Recycling of respiratory CO2 during Crassulacean acid metabolism: alleviation of photoinhibition in Pyrrosia piloselloides

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Abstract
The regulation of Crassulacean acid metabolism (CAM) in the fern Pyrrosia piloselloides (L.) Price was investigated in Singapore on two epiphytic populations acclimated to sun and shade conditions. The shade fronds were less succulent and had a higher chlorophyll content although the chlorophyll a:b ratio was lower and light compensation points and dark-respiration rates were reduced. Dawn-dusk variations in titratable acidity and carbohydrate pools were two to three times greater in fronds acclimated to high photosynthetically active radiation (PAR), although water deficits were also higher than in shade fronds. External and internal CO2 supply to attached fronds of the fern was varied so as to regulate the magnitude of CAM activity. A significant proportion of titratable acidity was derived from the refixation of respiratory CO2 (27% and 35% recycling for sun and shade populations, respectively), as measured directly under CO2-free conditions. Starch was shown to be the storage carbodydrate for CAM in Pyrrosia, with a stoichiometric reduction of “C3-skeleton” units in proportion to malic-acid accumulation. Measurements of photosynthetic O2 evolution under saturating CO2 were used to compare the light responses of sun and shade fronds for each CO2 supply regime, and also following the imposition of a photoinhibitory PAR treatment (1600 μmol·m-2·s-1 for 3 h). Apparent quantum yield declined following the high-PAR treatment for sun- and shade-adapted plants, although for sun fronds CAM activity derived from respiratory CO2 prevented any further reduction in photosynthetic efficiency. Recycling of respiratory CO2 by shade plants could only partly prevent photoinhibitory damage. These observations provide experimental evidence that respiratory CO2 recycling, ubiquitous in CAM plants, may have developed so as to alleviate photoinhibition.