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The kinetics of oxidation/reduction of the primary donor of photosystem 1 (PS1) P700, plastocyanin (PC), and ferredoxin (Fd) in the first leaves of barley seedlings under exposure to high-intensity light (2000 μmol∙quanta∙m–2·s–1, 30 min) and elevated temperature (40°C, 3 h) were studied using differential absorption photometry. Exposure to high-intensity light increased the accumulation of the oxidized form of PS1 reaction center P700+, oxidized PC, and reduced Fd. Reduced Fd was not reoxidized under the same conditions on exposure to red and far-red light. The accumulation of P700+, PC+, and Fd – increased in barley seedlings exposed to elevated temperatures. Also, reoxidation of leaf Fd accelerated under red light. Oxidized Fd did not accumulate under far-red light. It was concluded that photo-independent electron flow through Fd under light stress and alternative electron flows involving the plastoquinone pool under heat stress were activated. [ABSTRACT FROM AUTHOR]

The involvement of the second pair of chlorophylls, termed A-1A and A-1B, in light-induced electron transfer in photosystem I (PSI) is currently debated. Asparagines at PsaA600 and PsaB582 are involved in coordinating the A-1B and A-1A pigments, respectively. Here we have mutated these asparagine residues to methionine in two single mutants and a double mutant in PSI from Synechocystis sp. PCC 6803, which we term NA600M, NB582M, and NA600M/NB582M mutants. (P700+–P700) FTIR difference spectra (DS) at 293 K were obtained for the wild-type and the three mutant PSI samples. The wild-type and mutant FTIR DS differ considerably. This difference indicates that the observed changes in the (P700+–P700) FTIR DS cannot be due to only the PA and PB pigments of P700. Comparison of the wild-type and mutant FTIR DS allows the assignment of different features to both A-1 pigments in the FTIR DS for wild-type PSI and assesses how these features shift upon cation formation and upon mutation. While the exact role the A-1 pigments play in the species we call P700 is unclear, we demonstrate that the vibrational modes of the A-1A and A-1B pigments are modified upon P700+ formation. Previously, we showed that the A-1 pigments contribute to P700 in green algae. In this manuscript, we demonstrate that this is also the case in cyanobacterial PSI. The nature of the mutation-induced changes in algal and cyanobacterial PSI is similar and can be considered within the same framework, suggesting a universality in the nature of P700 in different photosynthetic organisms. [ABSTRACT FROM AUTHOR]

I present here a personal perspective of the evolution of the two-light reaction two-pigment scheme for the electron transport in oxygenic photosynthesis - as I have lived through it - first as a graduate student of Robert Emerson, from September 1956-January 1959, and then of Eugene Rabinowitch from February 1959-September 1960. I have provided here some of the key published work in my way and have also provided a few photographs. It is essential to remind ourselves upfront that different individuals may have different recollections of the same event (see e.g., the Japanese movie 'Rashomon' https://en.wikipedia.org/wiki/Rashomon). Thus, I encourage others to write about their perspective. Further, I recognize, upfront, the support and encouragement I have received from Robert Blankenship for writing this perspective, and from Győző Garab for submission of this story to Photosynthetica. I begin my perspective with a dedication to Robert Emerson, whose pioneering work led to the concept of the two-light reaction two-pigment system in oxygenic photosynthesis.

I present here a personal perspective of the evolution of the two-light reaction two-pigment scheme for the electron transport in oxygenic photosynthesis – as I have lived through it – first as a graduate student of Robert Emerson, from September 1956–January 1959, and then of Eugene Rabinowitch from February 1959–September 1960. I have provided here some of the key published work in my way and have also provided a few photographs. It is essential to remind ourselves upfront that different individuals may have different recollections of the same event (see e.g., the Japanese movie ‘Rashomon’ https://en.wikipedia.org/wiki/Rashomon). Thus, I encourage others to write about their perspective. Further, I recognize, upfront, the support and encouragement I have received from Robert Blankenship for writing this perspective, and from Győző Garab for submission of this story to Photosynthetica. I begin my perspective with a dedication to Robert Emerson, whose pioneering work led to the concept of the two-light reaction two-pigment system in oxygenic photosynthesis. [ABSTRACT FROM AUTHOR]

