Chloroplast Electron Transport Chain Research Articles

Plastoquinol Oxidation: Rate-Limiting Stage in the Electron Transport Chain of Chloroplasts.

This work is devoted to theoretical study of functioning of the cytochrome (Cyt) b6f complex (plastoquinol:plastocyanin oxidoreductase) of the electron transport chain (ETC) in oxygenic photosynthesis. A composition of the chloroplast ETC and molecular mechanisms of functioning of the Cyt b6f complex, which stands between photosystems II and I (PSII and PSI), are briefly reviewed. The Cyt b6f complex oxidizes plastoquinol (PQH2) molecules formed in PSII, and reduces plastocyanin, which serves as an electron donor to PSI. PQH2 oxidation is the rate-limiting step in the chain of electron transfer processes between PSII and PSI. Using the density functional theory (DFT) method, we have analyzed the two-electron (bifurcated) oxidation of PQH2 in the catalytic center Qo of the Cyt b6f complex. Results of DFT calculations are consistent with the fact that the first step of PQH2 oxidation, electron transfer to the Fe2S2 cluster of the iron-sulfur protein (ISP), is an endergonic (energy-accepting) process (ΔE ≈ 15 kJ·mol-1) that can limit turnover of the Cyt b6f complex. The second stage of bifurcated oxidation of PQH2 - electron transfer from semiquinone (PQH•, formed after the first step of PQH2 oxidation) to heme b6L - is the exergonic (energy-donating) process (ΔE < 0). DFT modeling of this stage revealed that semiquinone oxidation should accelerate after the PQH• radical shift towards the heme b6L (an electron acceptor) and the carboxy group of Glu78 (a proton acceptor). The data obtained are discussed within the framework of the Mitchell Q-cycle model describing PQH2 oxidation at the Qo site of the Cyt b6f complex.

Potential of Asparagopsis armata as a Biopesticide for Weed Control under an Invasive Seaweed Circular-Economy Framework.

Simple SummaryThe invasive seaweed Asparagopsis armata has the potential to be used as a biopesticide. The application of its exudate shows severe impacts on energetic and carotenoid metabolism and induces significant oxidative stress in a model weed. This points to the potential use of this macroalga as a resource for a biopesticide cocktail, for sustainable and eco-friendly weed control and as a substitute for the chemical pesticides widely used nowadays.Marine macroalgae have been increasingly targeted as a source of bioactive compounds to be used in several areas, such as biopesticides. When harvesting invasive species, such as Asparagopsis armata, for this purpose, there is a two-folded opportunity: acquiring these biomolecules from a low-cost resource and controlling its spreading and impacts. The secondary metabolites in this seaweed’s exudate have been shown to significantly impact the physiology of species in the ecosystems where it invades, indicating a possible biocidal potential. Considering this in the present work, an A. armata exudate cocktail was applied in the model weed Thellungiella halophila to evaluate its physiological impact and mode of action, addressing its potential use as a natural biocide. A. armata greatly affected the test plants’ physiology, namely, their photochemical energy transduction pathway (impairing light-harvesting and chemical energy production throughout the chloroplast electron transport chain), carotenoid metabolism and oxidative stress. These mechanisms of action are similar to the ones triggered when using the common chemical pesticides, highlighting the potential of the A. armata exudate cocktail as an eco-friendly biopesticide.

The Assembly of Super-Complexes in the Plant Chloroplast.

Increasing evidence has revealed that the enzymes of several biological pathways assemble into larger supramolecular structures called super-complexes. Indeed, those such as association of the mitochondrial respiratory chain complexes play an essential role in respiratory activity and promote metabolic fitness. Dynamically assembled super-complexes are able to alternate between participating in large complexes and existing in a free state. However, the functional significance of the super-complexes is not entirely clear. It has been proposed that the organization of respiratory enzymes into super-complexes could reduce oxidative damage and increase metabolism efficiency. There are several protein complexes that have been revealed in the plant chloroplast, yet little research has been focused on the formation of super-complexes in this organelle. The photosystem I and light-harvesting complex I super-complex’s structure suggests that energy absorbed by light-harvesting complex I could be efficiently transferred to the PSI core by avoiding concentration quenching. Here, we will discuss in detail core complexes of photosystem I and II, the chloroplast ATPase the chloroplast electron transport chain, the Calvin–Benson cycle and a plastid localized purinosome. In addition, we will also describe the methods to identify these complexes.

