Photoprotection Of Photosystem II Research Articles

Harnessing the Role of Foliar Applied Salicylic Acid in Decreasing Chlorophyll Content to Reassess Photosystem II Photoprotection in Crop Plants.

Salicylic acid (SA), an essential plant hormone, has received much attention due to its role in modulating the adverse effects of biotic and abiotic stresses, acting as an antioxidant and plant growth regulator. However, its role in photosynthesis under non stress conditions is controversial. By chlorophyll fluorescence imaging analysis, we evaluated the consequences of foliar applied 1 mM SA on photosystem II (PSII) efficiency of tomato (Solanum lycopersicum L.) plants and estimated the reactive oxygen species (ROS) generation. Tomato leaves sprayed with 1 mM SA displayed lower chlorophyll content, but the absorbed light energy was preferentially converted into photochemical energy rather than dissipated as thermal energy by non-photochemical quenching (NPQ), indicating photoprotective effects provided by the foliar applied SA. This decreased NPQ, after 72 h treatment by 1 mM SA, resulted in an increased electron transport rate (ETR). The molecular mechanism by which the absorbed light energy was more efficiently directed to photochemistry in the SA treated leaves was the increased fraction of the open PSII reaction centers (qp), and the increased efficiency of open reaction centers (Fv’/Fm’). SA induced a decrease in chlorophyll content, resulting in a decrease in non-regulated energy dissipated in PSII (ΦNO) under high light (HL) treatment, suggesting a lower amount of triplet excited state chlorophyll (3Chl*) molecules available to produce singlet oxygen (1O2). Yet, the increased efficiency, compared to the control, of the oxygen evolving complex (OEC) on the donor side of PSII, associated with lower formation of hydrogen peroxide (H2O2), also contributed to less creation of ROS. We conclude that under non stress conditions, foliar applied SA decreased chlorophyll content and suppressed phototoxicity, offering PSII photoprotection; thus, it can be regarded as a mechanism that reduces photoinhibition and photodamage, improving PSII efficiency in crop plants.

Is Photoprotection of PSII One of the Key Mechanisms for Drought Tolerance in Maize?

Drought is one of the most important abiotic stress factors limiting maize production worldwide. The objective of this study was to investigate whether photoprotection of PSII was associated with the degree of drought tolerance and yield in three maize hybrids (30Y87, 31R88, P3939). To do this, three maize hybrids were subjected to three cycles of drought, and we measured the activities of photosystem II (PSII) and photosystem I (PSI). In a second field experiment, three maize hybrids were subjected to drought by withholding irrigation, and plant water status, yield and yield attributes were measured. Drought stress decreased leaf water potential (ΨL) in three maize hybrids, and this reduction was more pronounced in hybrid P3939 (−40%) compared to that of 30Y87 (−30%). Yield and yield attributes of three maize hybrids were adversely affected by drought. The number of kernels and 100-kernel weight was the highest in maize hybrid 30Y87 (−56%, −6%), whereas these were lowest in hybrid P3939 (−88%, −23%). Drought stress reduced the quantum yield of PSII [Y(II)], photochemical quenching (qP), electron transport rate through PSII [ETR(II)] and NPQ, except in P3939. Among the components of NPQ, drought increased the Y(NPQ) with concomitant decrease in Y(NO) only in P3939, whereas Y(NO) increased in drought-stressed plants of hybrid 30Y87 and 31R88. However, an increase in cyclic electron flow (CEF) around PSI and Y(NPQ) in P3939 might have protected the photosynthetic machinery but it did not translate in yield. However, drought-stressed plants of 30Y87 might have sufficiently downregulated PSII to match the energy consumption in downstream biochemical processes. Thus, changes in PSII and PSI activity and development of NPQ through CEF are physiological mechanisms to protect the photosynthetic apparatus, but an appropriate balance between these physiological processes is required, without which plant productivity may decline.