Environmental stress increases the risk of electron accumulation in photosystem I (PSI) of chloroplasts, which can cause oxygen (O2) reduction to superoxide radicals and decreased photosynthetic ability. We used three Arabidopsis thaliana lines: wild-type (WT) and the mutants pgr5hope1 and paa1-7/pox1. These lines have different reduced states of iron/sulfur (Fe/S) signals, including Fx, FA/FB, and ferredoxin, the electron carriers at the acceptor side of PSI. In the dark, short-pulse light was repetitively illuminated to the intact leaves of the plants to provide electrons to the acceptor side of PSI. WT and pgr5hope1 plants showed full reductions of Fe/S during short-pulse light and PSI inactivation. In contrast, paa1-7/pox1 showed less reduction of Fe/S and its PSI was not inactivated. Under continuous actinic-light illumination, pgr5hope1 showed no P700 oxidation with higher Fe/S reduction due to the loss of photosynthesis control and PSI inactivation. These results indicate that the accumulation of electrons at the acceptor side of PSI may trigger the production of superoxide radicals. P700 oxidation, responsible for the robustness of photosynthetic organisms, participates in reactive oxygen species suppression by oxidizing the acceptor side of PSI. [ABSTRACT FROM AUTHOR]

Photosynthesis is the process that harnesses, converts and stores light energy in the form of chemical energy in bonds of organic compounds. Oxygenic photosynthetic organisms (i.e., plants, algae and cyanobacteria) employ an efficient apparatus to split water and transport electrons to high-energy electron acceptors. The photosynthetic system must be finely balanced between energy harvesting and energy utilisation, in order to limit generation of dangerous compounds that can damage the integrity of cells. Insight into how the photosynthetic components are protected, regulated, damaged, and repaired during changing environmental conditions is crucial for improving photosynthetic efficiency in crop species. Photosystem I (PSI) is an integral component of the photosynthetic system located at the juncture between energy-harnessing and energy consumption through metabolism. Although the main site of photoinhibition is the photosystem II (PSII), PSI is also known to be inactivated by photosynthetic energy imbalance, with slower reactivation compared to PSII; however, several outstanding questions remain about the mechanisms of damage and repair, and about the impact of PSI photoinhibition on signalling and metabolism. In this review, we address the knowns and unknowns about PSI activity, inhibition, protection, and repair in plants. We also discuss the role of PSI in retrograde signalling pathways and highlight putative signals triggered by the functional status of the PSI pool. [ABSTRACT FROM AUTHOR]

Leaf-type ferredoxin-NADP(H) oxidoreductase, encoded by AtLFNR1 gene in Arabidopsis, extensively exists in chloroplast stroma and thylakoids and is responsible for the reduction of NADP+ to NADPH in PSI. To investigate the relationship between the characteristics of chlorophyll a fluorescence transients OJIP and the function of AtLFNR1 protein, the fluorescence rise kinetics OJIP curves of AtLFNR1 mutant leaves were determined. It is observed that AtLFNR1 mutant has a lower level of variable fluorescence intensity relative to wild type, especially the J-step and the IP-phase. The JIP-test analysis suggests that the loss of AtLFNR1 function decreases clearly the oxidation rate of plastocyanin as well as PSI reaction center, damages slightly PSII antenna size and pigment assemblies, but stimulates distinctly the photosynthetic activity of PSII. Such multiple effects contribute to the change of shape characteristics of the OJIP transient curve in AtLFNR1 mutant.

Photosynthesis is the process that harnesses, converts and stores light energy in the form of chemical energy in bonds of organic compounds. Oxygenic photosynthetic organisms (i.e., plants, algae and cyanobacteria) employ an efficient apparatus to split water and transport electrons to high-energy electron acceptors. The photosynthetic system must be finely balanced between energy harvesting and energy utilisation, in order to limit generation of dangerous compounds that can damage the integrity of cells. Insight into how the photosynthetic components are protected, regulated, damaged, and repaired during changing environmental conditions is crucial for improving photosynthetic efficiency in crop species. Photosystem I (PSI) is an integral component of the photosynthetic system located at the juncture between energy-harnessing and energy consumption through metabolism. Although the main site of photoinhibition is the photosystem II (PSII), PSI is also known to be inactivated by photosynthetic energy imbalance, with slower reactivation compared to PSII; however, several outstanding questions remain about the mechanisms of damage and repair, and about the impact of PSI photoinhibition on signalling and metabolism. In this review, we address the knowns and unknowns about PSI activity, inhibition, protection, and repair in plants. We also discuss the role of PSI in retrograde signalling pathways and highlight putative signals triggered by the functional status of the PSI pool.

