Independent fluctuations of malate and citrate in the CAM species Clusia hilariana Schltdl. under low light and high light in relation to photoprotection.

Clusia hilariana Schltdl. is described in literature as an obligate Crassulacean acid metabolism (CAM) species. In the present study we assessed the effect of irradiance with low light (LL, 200µmolm(-2)s(-1)) and high light (HL, 650-740µmolm(-2)s(-1)), on the interdependency of citrate and malate diurnal fluctuations. In plants grown at HL CAM-type oscillations of concentration of citrate and malate were obvious. However, at LL daily courses of both acids do not seem to indicate efficient utilization of these compounds as CO2 and NADPH sources. One week after transferring plants from LL to HL decarboxylation of malate was accelerated. Thus, in the CAM plant C. hilariana two independent rhythms of accumulation and decarboxylation of malate and citrate take place, which appear to be related to photosynthesis and respiration, respectively. Non photochemical quenching (NPQ) of photosystem II, especially well expressed during the evening hours was enhanced. Exposure to HL for 7 d activated oxidative stress protection mechanisms such as the interconversion of violaxanthin (V), antheraxanthin (A) and zeaxanthin (Z) (epoxydation/de-epoxydation) measured as epoxydation state (EPS). This was accompanied by a slight increase in the total amount of these pigments. However, all these changes were not observed in plants exposed to HL for only 2 d. Besides violaxanthin cycle components also lutein, which shows a small, but not significant increase, may be involved in dissipating excess light energy in C. hilariana.

Light stress is not effective to enhanced crassulacean acid metabolism.

Clusia minor L., a C3-CAM intermediate, and Clusia multiflora H. B. K., a C3 obligate, present two physiotypes of a similar morphotype occurring sympatrically in the field. Both species, exposed 2 days to high light, show similar responses to this kind of stress: (i) the level of xanthophyll pigments in tested plants during the daycourse adapts to stress, (ii) the levels of antheraxanthin and zeaxanthin clearly increase during the afternoon showing increased de-epoxidation, (iii) the changes in the xanthophyll cycle are similar. Exposure to high light increases the malate levels in C. minor during the afternoon while decreases the day/night changes of the malate levels, and hence the Crassulacean Acid Metabolism (CAM) expression. It can be concluded that strong light applied as a single stress factor to well-watered plants is not effective in strengthing the CAM metabolism in a C3-CAM intermediate plant but rather suppresses the CAM activity despite exposure to high light energy. It is suggested that, when water supply is not limiting and other stresses do not prevail, C3 allows to use up the citrate pool, especially in the afternoon and enables a superior daily photon utilization.

Adaptation of the obligate CAM plant Clusia alata to light stress: Metabolic responses.

In the Crassulacean acid metabolism (CAM) plants Clusia alata Triana and Planch., decarboxylation of citrate during phase III of CAM took place later than malate decarboxylation. The interdependence of these two CO(2) and NADPH sources is discussed. High light accelerated malate decarboxylation during the day and lowered citrate levels. Strong light stress also activated mechanisms that can protect the plant against oxidative stress. Upon transfer from low light (200micromol m(-2)s(-1)) to high light (650-740micromol m(-2)s(-1)), after 2 days, there was a transient increase of non-photochemical quenching (NPQ) of fluorescence of chlorophyll a of photosystem II. This indicated acute photoinhibition, which declined again after 7 days of exposure. Conversely, after 1 week exposure to high light, the mechanisms of interconversion of violaxanthin (V), antheraxanthin (A), zeaxanthin (Z) (epoxydation/de-epoxydation) were activated. This was accompanied by an increase in pigment levels at dawn and dusk.

Canopy CO2 exchange of two neotropical tree species exhibiting constitutive and facultative CAM photosynthesis, Clusia rosea and Clusia cylindrica.

Photon flux density (PFD) and water availability, the daily and seasonal factors that vary most in tropical environments, were examined to see how they influenced expression of crassulacean acid metabolism (CAM) in 3-year-old Clusia shrubs native to Panama. Instead of the commonly used single-leaf approach, diel CO(2) exchange was measured for whole individual canopies of plants in large soil containers inside a naturally illuminated 8.8 m(3) chamber. In well-watered C. rosea, a mainly constitutive CAM species, nocturnally fixed CO(2) contributed about 50% to 24 h carbon gain on sunny days but the contribution decreased to zero following overcast days. Nonetheless, CO(2) fixation in the light responded in such a way that 24 h carbon gain was largely conserved across the range of daily PFDs. The response of C. rosea to drought was similarly buffered. A facultative component of CAM expression led to reversible increases in nocturnal carbon gain that offset drought-induced reductions of CO(2) fixation in the light. Clusia cylindrica was a C(3) plant when well-watered but exhibited CAM when subjected to water stress. The induction of CAM was fully reversible upon rewatering. C. cylindrica joins C. pratensis as the most unambiguous facultative CAM species reported in the genus Clusia.