Coupled cyclic electron transport is assigned a role in the protection of leaves against photoinhibition in addition to its role in ATP synthesis. In leaves, as in reconstituted thylakoid systems, cyclic electron transport requires 'poising," i.e. availability of electrons at the reducing side of photosystem I (PSI) and the presence of some oxidized plastoquinone between photosystem 11 (PSII) and PSI. Under self-regulatory poising conditions that are established when carbon dioxide limits photosynthesis at high light intensities, and particularly when stomata are partially or fully closed as a result of water stress, coupled cyclic electron transport controls linear electron transport by helping to establish a proton gradient large enough to decrease PSII activity and electron flow to PSI. This brings electron donation by PSII, and electron consumption by available electron acceptors, into a balance in which PSI becomes more oxidized than it is during fast carbon assimilation. Avoidance of overreduction of the electron transport chain is a prerequisite for the efficient protection of the photosynthetic apparatus against photoinactivation.Despite several decades of intensive research, it is still not entirely clear how in leaves the reactions of the photosynthetic electron transport chain, which yield reductant (in the form of NADPH) and phosphate energy (in form of ATP), are geared to the reactions of stromal enzymes. These use this "assimilatory power' (1) from the chloroplast stroma to the intrathylakoid space of the chloroplasts, and that the free energy of the proton gradient thus formed is used for ATP synthesis, neither the stoichiometry of H+/e coupling nor that of ATP synthesis are known for certain. During linear electron transport from water to NADP, two protons are released inside the thylakoids per H20 oxidized by PSII and at least another two by the subsequent oxidation of the plastoquinol formed as a consequence of the transfer of the two electrons of water to plastoquinone. Unfortunately, it is still not quite clear how the Cyt b/f complex mediates oxidation of plastoquinol. If it passes electrons of this two-electron carrier straight on to PSI, which lifts them to the redox levels of Fd and NADP, the H+/e ratio of linear electron transport is 2. If only one electron from the plastoquinol is transferred to PSI and the other one is shuttled back to plastoquinone in the loop of a Mitchellian Q-cycle, the H+/e ratio is 3. According to Rich (19), Q-cycle coupling is obligatory. Others consider it to be facultative, with decreasing coupling efficiency at increasing light intensities (17, 18). Crowther and Hind (4) proposed a modified Q-cycle, with electron input from the reducing side of PSI.
CONSEQUENCES OF THE H+/ATP RATIO