Environmental stress increases the risk of electron accumulation in photosystem I (PSI) of chloroplasts, which can cause oxygen (O2) reduction to superoxide radicals and decreased photosynthetic ability. We used three Arabidopsis thaliana lines: wild-type (WT) and the mutants pgr5hope1 and paa1-7/pox1. These lines have different reduced states of iron/sulfur (Fe/S) signals, including Fx, FA/FB, and ferredoxin, the electron carriers at the acceptor side of PSI. In the dark, short-pulse light was repetitively illuminated to the intact leaves of the plants to provide electrons to the acceptor side of PSI. WT and pgr5hope1 plants showed full reductions of Fe/S during short-pulse light and PSI inactivation. In contrast, paa1-7/pox1 showed less reduction of Fe/S and its PSI was not inactivated. Under continuous actinic-light illumination, pgr5hope1 showed no P700 oxidation with higher Fe/S reduction due to the loss of photosynthesis control and PSI inactivation. These results indicate that the accumulation of electrons at the acceptor side of PSI may trigger the production of superoxide radicals. P700 oxidation, responsible for the robustness of photosynthetic organisms, participates in reactive oxygen species suppression by oxidizing the acceptor side of PSI.

The stoichiometry of photosystem II (PSII) and photosystem I (PSI) varies between photoautotrophic organisms. The cyanobacterium Synechocystis sp. PCC 6803 maintains two- to fivefold more PSI than PSII reaction center complexes, and we sought to modify this stoichiometry by changing the promoter region of the psaAB operon. We thus generated mutants with varied psaAB expression, ranging from ~3% to almost 200% of the wild-type transcript level, but all showing a reduction in PSI levels, relative to wild type, suggesting a role of the psaAB promoter region in translational regulation. Mutants with 25%-70% of wild-type PSI levels were photoautotrophic, with whole-chain oxygen evolution rates on a per-cell basis comparable to that of wild type. In contrast, mutant strains with <10% of the wild-type level of PSI were obligate photoheterotrophs. Variable fluorescence yields of all mutants were much higher than those of wild type, indicating that the PSI content is localized differently than in wild type, with less transfer of PSII-absorbed energy to PSI. Strains with less PSI saturate at a higher light intensity, enhancing productivity at higher light intensities. This is similar to what is found in mutants with reduced antennae. With 3-(3,4-dichlorophenyl)-1,1-dimethylurea present, P700 + re-reduction kinetics in the mutants were slower than in wild type, consistent with the notion that there is less cyclic electron transport if less PSI is present. Overall, strains with a reduction in PSI content displayed surprisingly vigorous growth and linear electron transport., Importance: Consequences of reduction in photosystem I content were investigated in the cyanobacterium Synechocystis sp. PCC 6803 where photosystem I far exceeds the number of photosystem II complexes. Strains with less photosystem I displayed less cyclic electron transport, grew more slowly at lower light intensity and needed more light for saturation but were surprisingly normal in their whole-chain electron transport rates, implying that a significant fraction of photosystem I is dispensable for linear electron transport in cyanobacteria. These strains with reduced photosystem I levels may have biotechnological relevance as they grow well at higher light intensities., Competing Interests: The authors declare no conflict of interest.

