ApcD is required for state transition but not involved in blue-light induced quenching in the cyanobacterium <Emphasis Type="Italic">Anabaena</Emphasis> sp. PCC7120

Pbycobilisomes （PBS） are able to transfer absorbed energy to photosystem Ⅰ and Ⅱ, and the distribution of light energy between two photosystems is regulated by state transitions. In this study we show that energy transfer from PBS to photosystem Ⅰ （PSI） requires ApcD. Cells were unable to perform state transitions in the absence of ApcD. The apcD mutant grows more slowly in light mainly absorbed by PBS, indicating that ApcD-dependent energy transfer to PSI is required for optimal growth under this condition. The apcD mutant showed normal blue-light induced quenching, suggesting that ApcD is not required for this process and state transitions are independent of blue-light induced quenching. Under nitrogen fixing condition, the growth rates of the wild type and the mutant were the same, indicating that energy transfer from PBS to PSI in heterocysts was not required for nitrogen fixation.     

State transitions and light adaptation require chloroplast thylakoid protein kinase STN7

Photosynthetic organisms are able to adjust to changing light conditions through state transitions, a process that involves the redistribution of light excitation energy between photosystem II (PSII) and photosystem I (PSI). Balancing of the light absorption capacity of these two photosystems is achieved through the reversible association of the major antenna complex (LHCII) between PSII and PSI (ref. 3). Excess stimulation of PSII relative to PSI leads to the reduction of the plastoquinone pool and the activation of a kinase; the phosphorylation of LHCII; and the displacement of LHCII from PSII to PSI (state 2). Oxidation of the plastoquinone pool by excess stimulation of PSI reverses this process (state 1). The Chlamydomonas thylakoid-associated Ser-Thr kinase Stt7, which is required for state transitions, has an orthologue named STN7 in Arabidopsis. Here we show that loss of STN7 blocks state transitions and LHCII phosphorylation. In stn7 mutant plants the plastoquinone pool is more reduced and growth is impaired under changing light conditions, indicating that STN7, and probably state transitions, have an important role in response to environmental changes.     

New mechanism revealed for light-state transition in cyanobacterium Arthrospira platensis according to 77-K fluorescence kinetics

The mechanisms of oxygen evolution and carbon fixation in oxygenic organisms depend on the equal distribution of excitation energy to photosystems Ⅰ and Ⅱ, which is regulated by a mechanism referred to as light-state transition. In this work, a novel mechanism, energy spillover from PS Ⅰ to PS Ⅱ referred to as "inverse spillover", was revealed besides "mobile phycobilisome (PBS)" and the "spillover" of energy from PS Ⅱ to PS Ⅰ in cyanobacteria. Under continuous illumination with blue light, time-dependent 77-K fluorescence spectra demonstrated heterogeneous kinetics for the PBS and photosystem components, indicating that inverse spillover and mobile PBS work successively to regulate the excitation to a balanced distribution in cyanobacterial cells under blue light. Inverse spillover and mobile PBS occur under both 100 and 300 μmol m-2 s-1 blue-light conditions but they are accelerated under the latter.     

The PSI-H subunit of photosystem I is essential for state transitions in plant photosynthesis

Photosynthesis in plants involves two photosystems responsible for converting light energy into redox processes. The photosystems, PSI and PSII, operate largely in series, and therefore their excitation must be balanced in order to optimize photosynthetic performance. When plants are exposed to illumination favouring either PSII or PSI they can redistribute excitation towards the light-limited photosystem. Long-term changes in illumination lead to changes in photosystem stoichiometry. In contrast, state transition is a dynamic mechanism that enables plants to respond rapidly to changes in illumination. When PSII is favoured (state 2), the redox conditions in the thylakoids change and result in activation of a protein kinase. The kinase phosphorylates the main light-harvesting complex (LHCII) and the mobile antenna complex is detached from PSII. It has not been clear if attachment of LHCII to PSI in state 2 is important in state transitions. Here we show that in the absence of a specific PSI subunit, PSI-H, LHCII cannot transfer energy to PSI, and state transitions are impaired.     