Loss of carbonic anhydrase in the thylakoid lumen causes unusual moderate-light-induced rearrangement of the chloroplast in Chlamydomonas reinhardtii as a way of photosystem II photoprotection

Chlamydomonas reinhardtii cells have a single large cup-shaped chloroplast that can lose lobes under high light to prevent photodamage of the photosynthetic apparatus, including photosystem II (PSII). Here, under moderate light treatment, the development of the unusual morphology of the chloroplast is shown for mutant cia3, which is deficient in carbonic anhydrase (EC 4.2.1.1) CAH3 in the thylakoid lumen, while such light intensity is harmless for wild type (WT) cells for hours. Cia3 cells had more activated PSII photoprotective mechanisms and showed a tendency to shift in the balance of the PSII damage-repair cycle, whereas PSII retained the same photosynthetic efficiency as in the WT. These findings allow speculation about the unique PSII photoprotection strategy by rearranging the chloroplast in the absence of CAH3. CAH3, in turn, is suggested to be an important participant of the C. reinhardtii photosynthetic apparatus operation, functioning in close connection with PSII.

Analysis of the changes of electron transfer and heterogeneity of photosystem II in Deg1-reduced Arabidopsis plants.

Deg1 protease functions in protease and chaperone of PSII complex components, but few works were performed to study the effects of Deg1 on electron transport activities on the donor and acceptor side of PSII and its correlation with the photoprotection of PSII during photoinhibition. Therefore, we performed systematic and comprehensive investigations of electron transfers on the donor and acceptor sides of photosystem II (PSII) in the Deg1-reduced transgenic lines deg1-2 and deg1-4. Both the maximal quantum efficiency of PSII photochemistry (Fv/Fm) and the actual PSII efficiency (ΦPSII) decreased significantly in the transgenic plants. Increases in nonphotochemical quenching (NPQ) and the dissipated energy flux per reaction center (DI0/RC) were also shown in the transgenic plants. Along with the decreased D1, CP47, and CP43 content, these results suggested photoinhibition under growth light conditions in transgenic plants. Decreased Deg1 caused inhibition of electron transfer on the PSII reducing side, leading to a decline in the number of QB-reducing centers and accumulation of QB-nonreducing centers. The Tm of the Q band shifted from 5.7 °C in the wild-type plant to 10.4 °C and 14.2 °C in the deg1-2 and deg1-4 plants, respectively, indicating an increase in the stability of S2QA¯ in transgenic plants. PSIIα in the transgenic plants largely reduced, while PSIIβ and PSIIγ increased with the decline in the Deg1 levels in transgenic plants suggesting PSIIα centers gradually converted into PSIIβ and PSIIγ centers in the transgenic plants. Besides, the connectivity of PSIIα and PSIIβ was downregulated in transgenic plants. Our results reveal that downregulation of Deg1 protein levels induced photoinhibition in transgenic plants, leading to loss of PSII activities on both the donor and acceptor sides in transgenic plants. These results give a new insight into the regulation role of Deg1 in PSII electron transport.

Lutein-mediated photoprotection of photosynthetic machinery in Arabidopsis thaliana exposed to chronic low ultraviolet-B radiation

