Oxygen-evolving photosystem II structures during S1-S2-S3 transitions.

Photosystem II (PSII) catalyses the oxidation of water through a four-step cycle of Si states (i = 0-4) at the Mn4CaO5 cluster1-3, during which an extra oxygen (O6) is incorporated at the S3 state to form a possible dioxygen4-7. Structural changes of the metal cluster and its environment during the S-state transitions have been studied on the microsecond timescale. Here we use pump-probe serial femtosecond crystallography to reveal the structural dynamics of PSII from nanoseconds to milliseconds after illumination with one flash (1F) or two flashes (2F). YZ, a tyrosine residue that connects the reaction centre P680 and the Mn4CaO5 cluster, showed structural changes on a nanosecond timescale, as did its surrounding amino acid residues and water molecules, reflecting the fast transfer of electrons and protons after flash illumination. Notably, one water molecule emerged in the vicinity of Glu189 of the D1 subunit of PSII (D1-E189), and was bound to the Ca2+ ion on a sub-microsecond timescale after 2F illumination. This water molecule disappeared later with the concomitant increase of O6, suggesting that it is the origin of O6. We also observed concerted movements of water molecules in the O1, O4 and Cl-1 channels and their surrounding amino acid residues to complete the sequence of electron transfer, proton release and substrate water delivery. These results provide crucial insights into the structural dynamics of PSII during S-state transitions as well as O-O bond formation.

Regulation of Leaf Angle Protects Photosystem I under Fluctuating Light in Tobacco Young Leaves.

Fluctuating light is a typical light condition in nature and can cause selective photodamage to photosystem I (PSI). The sensitivity of PSI to fluctuating light is influenced by the amplitude of low/high light intensity. Tobacco mature leaves are tended to be horizontal to maximize the light absorption and photosynthesis, but young leaves are usually vertical to diminish the light absorption. Therefore, we tested the hypothesis that such regulation of the leaf angle in young leaves might protect PSI against photoinhibition under fluctuating light. We found that, upon a sudden increase in illumination, PSI was over-reduced in extreme young leaves but was oxidized in mature leaves. After fluctuating light treatment, such PSI over-reduction aggravated PSI photoinhibition in young leaves. Furthermore, the leaf angle was tightly correlated to the extent of PSI photoinhibition induced by fluctuating light. Therefore, vertical young leaves are more susceptible to PSI photoinhibition than horizontal mature leaves when exposed to the same fluctuating light. In young leaves, the vertical leaf angle decreased the light absorption and thus lowered the amplitude of low/high light intensity. Therefore, the regulation of the leaf angle was found for the first time as an important strategy used by young leaves to protect PSI against photoinhibition under fluctuating light. To our knowledge, we show here new insight into the photoprotection for PSI under fluctuating light in nature.

Identification of a Novel Mutation Exacerbated the PSI Photoinhibition in pgr5/pgrl1 Mutants; Caution for Overestimation of the Phenotypes in Arabidopsis pgr5-1 Mutant.

PSI photoinhibition is usually avoided through P700 oxidation. Without this protective mechanism, excess light represents a potentially lethal threat to plants. PGR5 is suggested to be a major component of cyclic electron transport around PSI and is important for P700 oxidation in angiosperms. The known Arabidopsis PGR5 deficient mutant, pgr5-1, is incapable of P700 oxidation regulation and has been used in numerous photosynthetic studies. However, here it was revealed that pgr5-1 was a double mutant with exaggerated PSI photoinhibition. pgr5-1 significantly reduced growth compared to the newly isolated PGR5 deficient mutant, pgr5hope1. The introduction of PGR5 into pgr5-1 restored P700 oxidation regulation, but remained a pale-green phenotype, indicating that pgr5-1 had additional mutations. Both pgr5-1 and pgr5hope1 tended to cause PSI photoinhibition by excess light, but pgr5-1 exhibited an enhanced reduction in PSI activity. Introducing AT2G17240, a candidate gene for the second mutation into pgr5-1 restored the pale-green phenotype and partially restored PSI activity. Furthermore, a deficient mutant of PGRL1 complexing with PGR5 significantly reduced PSI activity in the double-deficient mutant with AT2G17240. From these results, we concluded that AT2G17240, named PSI photoprotection 1 (PTP1), played a role in PSI photoprotection, especially in PGR5/PGRL1 deficient mutants.

