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Protein Science : a Publication of the... May 2020Photosystem II (PSII) is a membrane-spanning, multi-subunit pigment-protein complex responsible for the oxidation of water and the reduction of plastoquinone in oxygenic... (Review)
Review
Photosystem II (PSII) is a membrane-spanning, multi-subunit pigment-protein complex responsible for the oxidation of water and the reduction of plastoquinone in oxygenic photosynthesis. In the present review, the recent explosive increase in available structural information about the PSII core complex based on X-ray crystallography and cryo-electron microscopy is described at a level of detail that is suitable for a future structure-based analysis of light-harvesting processes. This description includes a proposal for a consistent numbering scheme of protein-bound pigment cofactors across species. The structural survey is complemented by an overview of the state of affairs in structure-based modeling of excitation energy transfer in the PSII core complex with emphasis on electrostatic computations, optical properties of the reaction center, the assignment of long-wavelength chlorophylls, and energy trapping mechanisms.
Topics: Crystallography, X-Ray; Light; Models, Molecular; Photosystem II Protein Complex; Protein Conformation
PubMed: 32067287
DOI: 10.1002/pro.3841 -
Plant, Cell & Environment Feb 2018Repair of photosystem II (PSII) during photoinhibition involves replacement of photodamaged D1 protein by newly synthesized D1 protein. In this review, we summarize... (Review)
Review
Repair of photosystem II (PSII) during photoinhibition involves replacement of photodamaged D1 protein by newly synthesized D1 protein. In this review, we summarize evidence for the indispensability of ATP in the degradation and synthesis of D1 during the repair of PSII. Synthesis of one molecule of the D1 protein consumes more than 1,300 molecules of ATP equivalents. The degradation of photodamaged D1 by FtsH protease also consumes approximately 240 molecules of ATP. In addition, ATP is required for several other aspects of the repair of PSII, such as transcription of psbA genes. These requirements for ATP during the repair of PSII have been demonstrated by experiments showing that the synthesis of D1 and the repair of PSII are interrupted by inhibitors of ATP synthase and uncouplers of ATP synthesis, as well as by mutation of components of ATP synthase. We discuss the contribution of cyclic electron transport around photosystem I to the repair of PSII. Furthermore, we introduce new terms relevant to the regulation of the PSII repair, namely, "ATP-dependent regulation" and "redox-dependent regulation," and we discuss the possible contribution of the ATP-dependent regulation of PSII repair under environmental stress.
Topics: Adenosine Triphosphate; Electron Transport; Light; Photosystem II Protein Complex; Plants
PubMed: 29210214
DOI: 10.1111/pce.13108 -
Frontiers in Plant Science 2022Light intensity is highly heterogeneous in nature, and plants have evolved a series of strategies to acclimate to dynamic light due to their immobile lifestyles....
Light intensity is highly heterogeneous in nature, and plants have evolved a series of strategies to acclimate to dynamic light due to their immobile lifestyles. However, it is still unknown whether there are differences in photoprotective mechanisms among different light-demanding plants in response to dynamic light, and thus the role of non-photochemical quenching (NPQ), electron transport, and light energy allocation of photosystems in photoprotection needs to be further understood in different light-demanding plants. The activities of photosystem II (PSII) and photosystem I (PSI) in shade-tolerant species , intermediate species , and sun-demanding species were comparatively measured to elucidate photoprotection mechanisms in different light-demanding plants under dynamic light. The results showed that the NPQ and PSII maximum efficiency ( '/ ') of were higher than the other two species under dynamic high light. Meanwhile, cyclic electron flow (CEF) of sun plants is larger under transient high light conditions since the slope of post-illumination, P700 dark reduction rate, and plastoquinone (PQ) pool were greater. NPQ was more active and CEF was initiated more readily in shade plants than the two other species under transient light. Moreover, sun plants processed higher quantum yield of PSII photochemistry (Φ), quantum yield of photochemical energy conversion [Y(I)], and quantum yield of non-photochemical energy dissipation due to acceptor side limitation (Y(NA), while the constitutive thermal dissipation and fluorescence (Φ) and quantum yield of non-photochemical energy dissipation due to donor side limitation [Y(ND)] of PSI were higher in shade plants. These results suggest that sun plants had higher NPQ and CEF for photoprotection under transient high light and mainly allocated light energy through Φ and Φ, while shade plants had a higher Φ and a larger heat dissipation efficiency of PSI donor. Overall, it has been demonstrated that the photochemical efficiency and photoprotective capacity are greater in sun plants under transient dynamic light, while shade plants are more sensitive to transient dynamic light.