Trace metal contaminations in natural waters, wetlands, and wastewaters pose serious threats to aquatic ecosystems—mainly via targeting microalgae. In this work, we investigated the effects of toxic amounts of chromium and cadmium ions on the structure and function of the photosynthetic machinery of Chlorella variabilis cells. To halt the propagation of cells, we used high concentrations of Cd and Cr, 50–50 mg L−1, in the forms of CdCl2 x 2.5 H2O and K2Cr2O7, respectively. Both treatments led to similar, about 50% gradual diminishment of the chlorophyll contents of the cells in 48 h, which was, however, accompanied by a small (~10%) but statistically significant enrichment (Cd) and loss (Cr) of ß-carotene. Both Cd and Cr inhibited the activity of photosystem II (PSII)—but with more severe inhibitions with Cr. On the contrary, the PsbA (D1) protein of PSII and the PsbO protein of the oxygen-evolving complex were retained more in Cr-treated cells than in the presence of Cd. These data and the higher susceptibility of P700 redox transients in Cr-treated cells suggest that, unlike with Cd, PSII is not the main target in the photochemical apparatus. These differences at the level of photochemistry also brought about dissimilarities at higher levels of the structural complexity of the photosynthetic apparatus. Circular dichroism (CD) spectroscopy measurements revealed moderate perturbations in the macro-organization of the protein complexes—with more pronounced decline in Cd-treated cells than in the cells with Cr. Also, as reflected by transmission electron microscopy and small-angle neutron scattering, the thylakoid membranes suffered shrinking and were largely fragmented in Cd-treated cells, whereas no changes could be discerned with Cr. The preservation of integrity of membranes in Cr-treated cells was most probably aided by high proportion of the de-epoxidized xanthophylls, which were absent with Cd. It can thus be concluded that beside strong similarities of the toxic effects of Cr and Cd, the response of the photosynthetic machinery of C. variabilis to these two trace metal ions substantially differ from each other—strongly suggesting different inhibitory and protective mechanisms following the primary toxic events.

The extremophilic red alga Galdieria partita is a facultative heterotroph that occupies mostly low-light microhabitats. However, the exceptional detection of abundant populations of G. partita in sunlight-exposed soil raises the possibility that exogenous organic carbon sources protect cells from photo-oxidative damage. The present study aimed to identify the photoprotective process activated by exogenous glucose under photo-oxidative stress. We demonstrated that exogenous glucose mitigated the photo-oxidative damage of cells exposed to 300 μmol photons m–2 s–1 photosynthetic active radiation. Photosynthesis carbon assimilation scarcely contributed to the cell growth in the presence of glucose, but the photosynthetic apparatus was nevertheless maintained and protected by glucose in a concentration-dependent manner. Supplementation of glucose increased expression of the L-gulonolactone oxidase gene essential for ascorbic acid biosynthesis, whereas no enhanced expression of the genes involved in carotenoid or tocopherol biosynthesis was observed. Under the photo-oxidative stress condition, the ascorbic acid content was strongly enhanced by exogenous glucose. We propose that the biosynthesis of ascorbic acid is one of the major photoprotective processes induced by exogenous glucose. The elucidation of how ascorbic acid is involved in scavenging reactive oxygen species provides key insights into the photoprotective mechanism in red algae.

Trace metal contaminations in natural waters, wetlands, and wastewaters pose serious threats to aquatic ecosystems—mainly via targeting microalgae. In this work, we investigated the effects of toxic amounts of chromium and cadmium ions on the structure and function of the photosynthetic machinery of Chlorella variabilis cells. To halt the propagation of cells, we used high concentrations of Cd and Cr, 50–50 mg L−1, in the forms of CdCl2 x 2.5 H2O and K2Cr2O7, respectively. Both treatments led to similar, about 50% gradual diminishment of the chlorophyll contents of the cells in 48 h, which was, however, accompanied by a small (~10%) but statistically significant enrichment (Cd) and loss (Cr) of ß-carotene. Both Cd and Cr inhibited the activity of photosystem II (PSII)—but with more severe inhibitions with Cr. On the contrary, the PsbA (D1) protein of PSII and the PsbO protein of the oxygen-evolving complex were retained more in Cr-treated cells than in the presence of Cd. These data and the higher susceptibility of P700 redox transients in Cr-treated cells suggest that, unlike with Cd, PSII is not the main target in the photochemical apparatus. These differences at the level of photochemistry also brought about dissimilarities at higher levels of the structural complexity of the photosynthetic apparatus. Circular dichroism (CD) spectroscopy measurements revealed moderate perturbations in the macro-organization of the protein complexes—with more pronounced decline in Cd-treated cells than in the cells with Cr. Also, as reflected by transmission electron microscopy and small-angle neutron scattering, the thylakoid membranes suffered shrinking and were largely fragmented in Cd-treated cells, whereas no changes could be discerned with Cr. The preservation of integrity of membranes in Cr-treated cells was most probably aided by high proportion of the deepoxidized xanthophylls, which were absent with Cd. It can thus be concluded that beside strong similarities of the toxic effects of Cr and Cd, the response of the photosynthetic machinery of C. variabilis to these two trace metal ions substantially differ from each other—strongly suggesting different inhibitory and protective mechanisms following the primary toxic events. [ABSTRACT FROM AUTHOR]