Cooperation of chloroplast ascorbate peroxidases and proton gradient regulation 5 is critical for protecting Arabidopsis plants from photo-oxidative stress.

High-light (HL) stress enhances the production of H2 O2 from the photosynthetic electron transport chain in chloroplasts, potentially causing photo-oxidative damage. Although stromal and thylakoid membrane-bound ascorbate peroxidases (sAPX and tAPX, respectively) are major H2 O2 -scavenging enzymes in chloroplasts, their knockout mutants do not exhibit a visible phenotype under HL stress. Trans-thylakoid proton gradient (∆pH)-dependent mechanisms exist for controlling H2 O2 production from photosynthesis, such as thermal dissipation of light energy and downregulation of electron transfer between photosystems II and I, and these may compensate for the lack of APXs. To test this hypothesis, we focused on a proton gradient regulation 5 (pgr5) mutant, wherein both ∆pH-dependent mechanisms are impaired, and an Arabidopsis sapx tapx double mutant was crossed with the pgr5 single mutant. The sapx tapx pgr5 triple mutant exhibited extreme sensitivity to HL compared with its parental lines. This phenotype was consistent with cellular redox perturbations and enhanced expression of many oxidative stress-responsive genes. These findings demonstrate that the PGR5-dependent mechanisms compensate for chloroplast APXs, and vice versa. An intriguing finding was that the failure of induction of non-photochemical quenching in pgr5 (because of the limitation in ∆pH formation) was partially recovered in sapx tapx pgr5. Further genetic studies suggested that this recovery was dependent on the NADH dehydrogenase-like complex-dependent pathway for cyclic electron flow around photosystem I. Together with data from the sapx tapx npq4 mutant, we discuss the interrelationship between APXs and ∆pH-dependent mechanisms under HL stress.

Integrated transcriptome and metabolome analysis reveals the physiological and molecular responses of allotetraploid rapeseed to ammonium toxicity

Ammonium (NH4+) and nitrate (NO3−) are the main nitrogen (N) sources accessible to plants. However, exclusive NH4+ may causes toxicity to crops. Transcriptomics in conjunction with metabolomics analysis in allotetraploid rapeseed under NH4+ stress showed that NH4+ taken up by roots was mainly transferred to leaves for assimilation. The concentrations of most amino acids, glucose, and glucose-6-phosphate increased whilst malate, succinate, and α-ketoglutarate concentrations decreased in NH4+-treated leaves. Moreover, NH4+ toxicity down-regulated genes related to photosynthesis and impaired the electron transport chain in chloroplasts. The expression of genes related to H2O2 decomposition, as well as the concentrations of monodehydro-ascorbic acid and ascorbic acid, reduced after exposure to NH4+-N, accompanied by higher O2- and H2O2 accumulation. Taken together, our results indicated that the excessive accumulation of NH4+ in plants caused an imbalance of carbon and N metabolism, impaired photosystems and generated reactive oxygen species, which resulted in toxicity to rapeseed.

The flexibility of metabolic interactions between chloroplasts and mitochondria in Nicotiana tabacum leaf.

