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Tonoplast transporters, including proton pumps and secondary transporters, are essential for plant cell function and for quality formation of fleshy fruits and omamentals.Vacuolar transport of anthocyanins, malate and other metabolites is directly or indirectly dependent on the H+-pumping activities of vacuolar H+-ATPase (VHA) and/or pyrophosphatase (VHP), but how these proton pumps are regulated in modulating vacuolar transport is largely unknown.