Ecologically relevant low UV-B is reported to alter reactive oxygen species metabolism and anti-oxidative systems through an up-regulation of enzymes of the phenylpropanoid pathway. However, little is known about low UV-B-induced changes in carotenoid profile and their impacts on light harvesting and photoprotection of photosystem II (PSII) in plants. We investigated carotenoids profile, chlorophyll pigments, phenolics, photosynthetic efficiency and growth in Arabidopsis thaliana (Col-0) plants grown under photosynthetically active radiation (PAR), PAR+ ultraviolet (UV)-A and PAR+UV-A+B regimes for 10 days in order to assess plant acclimation to low UV-B radiation. A chlorophyll fluorescence assay was used to examine UV-B tolerance in plants further exposed to acute high UV-B for 4 and 6 h following a 10-day growth under different PAR and UV regimes. We found that both PAR+ UV-A and PAR+UV-A+B regimes had no negative effect on quantum efficiency, electron transport rate, rosette diameter, relative growth rate and shoot dry weight of plants. Chronic PAR+ UV-A regime considerably (P < 0.05) increased violaxanthin (26 %) and neoxanthin (92 %) content in plants. Plant exposure to chronic PAR+UV-A+B significantly (P < 0.05) increased violaxanthin (48 %), neoxanthin (63 %), lutein (33 %), 9-cis ß-carotene (28 %), total ß-carotene (29 %) and total phenolics (108 %). The maximum photochemical efficiency (Fv/Fm) in leaves was found to be positively correlated with total phenolics (rho = 0.81 and rho = 0.91, P < 0.05 for 4 and 6 h, respectively) and non-photochemical quenching (qN) (rho = 0.81 and rho = 0.84, P < 0.05 for 4 and 6 h, respectively) in plants exposed to acute high UV-B for 4 and 6 h following a 10-day growth under chronic PAR+UV-A+B. There was also a significant positive correlation (rho = 0.93, P < 0.01) between qN and lutein content in the plants exposed to acute high UV-B stress for 4 h following plant exposure to chronic PAR+UV-A+B. The findings from our study indicate that plants grown under chronic PAR+UV-A+B displayed higher photoprotection of PSII against acute high UV-B stress than those grown under PAR and PAR+ UV-A regimes. An induction of phenolics and lutein-mediated development of qN were involved in the photoprotection of PSII against UV-B-induced oxidative stress.

Induction of cyclic electron flow around photosystem I during heat stress in grape leaves

Photosystem II (PSII) in plants is susceptible to high temperatures. The cyclic electron flow (CEF) around PSI is thought to protect both PSII and PSI from photodamage. However, the underlying physiological mechanisms of the photosynthetic electron transport process and the role of CEF in grape at high temperatures remain unclear. To investigate this issue, we examined the responses of PSII energy distribution, the P700 redox state and CEF to high temperatures in grape leaves. After exposing ‘Cabernet Sauvignon’ leaves to various temperatures (25, 30, 35, 40 and 45°C) in the light (600μmol photons m−2s−1) for 4h, the maximum quantum yield of PSII (Fv/Fm) significantly decreased at high temperatures (40 and 45°C), while the maximum photo-oxidizable P700 (Pm) was not affected. As the temperature increased, higher initial rates of increase in post-illumination Chl fluorescence were detected, which were accompanied by an increase in high energy state quenching (qE). The chloroplast NAD(P)H dehydrogenase-dependent CEF (NDH-dependent CEF) activities were different among grape cultivators. ‘Gold Finger’ with greater susceptibility to photoinhibition, exhibited lower NDH-dependent CEF activities under acute heat stress than a more heat tolerant ‘Cabernet Sauvignon’. These results suggest that overclosure of PSII reaction centers at high temperature resulted in the photoinhibition of PSII, while the stimulation of CEF in grape played an important role in the photoprotection of PSII and PSI at high temperatures through contributing to the generation of a proton gradient.

Contribution of the Alternative Respiratory Pathway to PSII Photoprotection in C3 and C4 Plants

The mechanism by which the mitochondrial alternative oxidase (AOX) pathway contributes to photosystem II (PSII) photoprotection is in dispute. It was generally thought that the AOX pathway protects photosystems by dissipating excess reducing equivalents exported from chloroplasts through the malate/oxaloacetate (Mal/OAA) shuttle and thus preventing the over-reduction of chloroplasts. In this study, using the aox1a Arabidopsis mutant and nine other C3 and C4 plant species, we revealed an additional action model of the AOX pathway in PSII photoprotection. Although the AOX pathway contributes to PSII photoprotection in C3 leaves treated with high light, this contribution was observed to disappear when photorespiration was suppressed. Disruption or inhibition of the AOX pathway significantly decreased the photorespiration in C3 leaves. Moreover, the AOX pathway did not respond to high light and contributed little to PSII photoprotection in C4 leaves possessing a highly active Mal/OAA shuttle but with little photorespiration. These results demonstrate that the AOX pathway contributes to PSII photoprotection in C3 plants by maintaining photorespiration to detoxify glycolate and via the indirect export of excess reducing equivalents from chloroplasts by the Mal/OAA shuttle. This new action model explains why the AOX pathway does not contribute to PSII photoprotection in C4 plants.