To examine the effect of mitochondrial function on photosynthesis, wild-type and transgenic Nicotiana tabacum with varying amounts of alternative oxidase (AOX) were treated with different respiratory inhibitors. Initially, each inhibitor increased the reduction state of the chloroplast electron transport chain, most severely in AOX knockdowns and least severely in AOX overexpressors. This indicated that the mitochondrion was a necessary sink for photo-generated reductant, contributing to the 'P700 oxidation capacity' of photosystem I. Initially, the Complex III inhibitor myxothiazol and the mitochondrial ATP synthase inhibitor oligomycin caused an increase in photosystem II regulated non-photochemical quenching not evident with the Complex III inhibitor antimycin A (AA). This indicated that the increased quenching depended upon AA-sensitive cyclic electron transport (CET). Following 12h with oligomycin, the reduction state of the chloroplast electron transport chain recovered in all plant lines. Recovery was associated with large increases in the protein amount of chloroplast ATP synthase and mitochondrial uncoupling protein. This increased the capacity for photophosphorylation in the absence of oxidative phosphorylation and enabled the mitochondrion to act again as a sink for photo-generated reductant. Comparing the AA and myxothiazol treatments at 12h showed that CET optimized photosystem I quantum yield, depending upon the P700 oxidation capacity. When this capacity was too high, CET drew electrons away from other sinks, moderating the P700+ amount. When P700 oxidation capacity was too low, CET acted as an electron overflow, moderating the amount of reduced P700. This study reveals flexible chloroplast-mitochondrion interactions able to overcome lesions in energy metabolism.

Electrochemical energy for living systems

Plastoquinone and ubiquinone play essential roles in the electron transport chains of chloroplasts and mitochondria by coupling electron transport to the transfer of protons across membranes. The energy of the resulting proton gradient is used to synthesize adenosine triphosphate (ATP), the energy currency of all life. How did quinones first become involved in this process? We have detected several quinone compounds in carbonaceous meteorites that can transport protons in a liposome model system. It is possible that such compounds were available to transport protons in primitive versions of electron transport in early life.

Ecological Significance of the Interaction of Photosynthesis Light and Dark Processes

The kinetics of 14C incorporation into glycolate was studied after changing the export of photosynthetic products from the leaf. It has been shown that the ribulose-bisphosphate-oxygenase pathway of glycolate formation works in the stationary state of the plant. An excess of photosyntates or a decrease in the amount of light primary products, as well as nitrates in the leaves, immediately turns on the transketolase pathway of glycolate formation. In this case, part of the oxygen formed in the photochemical reactions of chloroplasts ceases to be released from the leaf. After oxygen receives an electron from ferredoxin in the electron transport chain of chloroplasts, it starts (through photorespiration) the formation of non-carbohydrate photosyntates and metabolic processes in the cytoplasm. It was concluded that the main function of photorespiration in the regulation of photosynthesis is maintaining a balance between light and dark processes of photosynthesis on change of living conditions.

Influence of elevated temperature on electron flows in chloroplasts of barley

The efficiency of electron carriers in thylakoid membranes untreated and exposed to heat 7-day-old barley seedlings was evaluated with PAM fluorescence. Darkness–light transitional states in chloroplasts after heat exposure are studied. Thermoinduced changes in linear and cyclic electron transport chain of chloroplasts are revealed. The activation of NADPH-dependent electron flux after exposure to elevated temperatures is shown. We assumed that ΔрН of thylakoid membranes employed the regulatory role in the distribution of electron flows and the adaptation of the photosynthetic apparatus to stressful effects.

Efficient Photosynthetic Functioning of Arabidopsis thaliana Through Electron Dissipation in Chloroplasts and Electron Export to Mitochondria Under Ammonium Nutrition.

