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1.
Jan M. Anderson 《Photosynthesis research》1992,34(3):341-357
The cytochrome b
6
f complex occupies a central position in photosynthetic electron transport and proton translocation by linking PS II to PS I in linear electron flow from water to NADP+, and around PS I for cyclic electron flow. Cytochrome b
6
f complexes are uniquely located in three membrane domains: the appressed granal membranes, the non-appressed stroma thylakoids and end grana membranes, and also the non-appressed grana margins, in contrast to the marked lateral heterogeneity of the localization of all other thylakoid multiprotein complexes. In addition to its vital role in vectorial electron transfer and proton translocation across the membrane, cytochrome b
6
f complex is also involved in the regulation of balanced light excitation energy distribution between the photosystems, since its redox state governs the activation of LHC II kinase (the kinase that phosphorylates the mobile peripheral fraction of the chlorophyll a/b-proteins of LHC II of PS II). Hence, cytochrome b
6
f complex is the molecular link in the interactive co-regulation of light-harvesting and electron transfer.The importance of a highly dynamic, yet flexible organization of the thylakoid membranes of plants and green algae has been highlighted by the exciting discovery that a lateral reorganization of some cytochrome b
6
f complexes occurs in the state transition mechanism both in vivo and in vitro (Vallon et al. 1991). The lateral redistribution of phosphorylated LHC II from stacked granal membrane regions is accompanied by a concomitant movement of some cytochrome b
6
f complexes from the granal membranes out to the PS I-containing stroma thylakoids. Thus, the dynamic movement of cytochrome b
6
f complex as a multiprotein complex is a molecular mechanism for short-term adaptation to changing light conditions. With the concept of different membrane domains for linear and cyclic electron flow gaining credence, it is thought that linear electron flow occurs in the granal compartments and cyclic electron flow is localised in the stroma thylakoids at non-limiting irradiances. It is postulated that dynamic lateral reversible redistribution of some cytochrome b
6
f complexes are part of the molecular mechanism involved in the regulation of linear electron transfer (ATP and NADPH) and cyclic electron flow (ATP only). Finally, the molecular significance of the marked regulation of cytochrome b
6
f complexes for long-term regulation and optimization of photosynthetic function under varying environmental conditions, particularly light acclimation, is discussed.Abbreviations Chl
chlorophyll
- cyt
cytochrome
- PS
Photosystem 相似文献
2.
3.
Jean Alric 《Photosynthesis research》2010,106(1-2):47-56
Cyclic electron flow around PSI, or cyclic photophosphorylation, is the photosynthetic process which recycles the reducing equivalents produced by photosystem I in the stroma towards the plastoquinone pool. Through the activity of cytochrome b 6 f, which also transfers protons across the membrane, it promotes the synthesis of ATP. The literature dealing with cyclic electron flow in unicellular algae is far less abundant than it is for plants. However, in the chloroplast of algae such as Chlorella or Chlamydomonas, an efficient carbohydrate catabolism renders the redox poise much more reducing than in plant chloroplasts. It is therefore worthwhile highlighting the specific properties of unicellular algae because cyclic electron flow is highly dependent upon the accumulation of these stromal reducing equivalents. Such an increase of reducing power in the stroma stimulates the reduction of plastoquinones, which is the limiting step of cyclic electron flow. In anaerobic conditions in the dark, this reaction can lead to a fully reduced plastoquinone pool and induce state transitions, the migration of 80% of light harvesting complexes II and 20% of cytochrome b 6 f complex from the PSII-enriched grana to the PSI-enriched lamella. These ultrastructural changes have been proposed to further enhance cyclic electron flow by increasing PSI antenna size, and forming PSI-cyt b 6 f supercomplexes. These hypotheses are discussed in light of recently published data. 相似文献
4.
