Optional use of CAM photosynthesis in two C4 species, Portulaca cyclophylla and Portulaca digyna

Low levels of crassulacean acid metabolism (CAM) are demonstrated in two species with C4 photosynthesis, Portulaca cyclophylla and P. digyna. The expression of CAM in P. cyclophylla and P. digyna is facultative, i.e. optional. Well-watered plants did not accumulate acid at night and exhibited gas-exchange patterns consistent with C4 photosynthesis. CAM-type nocturnal acidification was reversible in that it was induced following drought and lost when droughted plants were rewatered. In P. cyclophylla, droughting was accompanied by a small but discernible net uptake of CO2 during the dark, whereas in P. digyna, net CO2 exchange at night approached the CO2 compensation point but did not transition beyond it. This report brings the number of known C4 species with a capacity for expressing CAM to six. All are species of Portulaca. The observation of CAM in P. cyclophylla and P. digyna is the first for species in the opposite-leaved (OL) Portulacelloid-anatomy lineage of Portulaca and for the Australian clade therein. The other four species are within the alternate-leaved (AL) lineage, in the Atriploid-anatomy Oleracea and the Pilosoid-anatomy Pilosa clades. Studies of the evolutionary origins of C4 and CAM in Portulaca will benefit from a more wide-range survey of CAM across its species, particularly in the C3-C4 intermediate-containing Cryptopetala clade.