PubMed: 35463455
DOI: 10.3389/fpls.2022.819843 -
Frontiers in Plant Science 2019As a light-harvesting organelle, the chloroplast inevitably produces a substantial amount of reactive oxygen species (ROS) primarily through the photosystems. These ROS,... (Review)
Review
As a light-harvesting organelle, the chloroplast inevitably produces a substantial amount of reactive oxygen species (ROS) primarily through the photosystems. These ROS, such as superoxide anion, hydrogen peroxide, hydroxyl radical, and singlet oxygen, are potent oxidizing agents, thereby damaging the photosynthetic apparatus. On the other hand, it became increasingly clear that ROS act as beneficial tools under photo-oxidative stress conditions by stimulating chloroplast-nucleus communication, a process called retrograde signaling (RS). These ROS-mediated RS cascades appear to participate in a broad spectrum of plant physiology, such as acclimation, resistance, programmed cell death (PCD), and growth. Recent reports imply that ROS-driven oxidation of RS-associated components is essential in sensing and responding to an increase in ROS contents. ROS appear to activate RS pathways reversible or irreversible oxidation of sensor molecules. This review provides an overview of the emerging perspective on the topic of "oxidative modification-associated retrograde signaling."
PubMed: 32038693
DOI: 10.3389/fpls.2019.01729 -
Frontiers in Plant Science 2023Phycobilisomes serve as a light-harvesting antenna of both photosystem I (PSI) and II (PSII) in cyanobacteria, yet direct energy transfer from phycobilisomes to PSI is...
Phycobilisomes serve as a light-harvesting antenna of both photosystem I (PSI) and II (PSII) in cyanobacteria, yet direct energy transfer from phycobilisomes to PSI is not well documented. Here we recorded picosecond time-resolved fluorescence at wavelengths of 605-760 nm in isolated photosystem I (PSI), phycobilisomes and intact cells of a PSII-deficient mutant of sp. PCC 6803 at 77 K to study excitation energy transfer and trapping. By means of a simultaneous target analysis of the kinetics of isolated complexes and whole cells, the pathways and dynamics of energy transfer and were established. We establish that the timescale of the slowest equilibration between different terminal emitters in the phycobilisome is ≈800 ps. It was estimated that the terminal emitter in about 40% of the phycobilisomes transfers its energy with a rate constant of 42 ns to PSI. This energy transfer rate is higher than the rates of equilibration within the phycobilisome - between the rods and the core or between the core cylinders - and is evidence for the existence of specific phycobilisome-PSI interactions. The rest of the phycobilisomes remain unconnected or slowly transferring energy to PSI.
PubMed: 38078099
DOI: 10.3389/fpls.2023.1293813 -
Free Radical Biology & Medicine Aug 2019The ability to harvest light to drive chemical reactions and gain energy provided microbes access to high energy electron donors which fueled primary productivity,... (Review)
Review
The ability to harvest light to drive chemical reactions and gain energy provided microbes access to high energy electron donors which fueled primary productivity, biogeochemical cycles, and microbial evolution. Oxygenic photosynthesis is often cited as the most important microbial innovation-the emergence of oxygen-evolving photosynthesis, aided by geologic events, is credited with tipping the scale from a reducing early Earth to an oxygenated world that eventually lead to complex life. Anoxygenic photosynthesis predates oxygen-evolving photosynthesis and played a key role in developing and fine-tuning the photosystem architecture of modern oxygenic phototrophs. The release of oxygen as a by-product of metabolic activity would have caused oxidative damage to anaerobic microbiota that evolved under the anoxic, reducing conditions of early Earth. Photosynthetic machinery is particularly susceptible to the adverse effects of oxygen and reactive oxygen species and these effects are compounded by light. As a result, phototrophs employ additional detoxification mechanisms to mitigate oxidative stress and have evolved alternative oxygen-dependent enzymes for chlorophyll biosynthesis. Phylogenetic reconstruction studies and biochemical characterization suggest photosynthetic reactions centers, particularly in Cyanobacteria, evolved to both increase efficiency of electron transfer and avoid photodamage caused by chlorophyll radicals that is acute in the presence of oxygen. Here we review the oxygen and reactive oxygen species detoxification mechanisms observed in extant anoxygenic and oxygenic photosynthetic bacteria as well as the emergence of these mechanisms over evolutionary time. We examine the distribution of phototrophs in modern systems and phylogenetic reconstructions to evaluate the emergence of mechanisms to mediate oxidative damage and highlight changes in photosystems and reaction centers, chlorophyll biosynthesis, and niche space in response to oxygen production. This synthesis supports an emergence of HS-driven anoxygenic photosynthesis in Cyanobacteria prior to the evolution of oxygenic photosynthesis and underscores a role for the former metabolism in fueling fine-tuning of the oxygen evolving complex and mechanisms to repair oxidative damage. In contrast, we note the lack of elaborate mechanisms to deal with oxygen in non-cyanobacterial anoxygenic phototrophs suggesting these microbes have occupied similar niche space throughout Earth's history.