Light acclimation in the lycophyte Selaginella martensii depends on changes in the amount of photosystems and on the flexibility of the light-harvesting complex II antenna association with both photosystems.

Vascular plants have evolved a long-term light acclimation strategy primarily relying on the regulation of the relative amounts of light-harvesting complex II (LHCII) and of the two photosystems, photosystem I (PSI) and photosystem II (PSII). We investigated whether such a model is also valid in Selaginella martensii, a species belonging to the early diverging group of lycophytes. Selaginellamartensii plants were acclimated to three natural light regimes (extremely low light (L), medium light (M) and full sunlight (H)) and thylakoid organization was characterized combining ultrastructural, biochemical and functional methods. From L to H plants, thylakoid architecture was rearranged from (pseudo)lamellar to predominantly granal, the PSII : PSI ratio changed in favour of PSI, and the photochemical capacity increased. However, regulation of light harvesting did not occur through variations in the amount of free LHCII, but rather resulted from the flexibility of the association of free LHCII with PSII and PSI. In lycophytes, the free interspersed LHCII serves a fixed proportion of reaction centres, either PSII or PSI, and the regulation of PSI-LHCII(-PSII) megacomplexes is an integral part of long-term acclimation. Free LHCII ensures photoprotection of PSII, allows regulated use of PSI as an energy quencher, and can also quench endangered PSI.

Redox potential of the terminal quinone electron acceptor QB in photosystem II reveals the mechanism of electron transfer regulation

Photosystem II (PSII) extracts electrons from water at a Mn4CaO5 cluster using light energy and then transfers them to two plastoquinones, the primary quinone electron acceptor QA and the secondary quinone electron acceptor QB. This forward electron transfer is an essential process in light energy conversion. Meanwhile, backward electron transfer is also significant in photoprotection of PSII proteins. Modulation of the redox potential (Em) gap of QA and QB mainly regulates the forward and backward electron transfers in PSII. However, the full scheme of electron transfer regulation remains unresolved due to the unknown Em value of QB. Here, for the first time (to our knowledge), the Em value of QB reduction was measured directly using spectroelectrochemistry in combination with light-induced Fourier transform infrared difference spectroscopy. The Em(QB (-)/QB) was determined to be approximately +90 mV and was virtually unaffected by depletion of the Mn4CaO5 cluster. This insensitivity of Em(QB (-)/QB), in combination with the known large upshift of Em(QA (-)/QA), explains the mechanism of PSII photoprotection with an impaired Mn4CaO5 cluster, in which a large decrease in the Em gap between QA and QB promotes rapid charge recombination via QA (-).

MPH1 is a thylakoid membrane protein involved in protecting photosystem II from photodamage in land plants

Photosystem II (PSII) is highly susceptible to photoinhibition caused by environmental stimuli such as high light; therefore plants have evolved multifaceted mechanisms to efficiently protect PSII from photodamage. We previously published data suggesting that Maintenance of PSII under High light 1 (MPH1, encoded by AT5G07020), a PSII-associated proline-rich protein found in land plants, participates in the maintenance of normal PSII activity under photoinhibitory stress. Here we provide additional evidence for the role of MPH1 in protecting PSII against photooxidative damage. Two Arabidopsis thaliana mutants lacking a functional MPH1 gene suffer from severe photoinhibition relative to the wild-type plants under high irradiance light. The mph1 mutants exhibit significantly decreased PSII quantum yield and electron transport rate after exposure to photoinhibitory light. The mutants also display drastically elevated photodamage to PSII reaction center proteins after high-light treatment. These data add further evidence that MPH1 is involved in PSII photoprotection in Arabidopsis. MPH1 homologs are found across phylogenetically diverse land plants but are not detected in algae or prokaryotes. Taken together, these results suggest that MPH1 protein began to play a role in protecting PSII against excess light following the transition from aquatic to terrestrial conditions.