The organization of the electron transport components in mesophyll and bundle sheath chloroplasts of Zea mays was investigated. Grana-containing mesophyll chloroplasts (chlorophyll a to chlorophyll b ratio of about 3.0) possessed the full complement of the various electron transport components, comparable to chloroplasts from C3 plants. Agranal bundle sheath chloroplasts () contained the full complement of photosystem (PS) I and of cytochrome (cyt) f but lacked a major portion of PS II and its associated Chl light-harvesting complex (LHC), and most of the cyt b559. The kinetic analysis of system I photoactivity revealed that the functional photosynthetic unit size of PS I was unchanged and identical in mesophyll and bundle sheath chloroplasts. The results suggest that PS I is contained in stroma-exposed thylakoids and that it does not receive excitation energy from the Chl LHC present in the grana. A stoichiometric parity between PS I and cyt f in mesophyll and bundle sheath chloroplasts indicates that biosynthetic and functional properties of cyt f and P700 are closely coordinated. Thus, it is likely that both cyt f and P700 are located in the membrane of the intergrana thylakoids only. The kinetic analysis of PS II photoactivity revealed the absence of PS IIαfrom the bundle sheath chloroplasts and helped identify the small complement of system II in bundle sheath chloroplasts as PS IIβ. The distribution of the main electron transport components in grana and stroma thylakoids is presented in a model of the higher plant chloroplast membrane system. 相似文献
5.
Flash excitation of isolated intact chloroplasts promoted absorbance transients corresponding to the electrochromic effect (P-518) and the α-bands of cytochrome b6 and cytochrome f. Under conditions supporting coupled cyclic electron flow, the oxidation of cytochrome b6 and the reduction of cytochrome f had relaxation half-times of 15 and 17 ms, respectively. Optimal poising of cyclic electron flow, achieved by addition of 0.1 μM 3-(3,4-dichlorophenyl)-1,1-dimethylurea, increased phosphorylation of endogenous ADP and prolonged these relaxation times. The presence of NH4Cl, or monensin plus NaCl, decreased the half-times for cytochrome relaxation to approximately 2 ms. Uncouplers also revealed the presence of a slow rise component in the electrochromic absorption shift, with formation half-time of about 2 ms. The inhibitors of cyclic phosphorylation antimycin and 2,5-dibromo-3-methyl-6-isopropyl-p-benzoquinone abolished the slow rise in the electrochromic shift and prolonged the uncoupled relaxation times of cytochromes b6 and f by factors of ten or more.These observations indicate that cytochrome b6, plastoquinone and cytochrome f participate in a coupled electron transport process responsible for cyclic phosphorylation in intact chloroplasts. Estimations of cyclic phosphorylation rates from 40 to 120 μmol ATP/mg chlorophyll per h suggest that this process can provide a substantial fraction of the ATP needed for CO2 fixation. 相似文献
6.
Matthew P. Johnson Cvetelin Vasilev John D. Olsen C. Neil Hunter 《The Plant cell》2014,26(7):3051-3061
The cytochrome b6f (cytb6f) complex plays a central role in photosynthesis, coupling electron transport between photosystem II (PSII) and photosystem I to the generation of a transmembrane proton gradient used for the biosynthesis of ATP. Photosynthesis relies on rapid shuttling of electrons by plastoquinone (PQ) molecules between PSII and cytb6f complexes in the lipid phase of the thylakoid membrane. Thus, the relative membrane location of these complexes is crucial, yet remains unknown. Here, we exploit the selective binding of the electron transfer protein plastocyanin (Pc) to the lumenal membrane surface of the cytb6f complex using a Pc-functionalized atomic force microscope (AFM) probe to identify the position of cytb6f complexes in grana thylakoid membranes from spinach (Spinacia oleracea). This affinity-mapping AFM method directly correlates membrane surface topography with Pc-cytb6f interactions, allowing us to construct a map of the grana thylakoid membrane that reveals nanodomains of colocalized PSII and cytb6f complexes. We suggest that the close proximity between PSII and cytb6f complexes integrates solar energy conversion and electron transfer by fostering short-range diffusion of PQ in the protein-crowded thylakoid membrane, thereby optimizing photosynthetic efficiency. 相似文献
7.
Denis Saint-Marcoux Francis-André Wollman Catherine de Vitry 《The Journal of cell biology》2009,185(7):1195-1207
In chloroplasts, binding of a c′-heme to cytochrome b6 on the stromal side of the thylakoid membranes requires a specific mechanism distinct from the one at work for c-heme binding to cytochromes f and c6 on the lumenal side of membranes. Here, we show that the major protein components of this pathway, the CCBs, are bona fide transmembrane proteins. We demonstrate their association in a series of hetero-oligomeric complexes, some of which interact transiently with cytochrome b6 in the process of heme delivery to the apoprotein. In addition, we provide preliminary evidence for functional assembly of cytochrome b6f complexes even in the absence of c′-heme binding to cytochrome b6. Finally, we present a sequential model for apo- to holo-cytochrome b6 maturation integrated within the assembly pathway of b6f complexes in the thylakoid membranes. 相似文献
8.