Cyclic electron flow around photosystem I is essential for photosynthesis

Photosynthesis provides at least two routes through which light energy can be used to generate a proton gradient across the thylakoid membrane of chloroplasts, which is subsequently used to synthesize ATP. In the first route, electrons released from water in photosystem II (PSII) are eventually transferred to NADP+ by way of photosystem I (PSI). This linear electron flow is driven by two photochemical reactions that function in series. The cytochrome b6f complex mediates electron transport between the two photosystems and generates the proton gradient (DeltapH). In the second route, driven solely by PSI, electrons can be recycled from either reduced ferredoxin or NADPH to plastoquinone, and subsequently to the cytochrome b6f complex. Such cyclic flow generates DeltapH and thus ATP without the accumulation of reduced species. Whereas linear flow from water to NADP+ is commonly used to explain the function of the light-dependent reactions of photosynthesis, the role of cyclic flow is less clear. In higher plants cyclic flow consists of two partially redundant pathways. Here we have constructed mutants in Arabidopsis thaliana in which both PSI cyclic pathways are impaired, and present evidence that cyclic flow is essential for efficient photosynthesis.     

Hydrogen ions directly regulating the oligomerization state of Photosystem I in intact <Emphasis Type="Italic">Spirulina platensis</Emphasis> cells

H^＋ concentration induced-monomerization or trimerization of photosystem Ⅰ （PSI） in cyanobacteria has never been directly observed. In this work, taking characteristic spectra for the trimers and monomers as the indicators, it was experimentally demonstrated that H^＋ could induce the oligomeric changes of PSI reaction centers in the intact Spirulina platensis cells and also in the isolated thylakoid membrane complexes. Especially, the higher concentration of H^＋ would induce the monomerization while the lower the trimerization, suggesting the electrostatic interaction should be mainly responsible for changes in the oligomeric state of PSI in Spirulina platensis.     

Low-light-adapted Prochlorococcus species possess specific antennae for each photosystem

Prochlorococcus, the most abundant genus of photosynthetic organisms, owes its remarkably large depth distribution in the oceans to the occurrence of distinct genotypes adapted to either low- or high-light niches. The pcb genes, encoding the major chlorophyll-binding, light-harvesting antenna proteins in this genus, are present in multiple copies in low-light strains but as a single copy in high-light strains. The basis of this differentiation, however, has remained obscure. Here we show that the moderate low-light-adapted strain Prochlorococcus sp. MIT 9313 has one iron-stress-induced pcb gene encoding an antenna protein serving photosystem I (PSI)--comparable to isiA genes from cyanobacteria--and a constitutively expressed pcb gene encoding a photosystem II (PSII) antenna protein. By comparison, the very low-light-adapted strain SS120 has seven pcb genes encoding constitutive PSI and PSII antennae, plus one PSI iron-regulated pcb gene, whereas the high-light-adapted strain MED4 has only a constitutive PSII antenna. Thus, it seems that the adaptation of Prochlorococcus to low light environments has triggered a multiplication and specialization of Pcb proteins comparable to that found for Cab proteins in plants and green algae.     

Direct electrochemistry and electrocatalysis of horseradish peroxidase in MnO2 nanosheet film

A novel material MnO2 nanosheet has been used as the support matrix for the immobilization of horseradish peroxidase （HRP）. HRP entrapped in MnO2 nanosheet film exhibits facile direct electron transfer with the electron transfer rate constant of 6.86 s^-1. The HRP/MnO2 nanosheet film gives a reversible redox couple with the apparent formal peak potential （E^0＇） of -0.315 V （vs. Ag/AgCl） in pH 6.5 phosphate buffer solution （PBS）. The formal potential E^0＇ of HRP shifts linearly with pH with a slope of -53.75 mV.pH^-1, denoting that an electron transfer accompanies single-proton transportation. The immobilized HRP shows an electrocatslytic activity to the reduction of H2O2. The response time of the biosensor for H2O2 is less than 3 s, and the detection limit is 0.21 μmol · L^-1 based on signal/noise = 3.     

A giant chlorophyll-protein complex induced by iron deficiency in cyanobacteria

Cyanobacteria are abundant throughout most of the world's water bodies and contribute significantly to global primary productivity through oxygenic photosynthesis. This reaction is catalysed by two membrane-bound protein complexes, photosystem I (PSI) and photosystem II (PSII), which both contain chlorophyll-binding subunits functioning as an internal antenna. In addition, phycobilisomes act as peripheral antenna systems, but no additional light-harvesting systems have been found under normal growth conditions. Iron deficiency, which is often the limiting factor for cyanobacterial growth in aquatic ecosystems, leads to the induction of additional proteins such as IsiA (ref. 3). Although IsiA has been implicated in chlorophyll storage, energy absorption and protection against excessive light, its precise molecular function and association to other proteins is unknown. Here we report the purification of a specific PSI-IsiA supercomplex, which is abundant under conditions of iron limitation. Electron microscopy shows that this supercomplex consists of trimeric PSI surrounded by a closed ring of 18 IsiA proteins binding around 180 chlorophyll molecules. We provide a structural characterization of an additional chlorophyll-containing, membrane-integral antenna in a cyanobacterial photosystem.     

