Chloroplastic ATP synthase plays an important role in the regulation of proton motive force in fluctuating light.

The proton motive force (pmf) across the thylakoid membranes plays a key role for photosynthesis in fluctuating light. However, the mechanisms underlying the regulation of pmf in fluctuating light are not well known. In this study, we aimed to identify the roles of chloroplastic ATP synthase and cyclic electron flow (CEF) around photosystem I (PSI) in the regulation of the pmf in fluctuating light. To do this, we measured chlorophyll fluorescence, P700 parameters, and the electrochromic shift signal in the fluctuating light alternating between 918 (high light) and 89 (low light) μmol photons m-2  s-1 every 5 min. We found that the activity of chloroplastic ATP synthase (gH + ), pmf, CEF activity, non-photochemical quenching (NPQ), and the P700 redox state changed rapidly in fluctuating light. During transition from low to high light, the decreased gH + and the stimulation of CEF both contributed to the rapid formation of pmf, activating NPQ and optimizing the redox state of P700 in PSI. During the low-light phases, gH + rapidly increased and the pmf declined sharply, leading to the relaxation of NPQ and down-regulation of photosynthetic control. These findings indicate that in fluctuating light the gH + and CEF are finely regulated to modulate the pmf formation, avoiding the over-accumulation of reactive intermediates and maximizing energy use efficiency.