Diurnal Response of Photosystem I to Fluctuating Light Is Affected by Stomatal Conductance.

Upon a sudden transition from low to high light, electrons transported from photosystem II (PSII) to PSI should be rapidly consumed by downstream sinks to avoid the over-reduction of PSI. However, the over-reduction of PSI under fluctuating light might be accelerated if primary metabolism is restricted by low stomatal conductance. To test this hypothesis, we measured the effect of diurnal changes in stomatal conductance on photosynthetic regulation under fluctuating light in tomato (Solanum lycopersicum) and common mulberry (Morus alba). Under conditions of high stomatal conductance, we observed PSI over-reduction within the first 10 s after transition from low to high light. Lower stomatal conductance limited the activity of the Calvin-Benson-Bassham cycle and aggravated PSI over-reduction within 10 s after the light transition. We also observed PSI over-reduction after transition from low to high light for 30 s at the low stomatal conductance typical of the late afternoon, indicating that low stomatal conductance extends the period of PSI over-reduction under fluctuating light. Therefore, diurnal changes in stomatal conductance significantly affect the PSI redox state under fluctuating light. Moreover, our analysis revealed an unexpected inhibition of cyclic electron flow by the severe over-reduction of PSI seen at low stomatal conductance. In conclusion, stomatal conductance can have a large effect on thylakoid reactions under fluctuating light.

Effects of solar radiation on photosynthetic physiology of barren stalk differentiation in maize.

Barren stalks and kernel abortion are the major obstacles that hinder maize production. After many years of inbreeding, our group produced a pair of barren stalk/non-barren stalk near-isogenic lines SN98A/SN98B. Under weak light stress, the barren stalk rate is up to 98 % in SN98A but zero in SN98B. Therefore, we consider that SN98A is a weak light-sensitive inbred line whereas SN98B is insensitive. In the present study, the near-isogenic lines SN98A/SN98B were used as test materials to conduct cytological and photosynthetic physiological analyses of the physiological mechanism associated with the differences in maize barren stalk induced by weak light stress. The results showed that weak light stress increased the accumulation of reactive oxygen species (ROS), decreased the function of chloroplasts, destroyed the normal rosette structure, inhibited photosynthetic electron transport, and enhanced lipid peroxidation. The actual photochemical quantum efficiency for PSI (Y(I)) and PSII (Y(II)), relative electron transfer rate for PSI (ETR(I)) and PSII (ETR(II)), and the P700 activities decreased significantly in the leaves of SN98A and SN98B under weak light stress, where the decreases were greater in SN98A than SN98B. After 10 days of shading treatment, the O2·- production rate, H2O2 contents, the yield of regulated energy dissipation (Y(NPQ)), the donor side restriction for PSI (Y(ND)) and the quantum efficiency of cyclic electron flow photochemistry were always higher in SN98A than SN98B, and the antioxidant enzyme activities were always lower in SN98A than those in SN98B. These results show that SN98B has a stronger ability to remove ROS at its source, and maintain the integrity of the structure and function of the photosynthetic system. This self-protection mechanism is an important physiological reason for its adaptation to weak light.

Resolving diurnal dynamics of the chloroplastic glutathione redox state in Arabidopsis reveals its photosynthetically derived oxidation.