Topics: Biological Evolution; Cyanobacteria; Oxidation-Reduction; Oxygen; Photosynthesis; Phototrophic Processes
PubMed: 31078729
DOI: 10.1016/j.freeradbiomed.2019.05.003 -
Frontiers in Plant Science 2022In this study, the differences in chlorophyll fluorescence transient (OJIP) and modulated 820 nm reflection (MR) of cucumber leaves were probed to demonstrate an insight...
In this study, the differences in chlorophyll fluorescence transient (OJIP) and modulated 820 nm reflection (MR) of cucumber leaves were probed to demonstrate an insight into the precise influence of melatonin (MT) on cucumber photosystems under low temperature stress. We pre-treated cucumber seedlings with different levels of MT (0, 25, 50, 100, 200, and 400 μmol · L) before imposing low temperature stress (10 °C/6 °C). The results indicated that moderate concentrations of MT had a positive effect on the growth of low temperature-stressed cucumber seedlings. Under low temperature stress conditions, 100 μmol · L (MT 100) improved the performance of the active photosystem II (PSII) reaction centers (PIabs), the oxygen evolving complex activity (OEC centers) and electron transport between PSII and PSI, mainly by decreasing the L-band, K-band, and G-band, but showed differences with different duration of low temperature stress. In addition, these indicators related to quantum yield and energy flux of PSII regulated by MT indicated that MT (MT 100) effectively protected the electron transport and energy distribution in the photosystem. According to the results of ≥ 1 and MR signals, MT also affected PSI activity. MT 100 decreased the minimal value of MR/MR and the oxidation rate of plastocyanin (PC) and PSI reaction center (P700) ( ), while increased △MR/MR and deoxidation rates of PC and P ( ). The loss of the slow phase of MT 200 and MT 400-treated plants in the MR kinetics was due to the complete prevention of electron movement from PSII to re-reduce the PC and P700 . These results suggest that appropriate MT concentration (100 μmol · L) can improve the photosynthetic performance of PS II and electron transport from primary quinone electron acceptor (Q) to secondary quinone electron acceptor (Q), promote the balance of energy distribution, strengthen the connectivity of PSI and PSII, improve the electron flow of PSII Q to PC and P from reaching PSI by regulating multiple sites of electron transport chain in photosynthesis, and increase the pool size and reduction rates of PSI in low temperature-stressed cucumber plants, All these modifications by MT 100 treatment promoted the photosynthetic electron transfer smoothly, and further restored the cucumber plant growth under low temperature stress. Therefore, we conclude that spraying MT at an appropriate concentration is beneficial for protecting the photosynthetic electron transport chain, while spraying high concentrations of MT has a negative effect on regulating the low temperature tolerance in cucumber.
PubMed: 36407604
DOI: 10.3389/fpls.2022.1029854 -
Biochimica Et Biophysica Acta.... Sep 2018The ability of photosynthetic organisms to use the sun's light as a sole source of energy sustains life on our planet. Photosystems I (PSI) and II (PSII) are large,...
The ability of photosynthetic organisms to use the sun's light as a sole source of energy sustains life on our planet. Photosystems I (PSI) and II (PSII) are large, multi-subunit, pigment-protein complexes that enable photosynthesis, but this intriguing process remains to be explained fully. Currently, crystal structures of these complexes are available for thermophilic prokaryotic cyanobacteria. The mega-Dalton trimeric PSI complex from thermophilic cyanobacterium, Thermosynechococcus elongatus, was solved at 2.5 Å resolution with X-ray crystallography. That structure revealed the positions of 12 protein subunits (PsaA-F, PsaI-M, and PsaX) and 127 cofactors. Although mesophilic organisms perform most of the world's photosynthesis, no well-resolved trimeric structure of a mesophilic organism exists. Our research model for a mesophilic cyanobacterium was Synechocystis sp. PCC6803. This study aimed to obtain well-resolved crystal structures of [1] a monomeric PSI with all subunits, [2] a trimeric PSI with a reduced number of subunits, and [3] the full, trimeric wild-type PSI complex. We only partially succeeded with the first two structures, but we successfully produced the trimeric PSI structure at 2.5 Å resolution. This structure was comparable to that of the thermophilic species, but we provided more detail. The PSI trimeric supercomplex consisted of 33 protein subunits, 72 carotenoids, 285 chlorophyll a molecules, 51 lipids, 9 iron-sulfur clusters, 6 plastoquinones, 6 putative calcium ions, and over 870 water molecules. This study showed that the structure of the PSI in Synechocystis sp. PCC6803 differed from previously described PSI structures. These findings have broadened our understanding of PSI structure.