The extremophilic red alga Galdieria partita is a facultative heterotroph that occupies mostly low-light microhabitats. However, the exceptional detection of abundant populations of G. partita in sunlight-exposed soil raises the possibility that exogenous organic carbon sources protect cells from photo-oxidative damage. The present study aimed to identify the photoprotective process activated by exogenous glucose under photo-oxidative stress. We demonstrated that exogenous glucose mitigated the photo-oxidative damage of cells exposed to 300 μmol photons m–2 s–1 photosynthetic active radiation. Photosynthesis carbon assimilation scarcely contributed to the cell growth in the presence of glucose, but the photosynthetic apparatus was nevertheless maintained and protected by glucose in a concentration-dependent manner. Supplementation of glucose increased expression of the L-gulonolactone oxidase gene essential for ascorbic acid biosynthesis, whereas no enhanced expression of the genes involved in carotenoid or tocopherol biosynthesis was observed. Under the photo-oxidative stress condition, the ascorbic acid content was strongly enhanced by exogenous glucose. We propose that the biosynthesis of ascorbic acid is one of the major photoprotective processes induced by exogenous glucose. The elucidation of how ascorbic acid is involved in scavenging reactive oxygen species provides key insights into the photoprotective mechanism in red algae. [ABSTRACT FROM AUTHOR]

Leaf-type ferredoxin-NADP(H) oxidoreductase, encoded by AtLFNR1 gene in Arabidopsis, extensively exists in chloroplast stroma and thylakoids and is responsible for the reduction of NADP+ to NADPH in PSI. To investigate the relationship between the characteristics of chlorophyll a fluorescence transients OJIP and the function of AtLFNR1 protein, the fluorescence rise kinetics OJIP curves of AtLFNR1 mutant leaves were determined. It is observed that AtLFNR1 mutant has a lower level of variable fluorescence intensity relative to wild type, especially the J-step and the IP-phase. The JIP-test analysis suggests that the loss of AtLFNR1 function decreases clearly the oxidation rate of plastocyanin as well as PSI reaction center, damages slightly PSII antenna size and pigment assemblies, but stimulates distinctly the photosynthetic activity of PSII. Such multiple effects contribute to the change of shape characteristics of the OJIP transient curve in AtLFNR1 mutant. [ABSTRACT FROM AUTHOR]

In response to decreases in the assimilation efficiency of CO2, plants oxidize the reaction center chlorophyll (P700) of photosystem I (PSI) to suppress reactive oxygen species (ROS) production. In hydro-cultured sunflower leaves experiencing essential mineral deficiencies, we analyzed the following parameters that characterize PSI and PSII: (1) the reduction-oxidation states of P700 [Y(I), Y(NA), and Y(ND)]; (2) the relative electron flux in PSII [Y(II)]; (3) the reduction state of the primary electron acceptor in PSII, QA (1 − qL); and (4) the non-photochemical quenching of chlorophyll fluorescence (NPQ). Deficiency treatments for the minerals N, P, Mn, Mg, S, and Zn decreased Y(II) with an increase in the oxidized P700 [Y(ND)], while deficiencies for the minerals K, Fe, Ca, B, and Mo decreased Y(II) without an increase in Y(ND). During the induction of photosynthesis, the above parameters showed specific responses to each mineral. That is, we could diagnose the mineral deficiency and identify which mineral affected the photosynthesis parameters.