Plants are subjected to fluctuations in light intensity, and this might cause unbalanced photosynthetic electron fluxes and overproduction of reactive oxygen species (ROS). Electrons needed for ROS detoxification are drawn, at least partially, from the cellular glutathione (GSH) pool via the ascorbate-glutathione cycle. Here, we explore the dynamics of the chloroplastic glutathione redox potential (chl-EGSH) using high-temporal-resolution monitoring of Arabidopsis (Arabidopsis thaliana) lines expressing the reduction-oxidation sensitive green fluorescent protein 2 (roGFP2) in chloroplasts. This was carried out over several days under dynamic environmental conditions and in correlation with PSII operating efficiency. Peaks in chl-EGSH oxidation during dark-to-light and light-to-dark transitions were observed. Increasing light intensities triggered a binary oxidation response, with a threshold around the light saturating point, suggesting two regulated oxidative states of the chl-EGSH. These patterns were not affected in npq1 plants, which are impaired in non-photochemical quenching. Oscillations between the two oxidation states were observed under fluctuating light in WT and npq1 plants, but not in pgr5 plants, suggesting a role for PSI photoinhibition in regulating the chl-EGSH dynamics. Remarkably, pgr5 plants showed an increase in chl-EGSH oxidation during the nights following light stresses, linking daytime photoinhibition and nighttime GSH metabolism. This work provides a systematic view of the dynamics of the in vivo chloroplastic glutathione redox state during varying light conditions.

Short-lived metal-centered excited state initiates iron-methionine photodissociation in ferrous cytochrome c.

The dynamics of photodissociation and recombination in heme proteins represent an archetypical photochemical reaction widely used to understand the interplay between chemical dynamics and reaction environment. We report a study of the photodissociation mechanism for the Fe(II)-S bond between the heme iron and methionine sulfur of ferrous cytochrome c. This bond dissociation is an essential step in the conversion of cytochrome c from an electron transfer protein to a peroxidase enzyme. We use ultrafast X-ray solution scattering to follow the dynamics of Fe(II)-S bond dissociation and 1s3p (Kß) X-ray emission spectroscopy to follow the dynamics of the iron charge and spin multiplicity during bond dissociation. From these measurements, we conclude that the formation of a triplet metal-centered excited state with anti-bonding Fe(II)-S interactions triggers the bond dissociation and precedes the formation of the metastable Fe high-spin quintet state.

Extracellular electron transfer through visible light induced excited-state outer membrane C-type cytochromes of Geobacter sulfurreducens.

Dissimilatory metal-reducing bacteria (DMRB) have a variety of c-type cytochromes (OM c-cyts) intercalated in their outer membrane, and this structure serves as the physiological basis for DMRB to carry out the extracellular electron transfer processes. Using Geobacter sulfurreducens as a model DMRB, we demonstrated that visible-light illumination could alter the electronic state of OM c-cyts from the ground state to the excited state in vivo. The existence of excited-state OM c-cyts in vivo was confirmed by spectroscopy. More importantly, excited-state OM c-cyts had a more negative potential compared to their ground-state counterparts, conferring DMRB with an extra pathway to transfer electrons to semi-conductive electron acceptors. To demonstrate this, using a TiO2-coated electrode as an electron acceptor, we showed that G. sulfurreducens could directly utilise the conduction band of TiO2 as an electron acceptor under visible-light illumination (λ > 420 nm) without causing TiO2 charge separation. When G. sulfurreducens was subject to visible-light illumination, the rate of extracellular electron transfer (EET) to TiO2 accelerated by over 8-fold compared to that observed under dark conditions. Results of additional electrochemical tests provided complementary evidence to support that G. sulfurreducens utilised excited-state OM c-cyts to enhance EET to TiO2.

Cellular autofluorescence is magnetic field sensitive.

We demonstrate, by direct, single-cell imaging kinetic measurements, that endogenous autofluorescence in HeLa cells is sensitive to the application of external magnetic fields of 25 mT and less. We provide spectroscopic and mechanistic evidence that our findings can be explained in terms of magnetic field effects on photoinduced electron transfer reactions to flavins, through the radical pair mechanism. The observed magnetic field dependence is consistent with a triplet-born radical pair and a B1/2 value of 18.0 mT with a saturation value of 3.7%.