Topics: Chlorophyll; Chlorophyll A; Crystallography, X-Ray; Models, Molecular; Photosynthesis; Photosystem I Protein Complex; Protein Conformation; Protein Subunits; Structure-Activity Relationship; Synechocystis
PubMed: 29414678
DOI: 10.1016/j.bbabio.2018.02.002 -
Communications Biology Dec 2021Water molecules play a pivotal functional role in photosynthesis, primarily as the substrate for Photosystem II (PSII). However, their importance and contribution to...
Water molecules play a pivotal functional role in photosynthesis, primarily as the substrate for Photosystem II (PSII). However, their importance and contribution to Photosystem I (PSI) activity remains obscure. Using a high-resolution cryogenic electron microscopy (cryo-EM) PSI structure from a Chlamydomonas reinhardtii temperature-sensitive photoautotrophic PSII mutant (TSP4), a conserved network of water molecules - dating back to cyanobacteria - was uncovered, mainly in the vicinity of the electron transport chain (ETC). The high-resolution structure illustrated that the water molecules served as a ligand in every chlorophyll that was missing a fifth magnesium coordination in the PSI core and in the light-harvesting complexes (LHC). The asymmetric distribution of the water molecules near the ETC branches modulated their electrostatic landscape, distinctly in the space between the quinones and FX. The data also disclosed the first observation of eukaryotic PSI oligomerisation through a low-resolution PSI dimer that was comprised of PSI-10LHC and PSI-8LHC.
Topics: Chlamydomonas; Cryoelectron Microscopy; Mutation; Photosystem I Protein Complex; Photosystem II Protein Complex; Temperature
PubMed: 34887518
DOI: 10.1038/s42003-021-02911-7 -
Planta Nov 2016The photosystem I/II ratio increased when antenna size was enlarged by transient induction of CAO in chlorophyll b -less mutants, thus indicating simultaneous regulation...
The photosystem I/II ratio increased when antenna size was enlarged by transient induction of CAO in chlorophyll b -less mutants, thus indicating simultaneous regulation of antenna size and photosystem I/II stoichiometry. Regulation of antenna size and photosystem I/II stoichiometry is an indispensable strategy for plants to acclimate to changes to light environments. When plants grown in high-light conditions are transferred to low-light conditions, the peripheral antennae of photosystems are enlarged. A change in the photosystem I/II ratio is also observed under the same light conditions. However, our knowledge of the correlation between antenna size modulation and variation in photosystem I/II stoichiometry remains limited. In this study, chlorophyll a oxygenase was transiently induced in Arabidopsis thaliana chlorophyll b-less mutants, ch1-1, to alter the antenna size without changing environmental conditions. In addition to the accumulation of chlorophyll b, the levels of the peripheral antenna complexes of both photosystems gradually increased, and these were assembled to the core antenna of both photosystems. However, the antenna size of photosystem II was greater than that of photosystem I. Immunoblot analysis of core antenna proteins showed that the number of photosystem I increased, but not that of photosystem II, resulting in an increase in the photosystem I/II ratio. These results clearly indicate that antenna size adjustment was coupled with changes in photosystem I/II stoichiometry. Based on these results, the physiological importance of simultaneous regulation of antenna size and photosystem I/II stoichiometry is discussed in relation to acclimation to light conditions.
Topics: Arabidopsis; Arabidopsis Proteins; Chlorophyll; Chlorophyll A; Chromatography, High Pressure Liquid; Electrophoresis, Polyacrylamide Gel; Fluorescence; Gene Expression Regulation, Plant; Immunoblotting; Light-Harvesting Protein Complexes; Models, Biological; Oxygenases; Photosynthesis; Photosystem I Protein Complex; Photosystem II Protein Complex; RNA, Messenger; Substrate Specificity; Temperature; Transformation, Genetic
PubMed: 27394155
DOI: 10.1007/s00425-016-2568-5