An improvement in photosynthetic rate promotes the growth of crop plants. The sink-regulation of photosynthesis is crucial in optimizing nitrogen fixation and integrating it with carbon balance. Studies on these processes are essential in understanding growth inhibition in plants with ammonium () syndrome. Hence, we sought to investigate the effects of using nitrogen sources with different states of reduction (during assimilation of versus ) on the photosynthetic performance of Arabidopsis thaliana. Our results demonstrated that photosynthetic functioning during long-term nutrition was not disturbed and that no indication of photoinhibition of PSII was detected, revealing the robustness of the photosynthetic apparatus during stressful conditions. Based on our findings, we propose multiple strategies to sustain photosynthetic activity during limited reductant utilization for assimilation. One mechanism to prevent chloroplast electron transport chain overreduction during nutrition is for cyclic electron flow together with plastid terminal oxidase activity. Moreover, redox state in chloroplasts was optimized by a dedicated type II NAD(P)H dehydrogenase. In order to reduce the amount of energy that reaches the photosynthetic reaction centers and to facilitate photosynthetic protection during nutrition, non-photochemical quenching (NPQ) and ample xanthophyll cycle pigments efficiently dissipate excess excitation. Additionally, high redox load may be dissipated in other metabolic reactions outside of chloroplasts due to the direct export of nucleotides through the malate/oxaloacetate valve. Mitochondrial alternative pathways can downstream support the overreduction of chloroplasts. This mechanism correlated with the improved growth of A. thaliana with the overexpression of the alternative oxidase 1a (AOX1a) during nutrition. Most remarkably, our findings demonstrated the capacity of chloroplasts to tolerate syndrome instead of providing redox poise to the cells.

Involvement of the chloroplast gene ferredoxin 1 in multiple responses of Nicotiana benthamiana to Potato virus X infection.

The chloroplast protein ferredoxin 1 (FD1), with roles in the chloroplast electron transport chain, is known to interact with the coat proteins (CPs) of Tomato mosaic virus and Cucumber mosaic virus. However, our understanding of the roles of FD1 in virus infection remains limited. Here, we report that the Potato virus X (PVX) p25 protein interacts with FD1, whose mRNA and protein levels are reduced by PVX infection or by transient expression of p25. Silencing of FD1 by Tobacco rattle virus-based virus-induced gene silencing (VIGS) promoted the local and systemic infection of plants by PVX. Use of a drop-and-see (DANS) assay and callose staining revealed that the permeability of plasmodesmata (PDs) was increased in FD1-silenced plants together with a consistently reduced level of PD callose deposition. After FD1 silencing, quantitative reverse transcription–real-time PCR (qRT–PCR) analysis and LC-MS revealed these plants to have a low accumulation of the phytohormones abscisic acid (ABA) and salicylic acid (SA), which contributed to the decreased callose deposition at PDs. Overexpression of FD1 in transgenic plants manifested resistance to PVX infection, but the contents of ABA and SA, and the PD callose deposition were not increased in transgenic plants. Overexpression of FD1 interfered with the RNA silencing suppressor function of p25. These results demonstrate that interfering with FD1 function causes abnormal plant hormone-mediated antiviral processes and thus enhances PVX infection.

Reactive Oxygen Species, Photosynthesis, and Environment in the Regulation of Stomata.

Stomata sense the intercellular carbon dioxide (CO2) concentration (Ci) and water availability under changing environmental conditions and adjust their apertures to maintain optimal cellular conditions for photosynthesis. Stomatal movements are regulated by a complex network of signaling cascades where reactive oxygen species (ROS) play a key role as signaling molecules. Recent Advances: Recent research has uncovered several new signaling components involved in CO2- and abscisic acid-triggered guard cell signaling pathways. In addition, we are beginning to understand the complex interactions between different signaling pathways. Plants close their stomata in reaction to stress conditions, such as drought, and the subsequent decrease in Ci leads to ROS production through photorespiration and over-reduction of the chloroplast electron transport chain. This reduces plant growth and thus drought may cause severe yield losses for agriculture especially in arid areas. The focus of future research should be drawn toward understanding the interplay between various signaling pathways and how ROS, redox, and hormonal balance changes in space and time. Translating this knowledge from model species to crop plants will help in the development of new drought-resistant crop species with high yields.

The impact of photosynthesis on initiation of leaf senescence.

