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1.
Variable chlorophyll a (Chl a) fluorescence is composed of a photochemical and a thermal phases of similar amplitudes. The photochemical phase can be induced
by a saturating single turnover flash (STF) and reflects the reduction of the Photosystem II (PS II) QA primary electron acceptor. The thermal phase requires multiple turnover flash (MTF) and is somehow related to the reduction
of the plastoquinone (PQ) molecules. This article aimed to determine the relative contributions of the QB-bound and the free oxidized PQ molecules to the thermal phase of Chl a fluorescence. We thus measured the interactive effects of exogenous PQ (PQex), of an inhibitor (DCMU) acting at the QB site of PS II and of an artificial quencher, 2-methyl-1,4-naphtoquinone, on Chl a fluorescence levels induced by STF (FF) and MTF (FM) in spinach thylakoids. We observed that: (1) the incorporation of PQex in thylakoids stimulated photosynthetic electron
transport but barely affected FF and FM in the absence of DCMU; (2) DCMU significantly increased the amplitude of FF but slightly quenched FM; (3) 2-methyl-1,4-naphtoquinone quenched FM to a larger-extent than FF; (4) DCMU increased the quenching effects of PQex on FF and FM and also, of methyl-1,4-naphtoquinone on FF. These results indicate that: (1) the QB-bound and the free PQ molecules contribute to about 56% and 25%, respectively, to the thermal phase Chl a fluorescence in dark-adapted thylakoids; and (2) the thermal phase of Chl a fluorescence is more susceptible than the photochemical phase to the non-photochemical quenching effect of oxidized quinones.
This revised version was published online in June 2006 with corrections to the Cover Date. 相似文献
2.
John F. Allen Conrad W. Mullineaux Christine E. Sanders Anastasios Melis 《Photosynthesis research》1989,22(2):157-166
Cells of the cyanobacterium Synechococcus 6301 were grown in yellow light absorbed primarily by the phycobilisome (PBS) light-harvesting antenna of photosystem II (PS II), and in red light absorbed primarily by chlorophyll and, therefore, by photosystem I (PS I). Chromatic acclimation of the cells produced a higher phycocyanin/chlorophyll ratio and higher PBS-PS II/PS I ratio in cells grown under PS I-light. State 1-state 2 transitions were demonstrated as changes in the yield of chlorophyll fluorescence in both cell types. The amplitude of state transitions was substantially lower in the PS II-light grown cells, suggesting a specific attenuation of fluorescence yield by a superimposed non-photochemical quenching of excitation. 77 K fluorescence emission spectra of each cell type in state 1 and in state 2 suggested that state transitions regulate excitation energy transfer from the phycobilisome antenna to the reaction centre of PS II and are distinct from photosystem stoichiometry adjustments. The kinetics of photosystem stoichiometry adjustment and the kinetics of the appearance of the non-photochemical quenching process were measured upon switching PS I-light grown cells to PS II-light, and vice versa. Photosystem stoichiometry adjustment was complete within about 48 h, while the non-photochemical quenching occurred within about 25 h. It is proposed that there are at least three distinct phenomena exerting specific effects on the rate of light absorption and light utilization by the two photoreactions: state transitions; photosystem stoichiometry adjustment; and non-photochemical excitation quenching. The relationship between these three distinct processes is discussed.Abbreviations Chl
chlorophyll
- DCMU
3-(3,4-dichlorophenyl)-1,1-dimethylurea
-
F
relative fluorescence intensity at emission wavelength nm
-
F
o
fluorescence intensity when all PS II traps are open
- light 1
light absorbed preferentially by PS I
- light 2
light absorbed preferentially by PS II
- PBS
phycobilisome
- PS
photosystem 相似文献
3.
