Oxygen is a natural acceptor of electrons in the respiratory pathway of aerobic organisms and in many other biochemical reactions. Aerobic metabolism is always associated with the formation of reactive oxygen species (ROS). ROS may damage biomolecules but are also involved in regulatory functions of photosynthetic organisms. This review presents the main properties of ROS, the formation of ROS in the photosynthetic electron transport chain and in the stroma of chloroplasts, and ROS scavenging systems of thylakoid membrane and stroma. Effects of ROS on the photosynthetic apparatus and their roles in redox signaling are discussed.

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Chlorophylls are degraded and flavonoids synthesized during autumn senescence of deciduous trees. In the present study, chlorophyll and flavonol contents of individual leaves were monitored non-destructively throughout the autumn. Loss of chlorophyll and synthesis of flavonols were not gradual. Instead, each leaf maintained steady chlorophyll content until rapid chlorophyll degradation, accompanied by flavonol synthesis, was triggered. In ~1 week, the leaf turns yellow and falls. The pattern was similar in birch (Betula pendula), maple (Acer platanoides) and bird cherry (Prunus padus); in rowan (Sorbus aucuparia), very slow gradual chlorophyll degradation occurred on top of the main pattern.

Nitrogen deficiency diminishes consumption of photosynthates in anabolic metabolism. We studied adjustments of the photosynthetic machinery in nitrogen-deficient bean plants and found four phenomena. First, the number of chloroplasts per cell decreased. Chloroplasts of nitrogen starved leaves contained less pigments than those of control leaves, but the in vitro activities of light reactions did not change when measured on chlorophyll basis. Second, nitrogen deficiency induced cyclic electron transfer. The amounts of Rubisco and ferredoxin-NADP(+) reductase decreased in nitrogen starved plants. Low activities of these enzymes are expected to lead to increase in reduction of oxygen by photosystem I. However, diaminobenzidine staining did not reveal hydrogen peroxide production in nitrogen starved plants. Measurements of far-red-light-induced redox changes of the primary donor of photosystem I suggested that instead of producing oxygen radicals, nitrogen starved plants develop a high activity of cyclic electron transport that competes with oxygen for electrons. Nitrogen starvation led to decrease in photochemical quenching and increase in non-photochemical quenching, indicating that cyclic electron transport reduces the plastoquinone pool and acidifies the lumen. A third effect is redistribution of excitation energy between the photosystems in favor of photosystem I. Thus, thylakoids of nitrogen starved plants appeared to be locked in state 2, which further protects photosystem II by decreasing its absorption cross-section. As a fourth response, the proportion of non-Q(B)-reducing photosystem II reaction centers increased and the redox potential of the Q(B)/Q(B)(-) pair decreased by 25 mV in a fraction of photosystem II centers of nitrogen starved plants.

The plastoquinone (PQ) pool mediates electron flow and regulates photoacclimation in plants. Here we report the action spectrum of the redox state of the PQ pool in Arabidopsis thaliana, showing that 470-500, 560 or 650-660 nm light favors Photosystem II (PSII) and reduces the PQ pool, whereas 420-440, 520 or 690 nm light favors Photosystem I (PSI) and oxidizes PQ. These data were used to construct a model predicting the redox state of PQ from the spectrum of any polychromatic light source. Moderate reduction of the PQ pool induced transition to light state 2, whereas state 1 required highly oxidized PQ. In low-intensity PSI light, PQ was more oxidized than in darkness and became gradually reduced with light intensity, while weak PSII light strongly reduced PQ. Natural sunlight was found to favor PSI, which enables plants to use the redox state of the PQ pool as a measure of light intensity.

Photoinhibition is light-induced inactivation of PSII, and action spectrum measurements have shown that UV light causes photoinhibition much more efficiently than visible light. In the present study, we quantified the contribution of the UV part of sunlight in photoinhibition of PSII in leaves. Greenhouse-grown pumpkin leaves were pretreated with lincomycin to block the repair of photoinhibited PSII, and exposed to sunlight behind a UV-permeable or UV-blocking filter. Oxygen evolution and Chl fluorescence measurements showed that photoinhibition proceeds 35% more slowly under the UV-blocking than under the UV-permeable filter. Experiments with a filter that blocks UV-B but transmits UV-A and visible light revealed that UV-A light is almost fully responsible for the UV effect. The difference between leaves illuminated through a UV-blocking and UV-transparent filter disappeared when leaves of field-grown pumpkin plants were used. Thylakoids isolated from field-grown and greenhouse-grown plants were equally sensitive to UV light, and measurements of UV-induced fluorescence from leaves indicated that the protection of the field-grown plants was caused by substances that block the passage of UV light to the chloroplasts. Thus, the UV part of sunlight, especially the UV-A part, is potentially highly important in photoinhibition of PSII but the UV-screening compounds of plant leaves may offer almost complete protection against UV-induced photoinhibition.

Singlet oxygen ( 1 O 2 ) is a harmful species that functions also as a signaling molecule. In chloroplasts, 1 O 2 is produced via charge recombination reactions in photosystem II, but which recombination pathway(s) produce triplet Chl and 1 O 2 remains open. Furthermore, the role of 1 O 2 in photoinhibition is not clear.We compared temperature dependences of 1 O 2 production, photoinhibition, and recombination pathways.1 O 2 production by pumpkin thylakoids increased from −2 to +35°C, ruling out recombination of the primary charge pair as a main contributor. S 2 Q A − or S 2 Q B − recombination pathways, in turn, had too steep temperature dependences. Instead, the temperature dependence of 1 O 2 production matched that of misses (failures of the oxygen (O 2 ) evolving complex to advance an S-state). Photoinhibition in vitro and in vivo (also in Synechocystis), and in the presence or absence of O 2 , had the same temperature dependence, but ultraviolet (UV)radiation-caused photoinhibition showed a weaker temperature response.We suggest that the miss-associated recombination of P 680 + Q A − is the main producer of 1 O 2 . Our results indicate three parallel photoinhibition mechanisms. The manganese mechanism dominates in UV radiation but also functions in white light. Mechanisms that depend on light absorption by Chls, having 1 O 2 or long-lived P 680 + as damaging agents, dominate in red light.

Recombination of the primary radical pair of photosystem II (PSII) of photosynthesis may produce the triplet state of the primary donor of PSII. Triplet formation is potentially harmful because chlorophyll triplets can react with molecular oxygen to produce the reactive singlet oxygen ( 1 O 2 ). The yield of 1 O 2 is expected to be directly proportional to the triplet yield and the triplet yield of charge recombination can be lowered with a magnetic field of 100-300 mT. In this study, we illuminated intact pumpkin leaves with strong light in the presence and absence of a magnetic field and found that the magnetic field protects against photoinhibition of PSII. The result suggests that radical pair recombination is responsible for significant part of 1 O 2 production in the chloroplast. The magnetic field effect vanished if leaves were illuminated in the presence of lincomycin, an inhibitor of chloroplast protein synthesis, or if isolated thylakoid membranes were exposed to light. These data, in turn, indicate that 1 O 2 produced by the recombination of the primary charge pair is not directly involved in photoinactivation of PSII but instead damages PSII by inhibiting the repair of photoinhibited PSII. We also found that an Arabidopsis thaliana mutant lacking α-tocopherol, a scavenger of 1 O 2 , is more sensitive to photoinhibition than the wild-type in the absence but not in the presence of lincomycin, confirming that the target of 1 O 2 is the repair mechanism.