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Cell organelles maintain their optimum pH through specialized mechanisms involving proton pumps, ion exchangers, buffering systems, and compartmentalization. Lysosomes, for example, require an acidic environment (pH ~4.5–5) for the activity of hydrolytic enzymes. This acidic pH is maintained by vacuolar-type H⁺-ATPases (V-ATPases) that actively transport protons (H⁺ ions) from the _______ ____ ____ _____ _______ __________ ________.
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