Aiming at understanding the odd case of CAM expression by a C4 plant, some properties of phosphoenolpyruvate carboxylase (PEPC, EC 4.1.1.31, orthophosphate: oxaloacetate carboxylyase, phosphorylating) were comparatively studied in leaves of CAM-expressing and non-expressing Portulaca oleracea L. plants. CAM expression was induced by growing plants under an 8-h photoperiod and under water-stress. CAM induction in leaves of these plants (designated as CAM) is indicated by the nocturnal acidification and by the clear diurnal oscillation pattern and amplitude of acidity, malic acid, and PEPC activity characteristic of CAM plants. Treatment of the other plant group (designated as C4) by growth under a 16-h photoperiod and well-watered conditions did not induce expression of the tested criteria of CAM in plants. In these C4 plants, the mentioned CAM criteria were undetectable. PEPC from CAM and C4Portulaca responded differently to any of the studied assay conditions or effectors. For example, extent and timing of sensitivity of PEPC to pH change, inhibition by malate, activation by glucose-6-phosphate or inorganic phosphate, and the enzyme affinity to the substrate PEP were reversed with induction of CAM from the C4-P. oleracea. These contrasting responses indicate distinct kinetic and regulatory properties of PEPC of the two modes. Thus by shifting to CAM in the C4Portulaca a new PEPC isoform may be synthesised to meet CAM requirements. Simultaneous occurrence of both C4 and CAM is suggested in P. oleracea when challenged with growth under stress.
In Sedum wrightii grown in a growth chamber, detached leaves could survive for at least 120 d with a high rate of success for propagule formation. The pattern of gas exchange, associated with CAM, may be important in extending the period during which the detached leaf remains physiologically active. The added benefit for the developing propagule, still attached to the "parent" leaf, is an additional source of water and saccharide reserves over an extended period necessary for rooting. Drought survival of propagules may be determined by the amount of water-storing tissue in the detached leaf.
Orcuttieae is a small tribe of C4 grasses endemic to seasonal pools in the southwestern U.S., comprising the basal genus Neostapfia, Tuctoria, and the most derived group, Orcuttia. Growth is initiated underwater, and when pools dry, species undergo a metamorphosis replacing aquatic foliage with terrestrial foliage. O. californica and O. viscida exhibit CAM-like diel fluctuations in acidity in the aquatic foliage. Pulse-chase studies showed that although CO2 was fixed into malic acid in the dark, an overnight chase in the dark revealed that most label was not retained in organic acids, indicating a role other than CAM. Terrestrial foliage exhibited a very different diel fluctuation; acids accumulated during the day, and diminished overnight. Malic acid predominated and was secreted on the surface of the leaf in a manner similar to another arid land species. This terrestrial daytime acid accumulation may not be related to photosynthetic pathway but may play an anti-herbivore function. No acid fluctuations were observed in either N. colusana or T. greenei.