Deficiency in flavodiiron protein Flv3 promotes cyclic electron flow and state transition under high light in the cyanobacterium Synechocystis sp. PCC 6803.

Photosynthetic organisms adjust their activity to changes in irradiance by different ways, including the operation of cyclic electron flow around photosystem I (PSI) and state transitions that redistribute amounts of light energy absorbed by PSI and PSII. In dark-acclimated wild type cells of Synechocystis PCC 6803, linear electron transport was activated after the first 500 ms of illumination, while cyclic electron flow around PSI was long predominant in the mutant deficient in flavodiiron protein Flv3. Chlorophyll P700 oxidation associated with activation of linear electron flow extended in the Flv3- mutant to several tens of seconds and included a P700+ re-reduction phase. Parallel monitoring of chlorophyll fluorescence and the redox state of P700 indicated that, at low light intensity both in wild type and in the Flv3- mutant, the transient re-reduction step coincided in time with S-M fluorescence rise, which reflected state 2-state 1 transition (Kana et al., 2012). Despite variations in the initial redox state of plastoquinone pool, the oxidases-deficient mutant, succinate dehydrogenase-deficient mutant, and wild type cells did not show the S-M rise under high-intensity light until additional Flv3- mutation was introduced in these strains. Thus, the lack of available electron acceptor for PSI was the main cause for the appearance of S-M fluorescence rise under high light. It is concluded that the lack of Flv3 protein promotes cyclic electron flow around PSI and facilitates the subsequent state 2-state 1 transition in the absence of strict relation to the dark-operated pathways of plastoquinone reduction or oxidation.

Acclimation of Chlamydomonas reinhardtii to extremely strong light.

Most photosynthetic organisms are sensitive to very high light, although acclimation mechanisms enable them to deal with exposure to strong light up to a point. Here we show that cultures of wild-type Chlamydomonas reinhardtii strain cc124, when exposed to photosynthetic photon flux density 3000 µmol m-2 s-1 for a couple of days, are able to suddenly attain the ability to grow and thrive. We compared the phenotypes of control cells and cells acclimated to this extreme light (EL). The results suggest that genetic or epigenetic variation, developing during maintenance of the population in moderate light, contributes to the acclimation capability. EL acclimation was associated with a high carotenoid-to-chlorophyll ratio and slowed down PSII charge recombination reactions, probably by affecting the pre-exponential Arrhenius factor of the rate constant. In agreement with these findings, EL acclimated cells showed only one tenth of the 1O2 level of control cells. In spite of low 1O2 levels, the rate of the damaging reaction of PSII photoinhibition was similar in EL acclimated and control cells. Furthermore, EL acclimation was associated with slow PSII electron transfer to artificial quinone acceptors. The data show that ability to grow and thrive in extremely strong light is not restricted to photoinhibition-resistant organisms such as Chlorella ohadii or to high-light tolerant mutants, but a wild-type strain of a common model microalga has this ability as well.

Mimicking Photosystem I with a Transmembrane Light Harvester and Energy Transfer-Induced Photoreduction in Phospholipid Bilayers.

Photosystem I (PS I) is a transmembrane protein that assembles perpendicular to the membrane, and performs light harvesting, energy transfer, and electron transfer to a final, water-soluble electron acceptor. We present here a supramolecular model of it formed by a bicationic oligofluorene 12+ bound to the bisanionic photoredox catalyst eosin Y (EY2- ) in phospholipid bilayers. According to confocal microscopy, molecular modeling, and time dependent density functional theory calculations, 12+ prefers to align perpendicularly to the lipid bilayer. In presence of EY2- , a strong complex is formed (Ka =2.1±0.1×106 m-1 ), which upon excitation of 12+ leads to efficient energy transfer to EY2- . Follow-up electron transfer from the excited state of EY2- to the water-soluble electron donor EDTA was shown via UV-Vis absorption spectroscopy. Overall, controlled self-assembly and photochemistry within the membrane provides an unprecedented yet simple synthetic functional mimic of PS I.

