Role of cyclic electron transport around photosystem I in regulating proton motive force

In addition to ∆pH formed across the thylakoid membrane, membrane potential contributes to proton motive force (pmf) in chloroplasts. However, the regulation of photosynthetic electron transport is mediated solely by ∆pH. To assess the contribution of two cyclic electron transport pathways around photosystem I (one depending on PGR5/PGRL1 and one on NDH) to pmf formation, electrochromic shift (ECS) was analyzed in the Arabidopsis pgr5 mutant, NDH-defective mutants (ndhs and crr4-2), and their double mutants (ndhs pgr5 and crr4-2 pgr5). In pgr5, the size of the pmf, as represented by ECSt, was reduced by 30% to 47% compared with that in the wild type (WT). A gH+ parameter, which is considered to represent the activity of ATP synthase, was enhanced at high light intensities. However, gH+ recovered to its low-light levels after 20 min in the dark, implying that the elevation in gH+ is due to the disturbed regulation of ATP synthase rather than to photodamage. After long dark adaptation more than 2 h, gH+ was higher in pgr5 than in the WT. During induction of photosynthesis, gH+ was more rapidly elevated in pgr5 than that in the WT. Both results suggest that ATP synthase is not fully inactivated in the dark in pgr5. In the NDH-deficient mutants, ECSt was slightly but significantly lower than in the WT, whereas gH+ was not affected. In the double mutants, ECSt was even lower than in pgr5. These results suggest that both PGR5/PGRL1- and NDH-dependent pathways contribute to pmf formation, although to different extents. This article is part of a Special Issue entitled: Chloroplast Biogenesis.