Cytochrome b6 from spinach chloroplasts (either within the purified cytochrome b6f complex, or in its isolated form) exhibits two spectral species, which correspond to two midpoint potentials. This can be demonstrated by low temperature difference spectroscopy at fixed redox potentials. The high potential form of cytochrome b6 has a split α-peak at 557.5 and 561.5 nm, the low potential form has a symmetrical α-peak at 560.5 nm. Similar results were obtained with cytochrome b6 in the isolated cytochrome b6f complex from the cyanobacterium Anabaena variabilis. 相似文献
9.
《BBA》1986,851(2):229-238
We have analyzed the heme-associated peroxidase activity in thylakoid membranes from the green algae Chlamydomonas reinhardtii after electrophoresis in the presence of sodium dodecyl sulfate. Besides cytochrome f and cytochrome b6, we observed peroxidase activity in two other bands, of 34 and 11 kDa, of unknown origin. Characterization of the b6/f complex subunits was undertaken by means of a comparison of the polypeptide deficiencies in several b6/f mutants with the polypeptide content of preparations enriched in b6/f complexes. We conclude that the b6/f complex consists of five subunits. Using site-specific translation inhibitors, we show that cytochrome f, cytochrome b6 and subunit IV are of chloroplast origin, whereas the Rieske protein and probably subunit V are translated on cytoplasmic ribosomes. A model of assembly of the complex is proposed: a cytochrome moiety, comprising the subunits of chloroplast origin, is assembled in the thylakoid membranes prior to the insertion and assembly of the subunits encoded in the nuclear genome. 相似文献
10.
A.L. Ducluzeau 《BBA》2008,1777(9):1140-1146
Heliobacteria have a Rieske/cytochrome b complex composed of a Rieske protein, a cytochrome b6, a subunit IV and a di-heme cytochrome c. The overall structure of the complex seems close to the b6f complex from cyanobacteria and chloroplasts to the exception of the di-heme cytochrome. We show here by biochemical and biophysical studies that a heme ci is covalently attached to the Rieske/cytochrome b complex from Heliobacteria. We studied the EPR signature of this heme in two different species, Heliobacterium modesticaldum and Heliobacillus mobilis. In contrast to the case of b6f complex, a strong axial ligand to the heme is present, most probably a protonatable amino acid residue. 相似文献
11.
Magda Grabsztunowicz Paula Mulo Frauke Baymann Risa Mutoh Genji Kurisu Pierre Stif Peter Beyer Anja Krieger‐Liszkay 《The Plant journal : for cell and molecular biology》2019,99(2):245-256
During daffodil flower development, chloroplasts differentiate into photosynthetically inactive chromoplasts having lost functional photosynthetic reaction centers. Chromoplasts exhibit a respiratory activity reducing oxygen to water and generating ATP. Immunoblots revealed the presence of the plastid terminal oxidase (PTOX), the NAD(P)H dehydrogenase (NDH) complex, the cytochrome b6f complex, ATP synthase and several isoforms of ferredoxin‐NADP+ oxidoreductase (FNR), and ferredoxin (Fd). Fluorescence spectroscopy allowed the detection of chlorophyll a in the cytochrome b6f complex. Here we characterize the electron transport pathway of chromorespiration by using specific inhibitors for the NDH complex, the cytochrome b6f complex, FNR and redox‐inactive Fd in which the iron was replaced by gallium. Our data suggest an electron flow via two separate pathways, both reducing plastoquinone (PQ) and using PTOX as oxidase. The first oxidizes NADPH via FNR, Fd and cytochrome bh of the cytochrome b6f complex, and does not result in the pumping of protons across the membrane. In the second, electron transport takes place via the NDH complex using both NADH and NADPH as electron donor. FNR and Fd are not involved in this pathway. The NDH complex is responsible for the generation of the proton gradient. We propose a model for chromorespiration that may also be relevant for the understanding of chlororespiration and for the characterization of the electron input from Fd to the cytochrome b6f complex during cyclic electron transport in chloroplasts. 相似文献
12.
13.