Low concentrations of NaHSO3 enhance NAD(P)H dehydrogenase-dependent cyclic photophosphorylation and alleviate the oxidative damage to improve photosynthesis in tobacco

Bisulfite at low concentrations(L-NaHSO3) increases cyclic electron transport around photosystem I(PSI) and photosynthesis.However,little is known regarding the detailed contribution of cyclic electron transport to the promoted photosynthesis by L-NaHSO3.In the present work,we used tobacco mutant defective in ndhC-ndhK-ndhJ(ndhCKJ) to investigate the role of NAD(P)H dehydrogenase(NDH)-dependent cyclic electron transport around PSI in an increase in photosynthesis by L-NaHSO3.After the treatment of tobacco leaves with L-NaHSO3(10 μmol L-1),the NDH-dependent cyclic electron transport,monitored by a transient post-illumination increase in Chl fluorescence and the amount of NDH,was notably up-regulated in wild type(WT).The NDH-dependent cyclic electron transport was severely impaired in ndhCKJ and was not significantly affected by treatment with L-NaHSO3.Accordingly,the NDH-dependent transthylakoid membrane proton gradient(pH),as reflected by the slow phase of millisecond-delayed light emission(ms-DLE),was increased by L-NaHSO3 in WT,but not in ndhCKJ;the enhancement of cyclic photophosphorylation(PSP) activity by L-NaHSO3 was more obvious in WT than ndhCKJ.The accumulation of both superoxide and hydrogen peroxide was reduced in WT when subjected to L-NaHSO3 treatment,but not in ndhCKJ.Furthermore,the increase of photosynthetic O 2 evolution rate by L-NaHSO3 was more significant in WT than in ndhCKJ.We therefore conclude that L-NaHSO3 alleviates the photo-oxidative damage by the enhancement of NDH-dependent cyclic PSP,thereby improving photosynthesis.     

Crystal structure of plant photosystem I

Oxygenic photosynthesis is the principal producer of both oxygen and organic matter on Earth. The conversion of sunlight into chemical energy is driven by two multisubunit membrane protein complexes named photosystem I and II. We determined the crystal structure of the complete photosystem I (PSI) from a higher plant (Pisum sativum var. alaska) to 4.4 A resolution. Its intricate structure shows 12 core subunits, 4 different light-harvesting membrane proteins (LHCI) assembled in a half-moon shape on one side of the core, 45 transmembrane helices, 167 chlorophylls, 3 Fe-S clusters and 2 phylloquinones. About 20 chlorophylls are positioned in strategic locations in the cleft between LHCI and the core. This structure provides a framework for exploration not only of energy and electron transfer but also of the evolutionary forces that shaped the photosynthetic apparatus of terrestrial plants after the divergence of chloroplasts from marine cyanobacteria one billion years ago.     

pH Dependence of Chlorophyll States, Protein Structures and Function of the PSII Membranes

IntroductionPhotosystem II ( PSII) is a pigment- proteincomplex in the thylakoid membrane. Its reactioncenter( PSII- RC) ,which is composed of D1 andD2 proteins,generates the highly positive oxidantrequired for the oxidation of water by light- drivencharge separation. Water oxidation occurs at theMn4cluster positioned atthe center of the oxygen-evolving complex on the lumenal surface of PSII.In green plants,the highly reactive Mn4cluster isshielded by a number of extrinsic proteins( 33…     

Deletion of an electron donor-binding subunit of the NDH-1 complex,NdhS, results in a heat-sensitive growth phenotype in Synechocystis sp. PCC 6803

In cyanobacteria and higher plants, NdhS is suggested to be an electron donor-binding subunit of NADPH dehydrogenase （NDH-1） complexes and its absence impairs NDH-l-dependent cyclic electron trans- port around photosystem I （NDH-CET）. Despite significant advances in the study of NdhS during recent years, its functional role in resisting heat stress is poorly understood. Here, our results revealed that the absence of NdhS resulted in a serious heat-sensitive growth phenotype in the uni- cellular cyanobacterium Synechocystis sp. strain PCC 6803. Furthermore, the rapid and significant increase in NDH-CET caused by heat treatment was completely abolished, and the repair of photosystem II under heat stress conditions was greatly impaired when compared to that of other photosynthetic apparatus in the thylakoid membrane. We therefore conclude that NdhS plays an important role in resistance to heat stress, possibly by stabilizing the electron input module of cyanobacterial NDH-1 complexes.     