Senescence is the last stage of leaf development preceding the death of the organ, and it is important for nutrient remobilization and for feeding sink tissues. There are many reports on leaf senescence, but the mechanisms initiating leaf senescence are still poorly understood. Leaf senescence is affected by many environmental factors and seems to vary in different species and even varieties of plants, which makes it difficult to generalize the mechanism. Here, we give an overview on studies reporting about alterations in the composition of the photosynthetic electron transport chain in chloroplasts during senescence. We hypothesize that alternative electron flow and related generation of the proton motive force required for ATP synthesis become increasingly important during progression of senescence. We address the generation of reactive oxygen species (ROS) in chloroplasts in the initiation of senescence, retrograde signaling from the chloroplast to the nucleus and ROS-dependent signaling associated with leaf senescence. Finally, a few ideas for increasing crop yields by increasing the chloroplast lifespan are presented.

Analysis of Correlations between the Indexes of Light-Dependent Reactions of Photosynthesis and the Photochemical Reflectance Index (PRI) in Pea Leaves under Short-Term Illumination

The photochemical reflectance index (PRI) is related to the conversion of violoxanthin into zeaxanthin in the xanthophyll cycle and thereby reflects the activity of electron-transport chain in chloroplasts. The PRI is recorded by a relatively simple and non-invasive technique; therefore it can be used in a perspective approach for the remote monitoring of photosynthetic processes and detection of photosynthetic stress. However, correlation coefficients between PRI and photosynthetic parameters vary widely in different studies; one of the possible causes of such variability under natural conditions may be the changes in illumination and the development of transient processes in the photosynthetic apparatus. In the present study, the influence of the duration of illumination (a minute interval) on the relationship between parameters of light-dependent reactions of photosynthesis and the PRI has been investigated. The photosynthetic parameters and PRI were determined in pea leaves. A high negative correlation has been found between changes of PRI induced by illumination and changes in the quantum yields of the photochemical reactions of photosystems I and II; the PRI changes did not correlate with changes of non-photochemical quenching. There was a positive correlation between the quantum yields of photochemical reactions and PRI averaged over the entire time range and a negative correlation in the case of non-photochemical quenching and PRI. The absolute value of the correlation coefficient of PRI with the parameters of the light-dependent reactions of photosynthesis increased with increasing duration of illumination. Thus, the correlation of PRI with the parameters of the light-dependent reactions of photosynthesis in a pea leaf may depend on the duration of illumination, at least in a minute interval.

The mitochondrial alternative oxidase from Chlamydomonas reinhardtii enables survival in high light

Photosynthetic organisms often experience extreme light conditions that can cause hyper-reduction of the chloroplast electron transport chain, resulting in oxidative damage. Accumulating evidence suggests that mitochondrial respiration and chloroplast photosynthesis are coupled when cells are absorbing high levels of excitation energy. This coupling helps protect the cells from hyper-reduction of photosynthetic electron carriers and diminishes the production of reactive oxygen species (ROS). To examine this cooperative protection, here we characterized Chlamydomonas reinhardtii mutants lacking the mitochondrial alternative terminal respiratory oxidases, CrAOX1 and CrAOX2. Using fluorescent fusion proteins, we experimentally demonstrated that both enzymes localize to mitochondria. We also observed that the mutant strains were more sensitive than WT cells to high light under mixotrophic and photoautotrophic conditions, with the aox1 strain being more sensitive than aox2 Additionally, the lack of CrAOX1 increased ROS accumulation, especially in very high light, and damaged the photosynthetic machinery, ultimately resulting in cell death. These findings indicate that the Chlamydomonas AOX proteins can participate in acclimation of C. reinhardtii cells to excess absorbed light energy. They suggest that when photosynthetic electron carriers are highly reduced, a chloroplast-mitochondria coupling allows safe dissipation of photosynthetically derived electrons via the reduction of O2 through AOX (especially AOX1)-dependent mitochondrial respiration.