Photosynthesis involves the conversion of sunlight energy into stored chemical energy, which is achieved through electron transport along a series of redox reactions. Excess photosynthetic electron transport might be dangerous due to the risk of molecular oxygen reduction, generating reactive oxygen species (ROS) over-accumulation. Avoiding excess ROS production requires the rate of electron transport to be coordinated with the capacity of electron acceptors in the chloroplast stroma. Imbalance between the donor and acceptor sides of photosystem I (PSI) can lead to inactivation, which is called PSI photoinhibition. We used a light-inducible PSI photoinhibition system in Arabidopsis thaliana to resolve the time dynamics of inhibition and to investigate its impact on ROS production and turnover. The oxidation state of the PSI reaction center and rates of CO2 fixation both indicated strong and rapid PSI photoinhibition upon donor side/acceptor side imbalance, while the rate of inhibition eased during prolonged imbalance. PSI photoinhibition was not associated with any major changes in ROS accumulation or antioxidant activity; however, a lower level of lipid oxidation correlated with lower abundance of chloroplast lipoxygenase in PSI-inhibited leaves. The results of this study suggest that rapid activation of PSI photoinhibition under severe photosynthetic imbalance protects the chloroplast from over-reduction and excess ROS formation.

Photosynthesis involves the conversion of sunlight energy into stored chemical energy, which is achieved through electron transport along a series of redox reactions. Excess photosynthetic electron transport might be dangerous due to the risk of molecular oxygen reduction, generating reactive oxygen species (ROS) over-accumulation. Avoiding excess ROS production requires the rate of electron transport to be coordinated with the capacity of electron acceptors in the chloroplast stroma. Imbalance between the donor and acceptor sides of photosystem I (PSI) can lead to inactivation, which is called PSI photoinhibition. We used a light-inducible PSI photoinhibition system in Arabidopsis thaliana to resolve the time dynamics of inhibition and to investigate its impact on ROS production and turnover. The oxidation state of the PSI reaction center and rates of CO2 fixation both indicated strong and rapid PSI photoinhibition upon donor side/acceptor side imbalance, while the rate of inhibition eased during prolonged imbalance. PSI photoinhibition was not associated with any major changes in ROS accumulation or antioxidant activity; however, a lower level of lipid oxidation correlated with lower abundance of chloroplast lipoxygenase in PSI-inhibited leaves. The results of this study suggest that rapid activation of PSI photoinhibition under severe photosynthetic imbalance protects the chloroplast from over-reduction and excess ROS formation. [ABSTRACT FROM AUTHOR]

(P700+ – P700) Fourier transform visible and infrared difference spectra (DS) have been obtained using photosystem I (PSI) complexes isolated from cells of Fischerella thermalis PCC 7521 grown under white light (WL) or far-red light (FRL). PSI from cells grown under FRL (FRL-PSI) contain ~8 chlorophyll f (Chl f) molecules (Shen et al., Photosynth. Res. Jan. 2019). Both the visible and infrared DS indicate that neither the P A or P B pigments of P700 are Chl f molecules, but do support the conclusion that at least one of the A –1 cofactors is a Chl f molecule. The FTIR DS indicate that the hydrogen bond to the 131-keto C O group of the P A pigment of P700 is weakened in FRL-PSI, as might be expected given that the proteins that bind the P700 pigments are substantially different in FRL-PSI (Gan et al., Science 345, 1312–1317, 2014). The FTIR DS obtained using FRL-PSI display a band at 1664 cm−1 that is assigned (based on density functional theory calculations) to the 21-formyl C O group of Chl f , that upshifts 5 cm−1 upon P700+ formation. This is much less than expected for a cation-induced upshift, indicating that the Chl f molecule is not one of the pigments of P700. In WL-PSI the A –1 cofactor is a Chl a molecule with 131-keto and 133-methylester C O mode vibrations at 1696 and 1750 cm−1, respectively. In FRL-PSI the A –1 cofactor is a Chl f molecule with 131-keto and 133-methylester C O mode vibrations at 1702 and 1754 cm−1, respectively. Unlabelled Image • A -1 cofactor in FRL-PSI is a Chl f molecule. • The hydrogen bond to the P A pigment of P700 is weakened in FRL-PSI. • Studying FRL-PSI leads to an improved interpretation of P700 FTIR difference spectra. [ABSTRACT FROM AUTHOR]