Effective binding of Tb3+ and La3+ cations on the donor side of Mn-depleted photosystem II.

The interaction of Tb3+ and La3+ cations with different photosystem II (PSII) membranes (intact PSII, Ca-depleted PSII (PSII[-Ca]) and Mn-depleted PSII (PSII[-Mn]) membranes) was studied. Although both lanthanide cations (Ln3+) interact only with Ca2+-binding site of oxygen-evolving complex (OEC) in PSII and PSII(-Ca) membranes, we found that in PSII(-Mn) membranes both Ln3+ ions tightly bind to another site localized on the oxidizing side of PSII. Binding of Ln3+ cations to this site is not protected by Ca2+ and is accompanied by very effective inhibition of Mn2+ oxidation at the high-affinity (HA) Mn-binding site ([Mn2+ + H2O2] couple was used as a donor of electrons). The values of the constant for inhibition of electron transport Ki are equal to 2.10 ± 0.03 µM for Tb3+ and 8.3 ± 0.4 µM for La3+, whereas OEC inhibition constant in the native PSII membranes is 323 ± 7 µM for Tb3+. The value of Ki for Tb3+ corresponds to Ki for Mn2+ cations in the reaction of diphenylcarbazide oxidation via HA site (1.5 µM) presented in the literature. Our results suggest that Ln3+ cations bind to the HA Mn-binding site in PSII(-Mn) membranes like Mn2+ or Fe2+ cations. Taking into account the fact that Mn2+ and Fe2+ cations bind to the HA site as trivalent cations after light-induced oxidation and the fact that Mn cation bound to the HA site (Mn4) is also in trivalent state, we can suggest that valency may be important for the interaction of Ln3+ with the HA site.

Blue Light Acclimation Reduces the Photoinhibition of Phalaenopsis aphrodite (Moth Orchid).

The moth orchid is an important ornamental crop. It is very sensitive to high light irradiation due to photoinhibition. In this study, young orchid tissue culture seedlings and 2.5" potted plants pretreated under blue light (BL, λmax = 450 nm) at 100 µmol m-2 s-1 for 12 days (BL acclimation) were found to have an increased tolerance to high light irradiation. After BL acclimation, orchids had an increased anthocyanin accumulation, enhanced chloroplast avoidance, and increased chlorophyll fluorescence capacity whenever they were exposed to high light of 1000 µmol m-2 s-1 for two weeks (HL). They had higher Fv/Fm, electron transport rate (ETR), chlorophyll content, catalase activity and sucrose content when compared to the control without BL acclimation. Quantitative reverse transcription polymerase chain reaction (qRT-PCR) showed that transcript levels of phototropins, D1, RbcS, PEPCK, Catalase and SUT2 were upregulated in the BL-acclimated orchids. Consequently, BL acclimation orchids had better growth when compared to the control under long-term high light stress. In summary, this study provides a solution, i.e., BL acclimation, to reduce moth orchid photoinhibition and enhance growth before transplantation of the young tissue culture seedlings and potted plants into greenhouses, where they usually suffer from a high light fluctuation problem.

Effects of red and blue light on leaf anatomy, CO2 assimilation and the photosynthetic electron transport capacity of sweet pepper (Capsicum annuum L.) seedlings.