Gene Expression Profiling Shows That NbFDN1 Is Involved in Modulating the Hypersensitive Response-Like Cell Death Induced by the Oat dwarf virus RepA Protein

In this study, we used high-throughput deep nucleotide sequencing to characterize the global transcriptional response of Nicotiana benthamiana plants to transient expression of the RepA protein from Oat dwarf virus (ODV). We identified 7,878 significantly differentially expressed genes (DEG) that mapped to 125 pathways, suggesting that comprehensive networks are involved in regulation of RepA-induced cell death. Of the 202 DEG associated with photosynthesis, expression of 195 was found to be downregulated, indicating a significant inhibition of photosynthesis in response to RepA expression, which is associated with chloroplast disruption and physiological changes. We focused our analysis on NbFDN1, a member of the ferredoxin protein family that participates in the chloroplast electron transport chain performing oxygenic photosynthesis, which was identified to directly interact with NbTsip1. We separately knocked down the expression of NbFDN1 and NbTsip1 using virus-induced gene silencing, and found that NbFDN1 silencing speeded up the development of RepA-induced cell death, unlike NbTsip1 silencing, which showed an opposite effect on RepA-induced response. Further study showed increased H2O2 accumulation and a negative correlation between the transcripts of NbFDN1 and NbTsip1 in NbFDN1-silenced plants. Hence, we speculate that NbFDN1 has an effect on RepA-induced hypersensitive response-like response by modulating NbTsip1 transcription as well as H2O2 production.

Climate Change and the Impact of Greenhouse Gasses: CO2 and NO, Friends and Foes of Plant Oxidative Stress.

Here, we review information on how plants face redox imbalance caused by climate change, and focus on the role of nitric oxide (NO) in this response. Life on Earth is possible thanks to greenhouse effect. Without it, temperature on Earth’s surface would be around -19°C, instead of the current average of 14°C. Greenhouse effect is produced by greenhouse gasses (GHG) like water vapor, carbon dioxide (CO2), methane (CH4), nitrous oxides (NxO) and ozone (O3). GHG have natural and anthropogenic origin. However, increasing GHG provokes extreme climate changes such as floods, droughts and heat, which induce reactive oxygen species (ROS) and oxidative stress in plants. The main sources of ROS in stress conditions are: augmented photorespiration, NADPH oxidase (NOX) activity, β-oxidation of fatty acids and disorders in the electron transport chains of mitochondria and chloroplasts. Plants have developed an antioxidant machinery that includes the activity of ROS detoxifying enzymes [e.g., superoxide dismutase (SOD), ascorbate peroxidase (APX), catalase (CAT), glutathione peroxidase (GPX), and peroxiredoxin (PRX)], as well as antioxidant molecules such as ascorbic acid (ASC) and glutathione (GSH) that are present in almost all subcellular compartments. CO2 and NO help to maintain the redox equilibrium. Higher CO2 concentrations increase the photosynthesis through the CO2-unsaturated Rubisco activity. But Rubisco photorespiration and NOX activities could also augment ROS production. NO regulate the ROS concentration preserving balance among ROS, GSH, GSNO, and ASC. When ROS are in huge concentration, NO induces transcription and activity of SOD, APX, and CAT. However, when ROS are necessary (e.g., for pathogen resistance), NO may inhibit APX, CAT, and NOX activity by the S-nitrosylation of cysteine residues, favoring cell death. NO also regulates GSH concentration in several ways. NO may react with GSH to form GSNO, the NO cell reservoir and main source of S-nitrosylation. GSNO could be decomposed by the GSNO reductase (GSNOR) to GSSG which, in turn, is reduced to GSH by glutathione reductase (GR). GSNOR may be also inhibited by S-nitrosylation and GR activated by NO. In conclusion, NO plays a central role in the tolerance of plants to climate change.

The Cytochrome b 6 f Complex: Biophysical Aspects of Its Functioning in Chloroplasts.