Long-range interaction between the Mn4CaO5 cluster and the non-heme iron center in photosystem II as revealed by FTIR spectroelectrochemistry.

It is known that inactivation of the Mn4CaO5 cluster, the catalytic center of water oxidation in photosystem II (PSII), induces a positive shift of the redox potential (Em) of the primary quinone electron acceptor QA by ∼+150 mV, resulting in suppression of the electron transfer from QA to the secondary quinone acceptor QB. Although the relevance of this Em(QA(-)/QA) shift to the photoprotection of PSII has been debated, its molecular mechanism is still enigmatic from a structural viewpoint because QA is ∼40 Å from the Mn4CaO5 cluster. In this work, we have investigated the influence of Mn depletion on the Em of the non-heme iron, which is located between QA and QB, and its surrounding structure. Electrochemical measurements in combination with Fourier transform infrared (FTIR) spectroscopy revealed that Mn depletion shifts Em(Fe(2+)/Fe(3+)) by +18 mV, which is ∼8 times smaller than the shift of Em(QA(-)/QA). Comparison of the Fe(2+)/Fe(3+) FTIR difference spectra between intact and Mn-depleted PSII samples showed that Mn depletion altered the pKa's of a His ligand to the non-heme iron, most probably the D1-His215 interacting QB, and a carboxylate group, possibly D1-Glu244, coupled with the non-heme iron. It was further shown that Mn depletion influences the C≡N vibration of bromoxynil bound to the QB site, indicative of the modification of the QB binding site. On the basis of these results, we discuss the mechanism of a long-range interaction between the donor and acceptor sides of PSII.

Glycine betaine application in grain filling wheat plants alleviates heat and high light-induced photoinhibition by enhancing the psbA transcription and stomatal conductance

Wheat (Triticum aestivum) plants often suffer from heat stress combined with high irradiance during the grain filling stage. Glycine betaine (GB) is a compatible solute that accumulates rapidly to enhance stress tolerance in wheat under abiotic stress. In this study, the effect of foliar application of GB on photosystem II (PSII) photochemistry in wheat leaves under the heat and high light stresses were investigated. The results showed that GB-supplemented wheat plants maintained higher chlorophyll content, photochemical activity of PSII and net photosynthetic rate (P n) than stressed only plants. Moreover, GB-treated plants could retard the decrease of psbA gene transcription and accelerate the endogenous accumulation of GB in leaves. The results suggested that the increasing stress tolerance by GB accumulation was associated with an improvement in D1 protein synthesis, which accelerated the repair of PSII following stress-enhanced photoinhibition. Moreover, the accumulation of abscisic acid under the heat and high light stresses was retarded by foliar application of GB, and the decrease of stomatal conductance was inhibited as well. The results suggested that GB accumulation in vivo was involved in the regulation of stomatal conductance. Further work is required to elucidate the mechanism of GB-induced stomatal movement and PSII photoprotection.

The relationship between maximum tolerated light intensity and photoprotective energy dissipation in the photosynthetic antenna: chloroplast gains and losses

