Alternative Photosystem I-driven Electron Transport Routes: Mechanisms and Functions

In addition to the linear electron transport, several alternative Photosystem I-driven (PS I) electron pathways recycle the electrons to the intersystem electron carriers mediated by either ferredoxin:NADPH reductase, NAD(P)H dehydrogenase, or putative ferredoxin:plastoquinone reductase. The following functions have been proposed for these pathways: adjustment of ATP/NADPH ratio required for CO(2) fixation, generation of the proton gradient for the down-regulation of Photosystem II (PS II), and ATP supply the active transport of inorganic carbon in algal cells. Unlike ferredoxin-dependent cyclic electron transport, the pathways supported by NAD(P)H can function in the dark and are likely involved in chlororespiratory-dependent energization of the thylakoid membrane. This energization may support carotenoid biosynthesis and/or maintain thylakoid ATPase in active state. Active operation of ferredoxin-dependent cyclic electron transport requires moderate reduction of both the intersystem electron carriers and the acceptor side of PS I, whereas the rate of NAD(P)H-dependent pathways under light depends largely on NAD(P)H accumulation in the stroma. Environmental stresses such as photoinhibition, high temperatures, drought, or high salinity stimulated the activity of alternative PS I-driven electron transport pathways. Thus, the energetic and regulatory functions of PS I-driven pathways must be an integral part of photosynthetic organisms and provides additional flexibility to environmental stress.