Molecular cloning and characterization of a sodium‐pump ATPase of the moss Physcomitrella patens

Physcomitrella patens grew slowly at 600 mm Na+, pH 6.0, affected by the low water potential but without signs of suffering Na+ toxicity. At pH 8.0, tolerance seemed to be lower but it grew at 200 mm Na+, again without signs of Na+ toxicity. The resistance of Physcomitrella cells to the toxic effects of Na+ can be accounted for by their capacity to keep high K+:Na+ ratios and to extrude Na+ by a system that is not dependent on DeltapH. Physcomitrella expresses two P-type ATPases similar in sequence to fungal ENA-type Na+-ATPases. A functional study in yeast demonstrated that one of these ATPases, PpENA1, is an Na+-pump. We also found that P. patens has a plant-type SOS1 Na+/H+ antiporter. We discuss that Na+-ATPases existed in early land plants but that they were lost during the evolution of bryophytes to flowering plants.