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The proton motive force arises from a combination of the concentration gradient of protons and a voltage gradient across the membrane (the protons are positively charged, so an imbalance of protons means that the side of the membrane with more protons has a positive charge relative to the side with fewer protons).
Proton-motive force, ATP synthetase, Coupling. I. Protonmotive force. A. Proof of proton gradient formation coupled to electron flow. Although scientists knew that electron transport led to the synthesis of ATP, for a. long time they did not know how.