Iron deficiency induces the formation of an antenna ring around trimeric photosystem I in cyanobacteria

Although iron is the fourth most abundant element in the Earth's crust, its concentration in the aquatic ecosystems-particularly the open oceans-is sufficiently low to limit photosynthetic activity and phytoplankton growth. Cyanobacteria, a major class of phytoplankton, respond to iron deficiency by expressing the 'iron-stress-induced' gene, isiA(ref. 3). The protein encoded by this gene has an amino-acid sequence that shows significant homology with one of the chlorophyll a-binding proteins (CP43) of photosystem II (PSII). The precise function of the CP43-like protein, here called CP43', has not been elucidated, although there have been many suggestions. Here we show that CP43' associates with photosystem I (PSI) to form a complex that consists of a ring of 18 CP43' molecules around a PSI trimer. This significantly increases the size of the light-harvesting system of PSI. The utilization of a PSII-like protein as an extra antenna for PSI emphasises the flexibility of cyanobacterial light-harvesting systems, and seems to be a strategy which compensates for the lowering of phycobilisome and PSI levels in response to iron deficiency.     

Transient Effects of Salt Shock on the Photosynthetic Machinery in Synechocystis sp. PCC 6803

IntroductionSalt stress is one of the main detrimental factors inthe environment that limit the growth andproductivity of plants.Salt stress causessignificant decreases in photosynthetic activity,such as the electron transport[1,2 ] ,but themechanisms by which salt stress inhibitsphotosynthesis remain poorly understood[3] .　Cyanobacteria are prokaryotes that performoxygenic photosynthesis using a photosyntheticapparatus similar to that in the chloroplasts ofgreen algae and higher plants[4 ] .…     

模拟酸雨对4种木本植物叶绿素荧光特性的影响

文中研究了不同pH（2.5、4.5）模拟酸雨对4种鼎湖山南亚热带森林优势树种:木荷（Schima superba）、黧蒴（Castanopsis fissa）、肖蒲桃（Acmena acuminatissima）、黄果厚壳桂（Cryptocarya concinna）叶片叶绿素荧光特性的影响，结果表明: 4种木本植物离体叶片经pH 4.5模拟酸雨处理7 d后未出现可见伤害， F_v/F_m与对照组相比无显著差异. 而在经pH 2.5 模拟酸雨处理7 d后，叶片均出现了明显的可见伤害，其中黄果厚壳桂的可见伤害程度和PSII受损程度较其他3种植物小. 经pH 2.5或pH 4.5的模拟酸雨处理后，黄果厚壳桂叶片光PSII电子传递速率（ETRII）大幅度降低，同时PSI环式电子流被激发，这可能是其提高自身抗酸能力的主动调节机制. 研究结果显示,黄果厚壳桂的抗酸雨胁迫的能力高于其他3种木本植物，酸沉降可能会提高南亚热带森林群落中黄果厚壳桂的相对优势地位.     

Mutagenesis of Ser24 of Cytochrome b559 α Subunit Affects PSII Acitivies in Chlamydomonas reinhardtii

In order to study the functions of cytochrome b559 (Cyt b559) in photosystem two (PSII) activity, mutant S24F of Chlamydomonas reinhardtii was constructed using site directed mutagenesis, in which Serine24 (Ser24) locating downstream of Histidine23 (His23) in α subunit of Cyt b559 was replaced by Phenylalanine (Phe). Physiological and biochemical analysis showed that mutant S24F could be grown photoautotrophically or photoheterotrophically. However, their growth rate was slower either on HSM or TAP medium than that of the control; Analysis of PSII activity revealed that its oxygen evolution was about 71% of wild type (WT); The Photochemical efficiency of PSII (Fv/Fm) of S24F was reduced 0.23 compared with WT; S24F was more sensitive to strong light irradiance than the wild type; Furthermore, SDS-PAGE and Western-blotting analysis indicated that the expression levels of α subunit of Cyt b559, LHCII and PsbO of S24F were a little less than those of the wild type. Overall, these data suggests that Ser24 plays a significant role in making Cyt b559 structure maintain PSII complex activity of oxygen evolution although it is not directly bound to heme group.     