Upon exposure to environmental stress, the primary electron donor in photosystem I (PSI), P700, is oxidized to suppress the production of reactive oxygen species that could oxidatively inactivate the function of PSI. The illumination of rice leaves with actinic light induces intrinsic fluctuations in the opening and closing of stomata, causing the net CO2 assimilation rate to fluctuate. We examined the effects of these intrinsic fluctuations on electron transport reactions. Under atmospheric O2 conditions (21 kPa), the effective quantum yield of photosystem II (PSII) (Y(II)) remained relatively high while the net CO2 assimilation rate fluctuated, which indicates the function of alternative electron flow. By contrast, under low O2 conditions (2 kPa), Y(II) fluctuated. These results suggest that photorespiration primarily drove the alternative electron flow. Photorespiration maintained the oxidation level of ferredoxin (Fd) throughout the fluctuation of the net CO2 assimilation rate. Moreover, the relative activity of photorespiration was correlated with both the oxidation level of P700 and the magnitude of the proton gradient across the thylakoid membrane in 21 kPa O2 conditions. These results show that photorespiration oxidized P700 by stimulating the proton gradient formation when CO2 assimilation was suppressed by stomatal closure.

Photosynthesis fixes CO2 and converts it to sugar, using chemical-energy compounds of both NADPH and ATP, which are produced in the photosynthetic electron transport system. The photosynthetic electron transport system absorbs photon energy to drive electron flow from Photosystem II (PSII) to Photosystem I (PSI). That is, both PSII and PSI are full of electrons. O2 is easily reduced to a superoxide radical (O2−) at the reducing side, i.e., the acceptor side, of PSI, which is the main production site of reactive oxygen species (ROS) in photosynthetic organisms. ROS-dependent inactivation of PSI in vivo has been reported, where the electrons are accumulated at the acceptor side of PSI by artificial treatments: exposure to low temperature and repetitive short-pulse (rSP) illumination treatment, and the accumulated electrons flow to O2, producing ROS. Recently, my group found that the redox state of the reaction center of chlorophyll P700 in PSI regulates the production of ROS: P700 oxidation suppresses the production of O2− and prevents PSI inactivation. This is why P700 in PSI is oxidized upon the exposure of photosynthesis organisms to higher light intensity and/or low CO2 conditions, where photosynthesis efficiency decreases. In this study, I introduce a new molecular mechanism for the oxidation of P700 in PSI and suppression of ROS production from the robust relationship between the light and dark reactions of photosynthesis. The accumulated protons in the lumenal space of the thylakoid membrane and the accumulated electrons in the plastoquinone (PQ) pool drive the rate-determining step of the P700 photo-oxidation reduction cycle in PSI from the photo-excited P700 oxidation to the reduction of the oxidized P700, thereby enhancing P700 oxidation.

The rapid detection of glyphosate resistance in goosegrass (Eleusine indica) will enhance our ability to respond to new resistant populations of this major weed. Chlorophyll fluorescence (Fluo) and P700 (reaction center chlorophyll of photosystem I) absorbance were analyzed in one biotype of goosegrass that is resistant to glyphosate and in another that remains sensitive to the herbicide. Both biotypes were treated with a foliar spray of glyphosate. Differences in photosystem II maximum quantum yield (Fv/Fm), effective photochemical quantum yield (Y(II)), and non-photochemical quenching (NPQ) between the biotypes increased over time. Values for Fv/Fm and Y(II) differed between the two biotypes 24 h after treatment (HAT). Differentiated activities and energy dissipation processes of photosystem II (PSII) and energy dissipation processes of photosystem I (PSI) were manifested in the two biotypes 24 HAT with 20 mmol L−1 glyphosate. Differentiated energy dissipation processes of PSI were still apparent 24 HAT with 200 mmol L−1 glyphosate. These results indicate that the Fluo parameters related to PSII activity and energy dissipation and the P700 parameters related to energy dissipation are suitable indicators that enable rapid detection of glyphosate resistance in goosegrass.

