younger ones. Irrigation of Welwitschia plants resulted in an increased CO uptake throughout the light period with maximum rate of 4.1 μmol m s. CO was only incorporated during the day.The water loss of Welwitschia by transpiration is considerable, reaching a peak value of 1.9 mmol m s around noon. Leaf conductance corresponds with the twopeaked pattern of CO uptake.Although there is no sign of a crassulacean acid metabolism in Welwitschia the leaf contains rather high amounts of malate (up to 200 μmol g dry matter) and citrate (up to 250 μmol g dry matter), which depend on leaf age but do not show any significant day-night oscillation.In spite of all this the δC values are in the range of-17.77 to-19.64‰. Possible reasons for such a high C content in a C plant are discussed.
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