The principle of quantifying the efficiency of protection of photosystem II (PSII) reaction centres against photoinhibition by non-photochemical energy dissipation (NPQ) has been recently introduced by Ruban & Murchie (2012 Biochim. Biophys. Acta 1817, 977-982 (doi:10.1016/j.bbabio.2012.03.026)). This is based upon the assessment of two key parameters: (i) the relationship between the PSII yield and NPQ, and (ii) the fraction of intact PSII reaction centres in the dark after illumination. In this paper, we have quantified the relationship between the amplitude of NPQ and the light intensity at which all PSII reaction centres remain intact for plants with different levels of PsbS protein, known to play a key role in the process. It was found that the same, nearly linear, relationship exists between the levels of the protective NPQ component (pNPQ) and the tolerated light intensity in all types of studied plants. This approach allowed for the quantification of the maximum tolerated light intensity, the light intensity at which all plant leaves become photoinhibited, the fraction of (most likely) unnecessary or 'wasteful' NPQ, and the fraction of photoinhibited PSII reaction centres under conditions of prolonged illumination by full sunlight. It was concluded that the governing factors in the photoprotection of PSII are the level and rate of protective pNPQ formation, which are often in discord with the amplitude of the conventional measure of photoprotection, the quickly reversible NPQ component, qE. Hence, we recommend pNPQ as a more informative and less ambiguous parameter than qE, as it reflects the effectiveness and limitations of the major photoprotective process of the photosynthetic membrane.

Photosynthetic electron transport and specific photoprotective responses in wheat leaves under drought stress

The photosynthetic responses of wheat (Triticum aestivum L.) leaves to different levels of drought stress were analyzed in potted plants cultivated in growth chamber under moderate light. Low-to-medium drought stress was induced by limiting irrigation, maintaining 20% of soil water holding capacity for 14days followed by 3days without water supply to induce severe stress. Measurements of CO2 exchange and photosystem II (PSII) yield (by chlorophyll fluorescence) were followed by simultaneous measurements of yield of PSI (by P700 absorbance changes) and that of PSII. Drought stress gradually decreased PSII electron transport, but the capacity for nonphotochemical quenching increased more slowly until there was a large decrease in leaf relative water content (where the photosynthetic rate had decreased by half or more). We identified a substantial part of PSII electron transport, which was not used by carbon assimilation or by photorespiration, which clearly indicates activities of alternative electron sinks. Decreasing the fraction of light absorbed by PSII and increasing the fraction absorbed by PSI with increasing drought stress (rather than assuming equal absorption by the two photosystems) support a proposed function of PSI cyclic electron flow to generate a proton-motive force to activate nonphotochemical dissipation of energy, and it is consistent with the observed accumulation of oxidized P700 which causes a decrease in PSI electron acceptors. Our results support the roles of alternative electron sinks (either from PSII or PSI) and cyclic electron flow in photoprotection of PSII and PSI in drought stress conditions. In future studies on plant stress, analyses of the partitioning of absorbed energy between photosystems are needed for interpreting flux through linear electron flow, PSI cyclic electron flow, along with alternative electron sinks.

Light-harvesting processes in the dynamic photosynthetic antenna

We present our perspective on the theoretical basis of light-harvesting within the photosynthetic membrane. Far from being a static structure, the photosynthetic membrane is a highly dynamic system, with protein mobility playing an important role in the damage/repair cycle of photosystem II (PSII), in balancing the input of energy between PSI and PSII, and in the photoprotection of PSII in response to a sudden excess of illumination. The concept of a photosynthetic antenna is illustrated and the state transition phenomenon is discussed as an example of purposeful antenna mobility. We discuss fluorescence recovery after photo-bleaching as a technique for visualising membrane mobility, before introducing light-induced grana membrane reorganisation as an integral part of the rapid photoprotective switch in plants. We then discuss current theoretical approaches to modelling the energy transfer dynamics of the PSII antenna: the atomistic models of intra-complex transfer and the coarse-grained approach to the inter-complex dynamics. Finally we discuss the future prospect of extending these methods, beyond the static picture of the membrane, to the dynamic PSII photosynthetic antenna.

