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The vacuolar-type H + -pyrophosphatase (H + -PPase) catalyzes a coupled reaction of pyrophosphate (PPi) hydrolysis and active proton translocation across the tonoplast. Overexpression of H + -PPase improves growth in various plant species, and loss-of-function mutants ( fugu5 s) of H + -PPase in  Arabidopsis thaliana  have post-germinative developmental defects. Here, to further clarify the physiological significance of this important enzyme, we newly generated three varieties of H + -PPase overexpressing lines with different levels of activity that we analyzed together with the loss-of-function mutant  fugu5-3 . The H + -PPase overexpressors exhibited enhanced activity of H + -PPase during vegetative growth, but no change in the activity of vacuolar H + -ATPase. Overexpressors with high enzymatic activity grew more vigorously with fresh weight increased by more than 24 and 44%, compared to the wild type and  fugu5-3 , respectively. Consistently, the overexpressors had larger rosette leaves and nearly 30% more cells in leaves than the wild type. When uncoupling mutated variants of H + -PPase, that could hydrolyze PPi but could not translocate protons, were introduced into the  fugu5-3  mutant background, shoot growth defects recovered to the same levels as when a normal H + -PPase was introduced. Taken together, our findings clearly demonstrate that additional expression of H + -PPase improves plant growth by increasing cell number, predominantly as a consequence of the PPi-hydrolyzing activity of the enzyme.

frontiers in plant science

Contribution of PPi-Hydrolyzing Function of Vacuolar H+-Pyrophosphatase in Vegetative Growth of Arabidopsis: Evidenced by Expression of Uncoupling Mutated Enzymes