The cytochrome bf complex was isolated from spinach thylakoids,and also from separated grana and stroma lamellae vesicles,by a procedure involving NaBr washing, detergent treatment andcentrifugation in sucrose gradients. The resulting complex fromall three types of membranes, were almost completely devoidof chlorophyll and carotenoids. The complexes have kinase activitytowards histone III-S and contain a 64 kDa protein claimed tobe a kinase. Electrophoretic analyses indicate that the complexesare in dimeric form and composed of six polypeptides with molecularmasses of 34/33, 23, 20, 17, 12 and 4 kDa. The complexes containtwo moles cytochrome b6 per mole cytochrome f and one mole RieskeFeS. The 17 kDa and 4 kDa polypeptides are the so called subunit4 and 5 respectively. The 12 kDa protein was identified as plastocyaninby immunoblotting. Plastocyanin and the 4 kDa protein were presentin the cytochrome bf complex even after a second repeated sucrosedensity gradient centrifugation. The sucrose gradient sedimentation pattern was different forthe grana and stroma lamellae complexes. The complex from thestroma lamellae arrives at a higher density than the grana complex.Furthermore, the gradient centrifugation diagram of the stromalamellae consists of one main peak while the diagram of thegrana complex shows two peaks. There is significantly more plastocyaninand 4 kDa protein in the bf complex isolated from stroma lamellaethan from grana. In addition there is a 15 kDa protein in thecomplex isolated from the grana vesicles. Immunoblot analysisafter crosslinking indicated that the 4 kDa protein and theplastocyanin are associated in the cytochrome bf complex. Theoxidoreductase activity is higher (about 50%) in the cytochromebf complex from the grana than from the stroma lamellae fraction.We suggest that a difference in composition of the cytochromebf complex between the two membranes might be important in theregulation of cyclic and non cyclic electron flow.
1Present address: Department of Plant Physiology II, Universityof Warsaw, 00 927 Warsaw, Poland 相似文献
14.
Cytochemical and immunocytochemical methods were used to localize photosystems I and II in barley (Hordeum vulgare L. cv Himalaya) chloroplasts. PSI activity, monitored by diaminobenzidine oxidation, was associated with the lumen side of the thylakoids of both grana and stroma lamellae. The P700 chlorophyll a protein, the reaction center of PSI, was localized on thin sections of barley chloroplasts using monospecific antibodies to this protein and the peroxidase-antiperoxidase procedure. Results obtained by immunocytochemistry were similar to those of the diaminobenzidine oxidation: both grana and stroma lamellae contained immunocytochemically reactive material. Both the grana and stroma lamellae were also labeled when isolated thylakoids were reacted with the P700 chlorophyll a protein antiserum and then processed by the peroxidase-antiperoxidase procedure. PSII activity was localized cytochemically by monitoring the photoreduction of thiocarbamyl nitroblue tetrazolium, a reaction sensitive to the PSII inhibitor, DCMU. PSII reactions occurred primarily on the grana lamellae, with weaker reactions on the stroma lamellae. 相似文献
15.
Dibromo- and diiodo-naphthoquinones are shown to be inhibitors of the cytochrome b6/f complex in isolated thylakoid membranes from spinach chloroplasts. Dibromo-naphthoquinone inhibits ferredoxin catalyzed cyclic photophosphorylation at 0.1 μM concentrations, but non cyclic e-flow only at 10 μM. It does not inhibit cyclic systems with artifical cofactors, nor non-cyclic electron flow from duroquinol through photosystem I via the cytochrome b6/f complex. Dibromo-naphthoquinone does however, lower the stoichiometry for ATP formation in the duroquinol donor system. This inhibitory pattern is quite different from that of DBMIB, but very similar to that of antimycin. This antimycin-like behaviour of these inhibitors is interpreted to indicate a) the existence of a Qc site in the cytochrome b6/f complex and its obligate function in ferredoxin catalyzed cyclic electron flow and b) a non-essential role of the Qc site in non-cyclic electron flow, but which — when operative — pumps an extra proton across the thylakoid membrane increasing the ATP yield. 相似文献
16.