BACKGROUND: The red (R) and blue (B) light wavelengths are known to influence many plant physiological processes during growth and development, particularly photosynthesis. To understand how R and B light influences plant photomorphogenesis and photosynthesis, we investigated changes in leaf anatomy, chlorophyll fluorescence and photosynthetic parameters, and ribulose-1, 5-bisphosphate carboxylase/oxygenase (Rubisco) and Calvin cycle-related enzymes expression and their activities in sweet pepper (Capsicum annuum L.) seedlings exposed to four light qualities: monochromatic white (W, control), R, B and mixed R and B (RB) light with the same photosynthetic photon flux density (PPFD) of 300 µmol/m2·s. RESULTS: The results revealed that seedlings grown under R light had lower biomass accumulation, CO2 assimilation and photosystem II (PSII) electron transportation compared to plants grown under other treatments. These changes are probably due to inactivation of the photosystem (PS). Biomass accumulation and CO2 assimilation were significantly enriched in B- and RB-grown plants, especially the latter treatment. Their leaves were also thicker, and photosynthetic electron transport capacity, as well as the photosynthetic rate were enhanced. The up-regulation of the expression and activities of Rubisco, fructose-1, 6-bisphosphatase (FBPase) and glyceraldehyde-phosphate dehydrogenase (GAPDH), which involved in the Calvin cycle and are probably the main enzymatic factors contributing to RuBP (ribulose-1, 5-bisphosphate) synthesis, were also increased. CONCLUSIONS: Mixed R and B light altered plant photomorphogenesis and photosynthesis, mainly through its effects on leaf anatomy, photosynthetic electron transportation and the expression and activities of key Calvin cycle enzymes.

Regulatory mechanisms of photosynthesis light reactions in higher plants / Mechanizmy regulacyjne fazy swietlnej fotosyntezy u roslin wyzszych.

The light phase of photosynthesis is a key energy process in higher plants. Its purpose is to convert light energy into chemical one stored in ATP and NADPH molecules, which are then used to assimilate CO2 and in numerous metabolic processes. Maintaining optimal photosynthesis performance requires strict regulation of thylakoid membranes organization and rapid response to changing environmental conditions. The main factor affecting photosynthesis is light, which, if applied in excessive amounts, leads to a slowdown in the process. Therefore, plants have developed many protective mechanisms regulating the light reactions of photosynthesis and operating at the level of light energy absorption, electron transport, and the distribution and use of reducing power. These include, among others: (i) non-photochemical energy quenching regulating the amount of excitation energy delivered to the photosystems; (ii) 'state transition' process redistributing excitation energy between photosystems; (iii) redundant electron transport pathways responsible for maintaining redox balance in chloroplasts. All these mechanisms, in combination with antioxidant systems, are designed to maintain the function of the photosynthetic apparatus in adverse growth conditions.

Electron transport in Tradescantia leaves acclimated to high and low light: thermoluminescence, PAM-fluorometry, and EPR studies.

Using thermoluminescence, PAM-fluorometry, and electron paramagnetic resonance (EPR) for assaying electron transport processes in chloroplasts in situ, we have compared photosynthetic characteristics in Tradescantia fluminensis leaves grown under low light (LL, 50-125 µmol photons m-2 s-1) or high light (HL, 875-1000 µmol photons m-2 s-1) condition. We found differences in the thermoluminescence (TL) spectra of LL- and HL-acclimated leaves. The LL and HL leaves show different proportions of the Q (~ 0 °C) and B (~ 25-30 °C) bands in their TL spectra; the ratios of the "light sums" of the Q and B bands being SQ/SB ≈ 1/1 (LL) and SQ/SB ≈ 1/3 (HL). This suggests the existence of different redox states of electron carriers on the acceptor side of PSII in LL and HL leaves, which may be affected, in particular, by different capacities of their photo-reducible PQ pools. Enhanced content of PQ in chloroplasts of LL leaves may be the reason for an efficient performance of photosynthesis at low irradiance. Kinetic studies of slow induction of Chl a fluorescence and measurements of P700 photooxidation by EPR demonstrate that HL leaves have faster (about 2 times) response to switching on actinic light as compared to LL leaves grown at moderate irradiation. HL leaves also show higher non-photochemical quenching (NPQ) of Chl a fluorescence. These properties of HL leaves (faster response to light and generation of enhanced NPQ) reflect the flexibility of their photosynthetic apparatus, providing sustainability and rapid response to fluctuations of environmental light intensity and solar stress resistance. Analysis of time-courses of the EPR signals of [Formula: see text] induced by far-red (λmax = 707 nm), exciting predominantly PSI, and white light, exciting both PSI and PSII, suggests that there is a contribution of cyclic electron flow around PSI to electron flow through PSI in HL leaves. The data obtained are discussed in terms of photosynthetic apparatus sustainability of HL and LL leaves under variable irradiation conditions.