海滨锦葵的耐冷性及低温弱光对其光系统的伤害

以海滨锦葵为实验材料,研究不同低温处理对海滨锦葵光合作用的伤害,探求海滨锦葵对低温胁迫的敏感温度,以及低温弱光对海滨锦葵的伤害．结果表明：在低温胁迫下,海滨锦葵的净光合速率（Pn）、光系统II实际光化学效率（ФPSII）、最大光化学效率（Fv／Fm）显著下降,说明随着温度的降低,海滨锦葵光化学活性受到抑制,13℃是其低温胁迫下的临界温度．低温弱光（6℃、200μmol·m^-2s^-1）处理4h后Fv/Fm下降了2．5％,而光系统I活性（△I／I0）下降了18．5％,说明在低温弱光条件下,海滨锦葵光系统I受到的伤害高于光系统II;在恢复过程中,光系统II在8h基本完全恢复,而光系统I和净光合速率在48h后仍没有恢复到正常水平,说明PSI的恢复速率成了光合作用的主要限制因素．     

The composition and distribution of metal clusters in the MoFe protein from a nifZ deletion strain （DJ 194） of Azotobacter vinelandii

Through the anaerobic chromatography on the columns of DEAE 52, Q-Sepharose and Sephacryl S-200, a nitrogenase MoFe protein (△nifZ Av1) was obtained from a nifZ deleted mutant of Azotobacter vinelandii (stain DJ194).The results of Western blotting after anoxic native electrophoresis and SDS-PAGE showed that △nifZ Av1 was similar to wild type MoFe protein (OP Av1) at the electrophoretic mobility, molecular weight and subunit composition. Furthermore, △nifZ Avl was also similar to OP Av1 at the molybdenum content, EPR signal (g≈4.3, 3.65 and 2.01), and the molar extinction coefficient (△ε) of circular dichroism (CD)at 660 nm region. All of these indicated that, besides having the same α2β2 composition as OP Av1, the △nifZ Av1 also contained equal amount of reductive FeMoco in the spin state of S=3/2 to OP Av1. However, the iron content and substrate (C2H2, H^ and N2)-reduction activity of △nifZ Av1 were 74% and 46%-50% of those of OP Av1, respectively. Furthermore, the △ε at around 450 nm, which reflects P-cluster in Av1, was obviously lower than that of OP Av1. It suggested that the difference between △nifZ Avl and OP Av1 resulted from P-cluster rather than FeMoco, and from the half number of P-cluster in △nifZ Av1, but the composition or redoxstate of P-cluster in △nifZ Av1 were not changed. Thus it could propose that △nifZ Av1 is composed of two different αβsubunit pairs. One is a FeMoco-and P-cluster-containing pair, and the other is a P-cluster-deficient but FeMoco-containing pair. Since the deletion of nifZ gene leads to the deficiency of only one of two P-clusters in a α2β2 tetramer, the assembly of P-cluster may not simply depend on one gene product, and so a possible mechanism of NifZ is supposed here.     

铯对小麦叶片光合特性影响的研究

随着环境中铯(Cs)污染日益恶化,铯对生物毒害作用的研究受到越来越多的关注.为了探讨铯对植物光合作用的影响,在石英砂和Hoagland营养液培养体系下,从三叶期开始用浓度为0、0.5、1、5、10、20 mmol·L-1的133Cs+[CsCl]处理小麦(Triticum aestivum L.)幼苗.在处理后的第0、7、14、21、28 d时,检测小麦光合特性的改变.结果表明:用0.5 mmol·L-1和1 mmol·L-1低浓度133Cs+处理7 d后,小麦叶片的叶绿素含量、叶圆片放氧活性、类囊体膜电子传递活性均显著高于对照;但随133Cs+浓度的增加及处理时间的延长,这些参数的数值均显著下降.此外,光合系统II(PSII)活性的下降速率高于光合系统I(PSI)下降速率,说明PSII相对于PSI更容易受到133Cs+的胁迫伤害.这些结果表明133Cs+对小麦的毒害效应受时间和浓度的双重制约.低浓度133Cs+可以促进小麦光合作用,而高浓度133Cs+则显著抑制小麦光合作用.