Characterization of Triazine-Resistant and -Susceptible Isolines of Canola (Brassica napus L.) 总被引:4,自引:2,他引:2
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Vaughn KC 《Plant physiology》1986,82(3):859-863
Morphometric, electrophoretic, and immunological procedures were used to probe the structural and physiological differences between triazine-resistant (R) and susceptible (S) isolines of canola (Brassica napus L.). The R biotype exhibited increased grana stacking and decreased amounts of starch compared to the S biotype. Likewise, characters associated with an increase in grana stacking (lower chlorophyll a/b ratio, increased chlorophyll a/b light-harvesting complex, and relatively lower amounts of the P700 chlorophyll a protein and chloroplast coupling factor) were all observed in the R isoline of canola. Proteins which occur with approximately equal frequency in grana and stroma lamellae (plastocyanin, cutochrome f) or present only in the stroma (ribulose 1,5-bisphosphate carboxylase/oxygenase) were not quantitatively different in the two biotypes. Gross anatomical parameters (volume of epidermis, palisade mesophyll, spongy mesophyll, and air space) were similar in the two isolines. Thus, the triazine-resistance mutation does not confer a shade-type anatomy despite the chloroplast changes that are characteristic of shade biotypes or shade adaptions. These data indicate that the differences in chloroplast structure noted previously in comparisons of nonisonuclear R and S weed biotypes reflect differences in the triazineresistance factor rather than characters unrelated to triazine resistance. 相似文献
17.
《BBA》1987,892(3):314-319
Cytochrome b6 in freshly prepared, active cytochrome t6f complex from spinach chloroplasts shows a broad, low-spin EPR signal around gz = 3.6. Maximally half of the hemes of cytochrome b6 can be changed to high spin with a signal at g = 6 by inactivating treatments, or by isolating cytochrome b6. In this state the heme reacts with NO. Reduction rates suggest that it is the low-potential heme which changes. The change is accompanied by the loss of the shift in the gy signal of the Rieske FeS-center by quinone analogs. 相似文献
18.
Sallie G. Sprague 《Journal of bioenergetics and biomembranes》1987,19(6):691-703
Photosynthetic membranes of higher plant chloroplasts are composed primarily of polar, but uncharged, galactolipids unlike most mammalian membranes which contain large amounts of phosphatidylcholine. It is unclear what role(s) the galactolipids play in maintaining the differentiated thylakoid membranes, or in stabilizing the photosynthetically active enzyme complexes. Some of the membrane complexes show no lipid selectivity for maintaining structural or functional integrity. Others are poisoned or dissociated in the presence of high concentrations of a trace lipid class. The efficiency of energy transfer and the reconstitution of protein complexes into liposomes are dependent on the lipid class employed. The lipids are asymmetrically arranged along and across the thylakoid membranes but not as distinctly as the proteins.Abbreviations DGDG
digalactosyldiglyceride
- MGDG
monogalactosyldiglyceride
- SQDG
sulfoquinovosyldiglyceride
- PG
phosphatidylglycerol
- PC
phosphatidylcholine
- PE
phosphatidylethanolamine
- PSI
photosystem I
- PSII
photosystem II
- LHC
chlorophylla/b lightharvesting complex
- cytb
6
f
cytochromeb
6
f complex
- CF0/CF1
coupling factor ATPase
- DCIP
2,6-dichlorophenolindophenol
- LRa
galactolipase fromRhizopus arrhis 相似文献
19.
Cytochrome b-559 was purified from spinach leaves and antibodies were made against it in rabbit. Using affinity-purified, monospecific antibodies, we have found that cytochrome b-559, which is closely associated with the primary photochemical activity of photosystem II, is localized exclusively in the grana thylakoids. 相似文献
20.
Xian-De Liu 《BBA》2005,1706(3):215-219
This study investigated the regulation of the major light harvesting chlorophyll a/b protein (LHCII) phosphorylation in Dunaliella salina thylakoid membranes. We found that both light and NaCl could induce LHCII phosphorylation in D. salina thylakoid membranes. Treatments with oxidants (ferredoxin and NADP) or photosynthetic electron flow inhibitors (DCMU, DBMIB, and stigmatellin) inhibited LHCII phosphorylation induced by light but not that induced by NaCl. Furthermore, neither addition of CuCl2, an inhibitor of cytochrome b6f complex reduction, nor oxidizing treatment with ferricyanide inhibited light- or NaCl-induced LHCII phosphorylation, and both salts even induced LHCII phosphorylation in dark-adapted D. salina thylakoid membranes as other salts did. Together, these results indicate that the redox state of the cytochrome b6f complex is likely involved in light- but not salt-induced LHCII phosphorylation in D. salina thylakoid membranes. 相似文献