Unit content
Photophosphorylation driven by the thylakoid proton gradient
Photophosphorylation is ATP formation driven by a proton-motive force generated by photosynthetic electron transfer across the thylakoid membrane.
Linear electron flow builds that gradient in several ways:
- water oxidation by PSII releases protons into the thylakoid lumen;
- electron transfer through the plastoquinone/cytochrome b6f region moves additional proton equivalents toward the lumen;
- reduction of NADP$^+$ to NADPH consumes reducing equivalents on the stromal side.
Together these processes make the lumen relatively proton-rich compared with the stroma.
ATP synthase uses proton return
Thylakoid ATP synthase allows protons to move downhill from lumen to stroma. That favorable electrochemical movement is coupled to
$$\mathrm{ADP+P_i\rightarrow ATP+H_2O}.$$
ATP is therefore produced on the stromal side, where it can be used by carbon-fixation reactions.
The energy-conversion chain is
light-driven electron flow
↓
H+ accumulation in thylakoid lumen
↓
proton-motive force
↓
H+ return through ATP synthase
↓
ATP made in stroma
Light does not phosphorylate ADP directly
A photon excites a photosystem; photosynthetic electron flow then builds an electrochemical gradient; ATP synthase converts that gradient into chemical free energy.
Thus
$$\text{light}\not\rightarrow\text{ATP directly}.$$
The membrane gradient is the intermediate energy store.
This is the same chemiosmotic principle used in respiration, but the upstream energy source differs: mitochondria commonly build the gradient using favorable oxidation of reduced fuel carriers, whereas chloroplasts use photon-driven electron transfer.