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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).
This energy can be used in different ways, particularly in the synthesis of ATP by ATP synthases, promoting different types of secondary transport, for example, fueling flagellar movement and, in some cases, generating heat (Fig. 2). Proton Motive Force.