Crassulacean acid metabolism (CAM) is considered to be an
adaptive mechanism allowing many terrestrial and epiphytic
succulents to survive in periodically droughted habitats. In CAM
plants, carbon dioxide uptake occurs during the night, when
stomata open; carbon dioxide is combined with phosphoenolpyruvate
to yield oxaloacetate, which is reduced to malate. Malate is
transported into the vacuole and malic acid accumulates during
the night. Nocturnal acid accumulation and nocturnal stomatal
aperture are the main diagnostic features of CAM. During the day
malate is decarboxylated in the cytoplasm, providing Rubisco with
increased partial pressures of carbon dioxide during C3 photosynthesis.
The net outcome of the functioning of CAM is that
carbon dioxide is fixed with significant water saving relative to C3
photosynthesis, since the process occurs at times of lower
evaporative demand and a larger air-leaf carbon dioxide concentration
gradient [1–3].
In C3 plants, water deficit may lead to an imbalance between the
production and consumption of reducing equivalents and ATP in
photosynthesis by inducing stomatal closure and consequently
decreasing carbon dioxide assimilation during the day. This may
enhance excess excitation energy in thylakoids of stressed plants,
which must activate adequate photoprotection mechanisms, as
those described for high light-acclimated plants, to dissipate
excess energy and afford protection to photosystem II (PSII) [4].
Osmond suggested that high daytime partial pressures of carbon
dioxide inside the leaves and re-fixation of respiratory CO2 might
contribute to the protection of the photosynthetic machinery
against high irradiance with closed stomata in CAM plants [5].
Since then, some evidence has been gathered, most of it in
facultative CAM plants, in favor of this hypothesis (reviewed by
[3]). However, it has been shown that photoprotection through the
xanthophyll cycle may not necessarily be more effective in CAM
than in C3 species, as shown in a comparison of Clusia multiflora
(obligate C3) and Clusia minor (facultative CAM). Zeaxanthin