The outermost phellems of Abies alba Mill., Acer pseudoplatanus L., Aesculus hippocastanum L., Betula potaninii L.C. Hue and Sambucus nigra L. have been isolated enzymatically, resulting in membranes with five to seven heavily suberized cork cell layers. Water and oxygen permeances were determined for the phellem areas without lenticels. A special diaphragm made it possible to quantify permeances of single lenticels for the first time. The water permeance of phellems was in the range of 3x10(-5) to 9x10(-5) ms(-1) and can be predicted from the density of the phellem membranes with 93% accuracy. Embedded waxes amounted to 3% ( Aesculus) and up to 35% ( Betula) of the dry weight but affected water permeance only to a small degree. The sorption isotherms describing the water content of the phellems in relation to relative humidities followed a hyperbolic shape and indicated varying water contents among plant species. It is argued that water transfer across the phellems occurs via the middle lamellae. Phellem membranes were impermeable to oxygen. Removal of the waxes hardly changed this situation. Single lenticels from Betula and Sambucus were significantly more permeable to water and oxygen than phellem areas without lenticels. The water permeance was elevated by factors of 39 for Betula and 12 for Sambucus, the oxygen permeance by factors of 1,202 for Betula and 53 for Sambucus. Extraction of lenticels did not affect permeance. A quantitative comparison of the gas-exchange capacity of lenticels and stomata demonstrated the superiority of stomata. However, differences may be not more than one order of magnitude.

Crassulacean acid metabolism (CAM) plants are dependent on the organic acids that accumulate overnight in the vacuoles as a source of CO(2) during the daylight deacidification period, when stomata are closed and high irradiances generally prevail. We performed an integrative analysis of diurnal changes in gas exchange, chlorophyll fluorescence parameters and organic acid decarboxylation to understand the adjustments in photochemical and non-photochemical processes during the different CAM phases in Clusia hilariana Schlecht., a dominant tree species in the sandy coastal plains of southeastern Brazil. A linear relationship was obtained between the quantum yields of photochemical and non-photochemical quenching, irrespective of the CAM phase and prevailing irradiance. Degradation of malic and citric acids during the midday stomatal closure period could lead to potential CO(2) fixation rates of 23 &mgr;mol m(-2) s(-1), whereas CO(2) losses, measured as CO(2) evolution, corresponded to about 3% of this value. Thus, decarboxylation of malate and citrate provided high internal CO(2) concentrations during phase III of CAM, even though the stomata were closed, allowing optimal utilization of light energy, as indicated by the non-saturating electron transport rates (ETR) in the light response curves, with highest rates of ETR occurring at midday in the diurnal curves. At the transition from phase III to IV of CAM, depletion of internal CO(2) sources and low stomatal conductances, which restricted the supply of exogenous CO(2), reduced the demand for photochemical energy to drive carbon assimilation. This was compensated by increases in thermal energy dissipation as indicated by higher rates of non-photochemical quenching, while high irradiances still prevailed. Shifts in the CAM phases and changes in protective thermal dissipation potential allowed C. hilariana to match changes in PPFD patterns for leaves of different orientations. Evidence that most of the decline in photochemical efficiency was probably related to the fast-relaxing component of non-photochemical quenching is provided by the high values of the quantum yield of photosystem II after 20 min of relaxation in darkness, and an almost complete recovery after sunset. These adjustments in photosynthetic machinery minimized the danger of photo-inhibition in C. hilariana, which is commonly found in fully exposed habitats.

The C3/CAM intermediate species, C/usia parviflora Saldanha et EngI., and the obligate CAM species Clusia hilariana Schlecht., occur sympatrically in the coastal sand dune vegetation of the Restinga of Brazil. Their photosynthetic activity at an exposed and at a shaded site was compared by measuring gas exchange (porometry), chlorophyll a fluorescence parameters, organic acid levels (malic and citric) and carbon isotope ratios. At the shaded site, low photosynthetic photon flux densities (PPFD) strongly restricted photosynthetic activity. However, C parviflora could readily make use of light flecks. At the exposed site, C. parviflora was much less affected by photoinhibition than C. hilariana. The CAM species showed higher apparent rates of linear photosynthetic electron transport (ETR) and higher effective quantum yield of PSII (ΔF/F'm) than did C. parviflora during high insolation in the middle of the day, i.e., the time of Phase Ill of CAM. Nevertheless, it suffered much more severe acute photoinhibition that was not reversible after 20 min of darkening during this time, and even some chronic photoinhibition not reversible overnight. Comparative studies of sympatric physiotypes with different modes of photosynthesis of a given leaf morphotype, as available in the genus Cksia, challenge some CAM dogmas, e.g., CAM may not always be superior at exposed sites and may not always provide better photoprotection at high PPFD. However, the idea that C3/CAM plasticity allows occupation of a wider range of habitats is supported.

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