This chapter presents an overview of structural properties of the cytochrome (Cyt) b 6 f complex and its functioning in chloroplasts. The Cyt b 6 f complex stands at the crossroad of photosynthetic electron transport pathways, providing connectivity between Photosystem (PSI) and Photosysten II (PSII) and pumping protons across the membrane into the thylakoid lumen. After a brief review of the chloroplast electron transport chain, the consideration is focused on the structural organization of the Cyt b 6 f complex and its interaction with plastoquinol (PQH2, reduced form of plastoquinone), a mediator of electron transfer from PSII to the Cyt b 6 f complex. The processes of PQH2 oxidation by the Cyt b 6 f complex have been considered within the framework of the Mitchell's Q-cycle. The overall rate of the intersystem electron transport is determined by PQH2 turnover at the quinone-binding site Qo of the Cyt b 6 f complex. The rate of PQH2 oxidation is controlled by the intrathylakoid pHin, which value determines the protonation/deprotonation events in the Qo-center. Two other regulatory mechanisms associated with the Cyt b 6 f complex are briefly overviewed: (i) redistribution of electron fluxes between alternative (linear and cyclic) pathways, and (ii) "state transitions" related to redistribution of solar energy between PSI and PSII.

Electron Transfer through the Acceptor Side of Photosystem I: Interaction with Exogenous Acceptors and Molecular Oxygen.

This review considers the state-of-the-art on mechanisms and alternative pathways of electron transfer in photosynthetic electron transport chains of chloroplasts and cyanobacteria. The mechanisms of electron transport control between photosystems (PS) I and II and the Calvin-Benson cycle are considered. The redistribution of electron fluxes between the noncyclic, cyclic, and pseudocyclic pathways plays an important role in the regulation of photosynthesis. Mathematical modeling of light-induced electron transport processes is considered. Particular attention is given to the electron transfer reactions on the acceptor side of PS I and to interactions of PS I with exogenous acceptors, including molecular oxygen. A kinetic model of PS I and its interaction with exogenous electron acceptors has been developed. This model is based on experimental kinetics of charge recombination in isolated PS I. Kinetic and thermodynamic parameters of the electron transfer reactions in PS I are scrutinized. The free energies of electron transfer between quinone acceptors A1A/A1B in the symmetric redox cofactor branches of PS I and iron-sulfur clusters FX, FA, and FB have been estimated. The second-order rate constants of electron transfer from PS I to external acceptors have been determined. The data suggest that byproduct formation of superoxide radical in PS I due to the reduction of molecular oxygen in the A1 site (Mehler reaction) can exceed 0.3% of the total electron flux in PS I.

Influence of the variation potential on photosynthetic flows of light energy and electrons in pea.

Local damage (mainly burning, heating, and mechanical wounding) induces propagation of electrical signals, namely, variation potentials, which are important signals during the life of plants that regulate different physiological processes, including photosynthesis. It is known that the variation potential decreases the rate of CO2 assimilation by the Calvin-Benson cycle; however, its influence on light reactions has been poorly investigated. The aim of our work was to investigate the influence of the variation potential on the light energy flow that is absorbed, trapped and dissipated per active reaction centre in photosystem II and on the flow of electrons through the chloroplast electron transport chain. We analysed chlorophyll fluorescence in pea leaves using JIP-test and PAM-fluorometry; we also investigated delayed fluorescence. The electrical signals were registered using extracellular electrodes. We showed that the burning-induced variation potential stimulated a nonphotochemical loss of energy in photosystem II under dark conditions. It was also shown that the variation potential gradually increased the flow of light energy absorbed, trapped and dissipated by photosystem II. These changes were likely caused by an increase in the fraction of absorbed light distributed to photosystem II. In addition, the variation potential induced a transient increase in electron flow through the photosynthetic electron transport chain. Some probable mechanisms for the influence of the variation potential on the light reactions of photosynthesis (including the potential role of intracellular pH decrease) are discussed in the work.