Low photosynthetic activity is linked to changes in the organization of photosystem II in the fruit of Arum italicum

The low photosynthetic activity of fleshy green fruits is currently attributed to their special anatomy rather than to a down-regulation of photosystem II (PSII). However, it is unclear whether the organization of PSII, which is highly conserved in leaves, is also shared by non-foliar structures, such as fleshy fruits. To obtain new information on this aspect, the photosynthetic activity and the organization of PSII were investigated in the berry of Arum italicum Miller during maturation (ivory to green) and early ripening (green to yellow). The berry developed an "internal CO(2) recycling" photosynthesis; gross photosynthesis at the green stage was 25% of the leaf lamina. SDS-PAGE, BN-PAGE and 77 K spectrofluorimetry showed that the thylakoid membrane accumulated a very high amount of free LHCII trimers and only few PSII and PSI complexes. The pattern of PSII forms was similar to that of the lamina (monomers, dimers, LHCII-PSII supercomplexes), but increase in CP43-less PSII cores and low F695/F680 fluorescence ratio at room temperature indicated that PSII was less stable than in the leaf lamina. Beside effective PSII photoprotection, we propose that LHCII serves as a temporary storage of chlorophylls to provide a visual signal that fruit is not mature for seed dispersal. We conclude that the low photosynthetic activity of A. italicum berry depends on the scantiness of reaction centres and the reduced functionality of PSII.

Acclimation of Norway spruce photosynthetic apparatus to the combined effect of high irradiance and temperature

Diurnal courses of photosynthetic gas exchange parameters, chlorophyll a fluorescence characteristics and the de-epoxidation state of the xanthophyll cycle pigments (DEPS) were measured during the gradual acclimation of 4-year-old Norway spruce seedlings to different photosynthetic photon flux density (PPFD) and air temperature ( T air) regimes, simulating cloudy days with moderate T air (LI, maximum PPFD 300 μmol m −2 s −1, T air range 15–25 °C), sunny days with moderate T air (HI, maximum PPFD 1000 μmol m −2 s −1, T air range 15–25 °C) and hot sunny days (HI-HT, maximum PPFD 1000 μmol m −2 s −1, T air range 20–35 °C). The plants were acclimated inside a growth chamber and each acclimation regime lasted for 13 d. Acclimation to HI conditions led to a strong depression of the net CO 2 assimilation rates ( A N), particularly during noon and afternoon periods. Exposure to the HI-HT regime led to a further decrease of A N even during the morning period. Insufficient stomatal conductance was found to be the main reason for depressed A N under HI and HI-HT conditions. Only slight changes of the maximum photosystem II (PSII) photochemical efficiency ( F V/ F M), in the range of 0.78–0.82, supported the resistance of the Norway spruce photosynthetic apparatus against PSII photoinhibition during acclimation to both HI and HI-HT conditions. The HI plants showed increased content of xanthophyll cycle pigments (VAZ) and enhanced efficiency of thermal energy dissipation within PSII ( D) that closely correlated with the increased DEPS. In contrast, acclimation to the HI-HT regime resulted in a slight reduction of VAZ content and significantly diminished D and DEPS values during the entire day in comparison with HI plants. These results indicate a minor role of the xanthophyll cycle-mediated thermal dissipation in PSII photoprotection under elevated temperatures. The different contributions of the thermal dissipation and non-assimilatory electron transport pathways in PSII photoprotection during acclimation of the Norway spruce photosynthetic apparatus to excess irradiance and heat stresses are discussed.

Flavodiiron Proteins in Oxygenic Photosynthetic Organisms: Photoprotection of Photosystem II by Flv2 and Flv4 in Synechocystis sp. PCC 6803