Reef building corals (phylum Cnidaria) harbor endosymbiotic dinoflagellate algae (genus Symbiodinium) that generate photosynthetic products to fuel their host's metabolism. Non-invasive techniques such as chlorophyll (Chl) fluorescence analyses of Photosystem II (PSII) have been widely used to estimate the photosynthetic performance of Symbiodinium in hospite. However, since the spatial origin of PSII chlorophyll fluorescence in coral tissues is uncertain, such signals give limited information on depth-integrated photosynthetic performance of the whole tissue. In contrast, detection of absorbance changes in the near infrared (NIR) region integrates signals from deeper tissue layers due to weak absorption and multiple scattering of NIR light. While extensively utilized in higher plants, NIR bio-optical techniques are seldom applied to corals. We have developed a non-intrusive measurement method to examine photochemistry of intact corals, based on redox kinetics of the primary electron donor in Photosystem I (P700) and chlorophyll fluorescence kinetics (Fast-Repetition Rate fluorometry, FRRf). Since the redox state of P700 depends on the operation of both PSI and PSII, important information can be obtained on the PSII-PSI intersystem electron transfer kinetics. Under moderate, sub-lethal heat stress treatments (33°C for ~20 min), the coral Pavona decussata exhibited down-regulation of PSII electron transfer kinetics, indicated by slower rates of electron transport from QA to plastoquinone (PQ) pool, and smaller relative size of oxidized PQ with concomitant decrease of a specifically-defined P700 kinetics area, which represents the active pool of PSII. The maximum quantum efficiency of PSII (Fv/Fm) and functional absorption cross-section of PSII (σPSII) remained unchanged. Based on the coordinated response of P700 parameters and PSII-PSI electron transport properties, we propose that simple P700 kinetics parameters as employed here serve as indicators of the integrity of PSII-PSI electron transfer dynamics in corals.

Oxygen (O2)-evolving photosynthetic organisms oxidize the reaction center chlorophyll, P700, in photosystem I (PSI) to suppress the production of reactive oxygen species. The oxidation of P700 is accompanied by alternative electron flow in PSI (AEF-I), which is not required for photosynthetic linear electron flow (LEF). To characterize AEF-I, we compared the redox reactions of P700 and ferredoxin (Fd) during the induction of carbon dioxide (CO2) assimilation in wheat leaves, using dark-interval relaxation kinetics analysis. Switching on an actinic light (1000 μmol photons m−2 s−1) at ambient CO2 partial pressure of 40 Pa and ambient O2 partial pressure of 21 kPa gradually oxidized P700 (P700+) and enhanced the reduction rate of P700+ (vP700) and oxidation rate of reduced Fd (vFd). The vFd showed a positive linear relationship with an apparent photosynthetic quantum yield of PSII (Y[II]) originating at point zero; the redox turnover of Fd is regulated by LEF via CO2 assimilation and photorespiration. The vP700 also showed a positive linear relationship with Y(II), but the intercept was positive, not zero. That is, the electron flux in PSI included the electron flux in AEF-I in addition to that in LEF. This indicates that the oxidation of P700 induces AEF-I. We propose a possible mechanism underlying AEF-I and its physiological role in the mitigation of oxidative damage.

Cyclic electron flow around PSI can protect photosynthetic electron carriers under conditions of stromal over-reduction. The goal of the research reported in this paper was to investigate the responses of both PSI and PSII to a short-term heat stress in two rice lines with different capacities of cyclic electron transfer, i.e. Q4149 with a high capacity (hcef) and C4023 with a low capacity (lcef). The absorbance change at 820 nm (ΔA820) was used here to assess the charge separation in the photosystem I (PSI) reaction center (P700). The results obtained show that short-term heat stress abolishes the FQR-dependent CEF in rice and accelerates the initial rate of P700+ re-reduction. The P700+ amplitude was slightly increased at a moderate heat-stress (35°C) because of a partial restriction of FQR but it was decreased following high heat-stress (42°C). Assessment of PSI and PSII activities shows that PSI is more susceptible to heat stress than photosystem II (PSII). Under high temperature, FQR-dependent CEF was completely removed and NDH-dependent CEF was up-regulated and strengthened to a higher extent in C4023 than in Q4149. Specifically, under normal growth temperature, hcef (Q4149) was characterized by higher FQR- and NDH-dependent CEF rates than lcef (C4023). Following thermal stress, the activation of NDH-pathway was 130% and 10% for C4023 and Q4149, respectively. Thus, the NDH-dependent CEF may constitute the second layer of plant protection and defence against heat stress after the main route, i.e. FQR-dependent CEF, reaches its capacity. We discuss the possibility that under high heat stress, the NDH pathway serves as a safety valve to dissipate excess energy by cyclic photophosphorylation and overcome the stroma over-reduction following inhibition of CO2 assimilation and any shortage or lack in the FQR pathway. The potential role of the NDH-dependent pathway during the evolution of C4 photosynthesis is briefly discussed.