This study deals with effects of oxygen on the kinetics of P(700) photoinduced redox transitions and on induction transients of chlorophyll fluorescence in leaves of C(3) plants Hibiscus rosa-sinensis and Vicia faba. It is shown that the removal of oxygen from the leaf environment has a conspicuous effect on photosynthetic electron transport. Under anaerobic conditions, the concentration of oxidized P700 centers in continuous white light was substantially lower than under aerobic conditions. The deficiency of oxygen released non-photochemical quenching of chlorophyll fluorescence, thus indicating a decrease in the trans-thylakoid pH gradient (DeltapH). Quantitative analysis of experimental data within the framework of an original mathematical model has shown that the steady-state electron flux toward oxygen in Chinese hibiscus leaves makes up to approximately 40% of the total electron flow passing through photosystem 1 (PS1). The decrease in P700+ content under anaerobic conditions can be due to two causes: i) the retardation of electron outflow from PS1, and ii) the release of photosynthetic control (acceleration of electron flow from PS2 to P700+) owing to lower acidification of the intra-thylakoid space. At the same time, cyclic electron transport around PS1 was not stimulated in the oxygen-free medium, although such stimulation seemed likely in view of possible rearrangement of electron flows on the acceptor side of PS1. This conclusion stems from observations that the rates of P700+ reduction in DCMU-poisoned samples, both under aerobic and anaerobic conditions, were negligibly small compared to rates of electron flow from PS2 toward P700+ in untreated samples.
In this work, we have investigated the effects of atmospheric CO(2) and O(2) on induction events in Hibiscus rosa-sinensis leaves. These effects manifest themselves as multiphase kinetics of P(700) redox transitions and non-monotonous changes in chlorophyll fluorescence. Depletion of CO(2) and O(2) in air causes a decrease in linear electron flux (LEF) and dramatic lowering of P(700)(+) level. This is explained by the impediment to electron efflux from photosystem 1 (PS1) at low acceptor capacity. With the release of the acceptor deficit, the rate of LEF significantly increases. We have found that oxygen promotes the outflow of electrons from PS1, providing the rise of P(700)(+) level. The effect of oxygen as an alternative electron acceptor becomes apparent at low and ambient concentrations of atmospheric CO(2) < or = 0.06-0.07%). A decrease in LEF at low CO(2) is accompanied by a significant (about 3-fold) rise of non-photochemical quenching (NPQ) of chlorophyll fluorescence. Such an increase in NPQ can be explained by more significant acidification of the thylakoid lumen. This occurs due to lessening the proton flux through the ATP synthases caused by a decrease in the ATP consumption in the Bassham-Benson-Calvin (BBC) cycle. pH-dependent mechanisms of electron transport control have been described within the frames of our mathematical model. The model describes the reciprocal changes in LEF and NPQ and predicts the redistribution of electron fluxes on the acceptor side of PS1. In particular, the contribution of cyclic electron flow around PS1 (CEF1) and water-water cycle gradually decays during the induction phase. This result is consistent with experimental data indicating that under the steady-state conditions the contribution of CEF1 to photosynthetic electron transport in Hibiscus rosa-sinensis is insignificant (< or = 10%).
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