BackgroundFlavodiiron proteins (FDPs) comprise a group of modular enzymes that function in oxygen and nitric oxide detoxification in Bacteria and Archaea. The FDPs in cyanobacteria have an extra domain as compared to major prokaryotic enzymes. The physiological role of cyanobacteria FDPs is mostly unknown. Of the four putative flavodiiron proteins (Flv1–4) in the cyanobacterium Synechocystis sp. PCC 6803, a physiological function in Mehler reaction has been suggested for Flv1 and Flv3.Principal FindingsWe demonstrate a novel and crucial function for Flv2 and Flv4 in photoprotection of photosystem II (PSII) in Synechocystis. It is shown that the expression of Flv2 and Flv4 is high under air level of CO2 and negligible at elevated CO2. Moreover, the rate of accumulation of flv2 and flv4 transcripts upon shift of cells from high to low CO2 is strongly dependent on light intensity. Characterization of FDP inactivation mutants of Synechocystis revealed a specific decline in PSII centers and impaired translation of the D1 protein in Δflv2 and Δflv4 when grown at air level CO2 whereas at high CO2 the Flvs were dispensable. Δflv2 and Δflv4 were also more susceptible to high light induced inhibition of PSII than WT or Δflv1 and Δflv3.SignificanceAnalysis of published sequences revealed the presence of cyanobacteria-like FDPs also in some oxygenic photosynthetic eukaryotes like green algae, mosses and lycophytes. Our data provide evidence that Flv2 and Flv4 have an important role in photoprotection of water-splitting PSII against oxidative stress when the cells are acclimated to air level CO2. It is conceivable that the function of FDPs has changed during evolution from protection against oxygen in anaerobic microbes to protection against reactive oxygen species thus making the sustainable function of oxygen evolving PSII possible. Higher plants lack FDPs and distinctly different mechanisms have evolved for photoprotection of PSII.

Photoprotection of residual functional photosystem II units that survive illumination in the absence of repair, and their critical role in subsequent recovery

Photosystem II (PSII) complexes, which split water into oxygen, protons and electrons in photosynthesis, require light but are also inactivated by it. Recovery of PSII from photoinactivation requires de novo protein synthesis. PSII in capsicum leaf segments were photoinactivated in the absence of chloroplast‐encoded protein synthesis. At large photon exposures and despite the absence of repair, a residual fraction of PSII remained functional, being ca 0.08–0.2 depending on the ease of gas exchange in the tissue. This study revealed that the residual functional PSII was photoprotected by both (1) reaction‐center quenching of excitation energy by photoinactivated PSII even when little or no PSII activity was permitted, and (2) antenna quenching, which was dependent on a trans‐thylakoid pH gradient sustained mainly by linear electron transport and facilitated by the residual functional PSII complexes themselves. Significantly, little or no contribution to photoprotection of PSII was observed from cyclic electron flow around PSI. Further, the small residual functional PSII population was critical for recovery of the photoinactivated PSII complexes. Thus, photoinactivated and residual functional PSII complexes in leaves play a mutually beneficial role in each other’s ultimate survival.

Characterization of photosynthetic pigment composition, photosystem II photochemistry and thermal energy dissipation during leaf senescence of wheat plants grown in the field

Photosynthesis, photosynthetic pigment composition and photosystem II (PSII) photochemistry during leaf senescence of wheat plants grown in the field were characterized. During leaf senescence, neoxanthin and b -carotene decreased concomitantly with chlorophyll, whereas lutein and xanthophyll cycle pigments were less affected, leading to increases in lutein/chlorophyll and xanthophyll cycle pigments/chlorophyll ratios. The chlorophyll a/b ratio also increased. With the progressing of senescence, maximal efficiency of PSII photochemstry decreased only slightly in the early morning but substantially at midday. Actual PSII efficiency, photochemical quenching, efficiency of excitation capture by open PSII centers, and the I-P phase of fluorescence induction curves decreased significantly both in the early morning and at midday and such decreases were much greater at midday than early in the morning, while the O-I phase of fluorescence induction curves remained unchanged. At the same time, non-photochemical quenching, thermal dissipation and de-epoxidation status of the xanthophyll cycle increased both early in the morning and at midday, with a greater increase at midday. Such increases indicated that the xanthophyll cycle played a role in photoprotection of PSII in senescent leaves by dissipating excess excitation energy and that D pH could be the main factor controlling thermal dissipation. Taking into consideration the substantial loss of photosynthetic capacity during senescence, our data seem to support the view that the decrease in actual PSII efficiency in senescent leaves may represent a mechanism to down-regulate photosynthetic electron transport to match the decreased CO2 assimilation capacity and avoid photodamage of PSII from excess excitation energy.