In planta study of photosynthesis and photorespiration using NADPH and NADH/NAD+ fluorescent protein sensors.

The challenge of monitoring in planta dynamic changes of NADP(H) and NAD(H) redox states at the subcellular level is considered a major obstacle in plant bioenergetics studies. Here, we introduced two circularly permuted yellow fluorescent protein sensors, iNAP and SoNar, into Arabidopsis thaliana to monitor the dynamic changes in NADPH and the NADH/NAD+ ratio. In the light, photosynthesis and photorespiration are linked to the redox states of NAD(P)H and NAD(P) pools in several subcellular compartments connected by the malate-OAA shuttles. We show that the photosynthetic increases in stromal NADPH and NADH/NAD+ ratio, but not ATP, disappear when glycine decarboxylation is inhibited. These observations highlight the complex interplay between chloroplasts and mitochondria during photosynthesis and support the suggestions that, under normal conditions, photorespiration supplies a large amount of NADH to mitochondria, exceeding its NADH-dissipating capacity, and the surplus NADH is exported from the mitochondria to the cytosol through the malate-OAA shuttle.

The water-water cycle is more effective in regulating redox state of photosystem I under fluctuating light than cyclic electron transport.

Photosynthetic electron flux from water via photosystem II (PSII) and PSI to oxygen (water-water cycle) may act as an alternative electron sink under fluctuating light in angiosperms. We measured the P700 redox kinetics and electrochromic shift signal under fluctuating light in 11 Camellia species and tobacco leaves. Upon dark-to-light transition, these Camellia species showed rapid re-oxidation of P700. However, this rapid re-oxidation of P700 was not observed when measured under anaerobic conditions, as was in experiment with tobacco performed under aerobic conditions. Therefore, photo-reduction of O2 mediated by water-water cycle was functional in these Camellia species but not in tobacco. Within the first 10 s after transition from low to high light, PSI was highly oxidized in these Camellia species but was over-reduced in tobacco leaves. Furthermore, such rapid oxidation of PSI in these Camellia species was independent of the formation of trans-thylakoid proton gradient (ΔpH). These results indicated that in addition to ΔpH-dependent photosynthetic control, the water-water cycle can protect PSI against photoinhibition under fluctuating light in these Camellia species. We here propose that the water-water cycle is an overlooked strategy for photosynthetic regulation under fluctuating light in angiosperms.

Chlororespiration Serves as Photoprotection for the Photo-Inactivated Oxygen-Evolving Complex in Zostera marina, a Marine Angiosperm.

As an alternative electron sink, chlororespiration, comprising the NAD(P)H dehydrogenase complex and plastid terminal plastoquinone oxidase, may play a significant role in sustaining the redox equilibrium between stroma and thylakoid membrane. This study identified a distinct role for chlororespiration in the marine angiosperm Zostera marina, whose oxygen-evolving complex (OEC) is prone to photo-inactivation as a result of its inherent susceptibility to excess irradiation. The strong connectivity between OEC peripheral proteins and key chlororespiratory enzymes, as demonstrated in the interaction network of differentially expressed genes, suggested that the recovery of photo-inactivated OEC was connected with chlororespiration. Chlorophyll fluorescence, transcriptome and Western blot data verified a new physiological role for chlororespiration to function as photoprotection and generate a proton gradient across the thylakoid membrane for the recovery of photo-inactivated OEC. Chlororespiration was only activated in darkness following excess irradiation exposure, which might be related to electron deficiency in the electron transport chain because of the continuous impairment of the OEC. The activation of chlororespiration in Z. marina was prone to proactivity, which was also supported by the further activation of the oxidative pentose-phosphate pathway synthesizing NADPH to meet the demand of chlororespiration during darkness. This phenomenon is distinct from the common assumption that chlororespiration is prone to consuming redundant reducing power during the short transition phase from light to dark.