《Plant Physiology and Biochemistry》2000,38(4):271-277
We have found that plastoquinone-A (PQ-A) and α-tocopherol (α-Toc) increased the reduction level of the high-potential form of cytochrome b-559 (cyt. b-559 HP) and α-tocopherol quinone (α-TQ) decreased the level of this cytochrome form in Scenedesmus obliquus wild-type, while the investigated prenyllipids were not active in the restoration of the cyt. b-559 HP form in Scenedesmus PS28 mutant and Synechococcus 6301 (Anacystis nidulans) where the cyt. b-559 HP form is naturally not present. Among the tested prenyllipids, α-TQ quenched fluorescence in thylakoids of S. obliquus wild-type, the PS28 mutant and tobacco to the highest extent, while PQ-A was less effective in this respect. α-Tocopherol showed the opposite effect to α-TQ and it was rather small. The fluorescence quenching measurements of thylakoids in the presence of DCMU (3-(3,4-dichlorophenyl)-1,1-dimethylurea) showed that the α-Toc and FCCP (carbonylcyanide-p-trifluoromethoxy-phenyl-hydrazone) did not quench non-photochemically chlorophyll fluorescence while PQ-9 and α-TQ were effective fluorescence quenchers at higher concentrations (> 15 μM). However, at the lower α-TQ concentrations where its effective fluorescence quenching was found in DCMU-free samples, there was nearly no quenching effect by α-TQ observed in DCMU-treated thylakoids. This suggested a specific, not non-photochemical, DCMU sensitive, fluorescence quenching of photosystem II (PSII) at low α-TQ concentrations which is probably connected with the cyclic electron transport around PSII and might have a function of excess light energy dissipation. The effects of α-TQ on PSII resembled those of FCCP under many respects which might suggest similar mechanism of action of these compounds on PSII, i.e. the catalytic deprotonation and/or redox changes of some components of PSII such as the water splitting system, tyrosine D, Chlz or cytochrome b-559. 相似文献
4.
The relaxation of the non-photochemical quenching of chlorophyll fluorescence has been investigated in cells of the green alga Dunaliella following illumination. The relaxation after the addition of DCMU or darkening was strongly biphasic. The uncoupler NH4Cl induced rapid relaxation of both phases, which were therefore both energy-dependent quenching, qE. The proportion of the slow phase of qE increased at increasing light intensity. In the presence of the inhibitors rotenone and antimycin the slow phase of qE was stabilised for in excess of 15 min. NaN3 inhibited the relaxation of almost all the qE. The implications of these results are discussed in terms of the interpretation of the non-photochemical quenching of chlorophyll fluorescence in vivo and the mechanism of qE.Abbreviations PS II
Photosystem II
- qQ
photochemical quenching of chlorophyll fluorescence
- qNP
non-photochemical quenching of chlorophyll fluorescence
- qE
energy-dependent quenching of chlorophyll fluorescence
-
F
m
maximum level of chlorophyll fluorescence for dark adapted cells
-
F
m
level of fluorescence at any time when qQ is zero 相似文献
5.
High energy state quenching of chlorophyll fluorescence (qE) is inhibited by low concentrations of the inhibitor antimycin A in intact and osmotically shocked chloroplasts isolated from spinach and pea plants. This inhibition is independent of any effect upon pH (as measured by 9-aminoacridine fluorescence quenching). A dual control of qE formation, by pH and the redox state of an unidentified chloroplast component, is implied. Results are discussed in terms of a role for qE in the dissipation of excess excitation energy within photosystem II.Abbreviations 9-AAmax =
Maximum yield of 9-aminoacridine fluorescence
- DCMU =
3(3,4-dichlorophenyl)-1,1-dimethylurea; Fmax ± Maximum yield of chlorophyll fluorescence
- hr =
hour
- PAR =
Photosynthetically Active Radiation
- QA =
Primary stable electron acceptor within photosystem II
- qE =
High energy state quenching of chlorophyll fluorescence
- qI =
quenching of chlorophyll fluorescence related to photoinhibition
- qP =
Quenching of chlorophyll fluorescence by oxidised plastoquinone
- qQ =
photochemical quenching of chlorophyll fluorescence
- qR =
(Fmax—maximum level of chlorophyll fluorescence induced by the addition of saturating DCMU)
- qT =
Quenching of chlorophyll fluorescence attributable to state transitions 相似文献
6.
The components of non-photochemical chlorophyll fluorescence quenching (qN) in barley leaves have been quantified by a combination of relaxation kinetics analysis and 77 K fluorescence measurements (Walters RG and Horton P 1991). Analysis of the behaviour of chlorophyll fluorescence parameters and oxygen evolution at low light (when only state transitions — measured as qNt — are present) and at high light (when only photoinhibition — measured as qNi — is increasing) showed that the parameter qNt represents quenching processes located in the antenna and that qNi measures quenching processes located in the reaction centre but which operate significantly only when those centres are closed. The theoretical predictions of a variety of models describing possible mechanisms for high-energy-state quenching, measured as the residual quenching, qNe, were then tested against the experimental data for both fluorescence quenching and quantum yield of oxygen evolution. Only one model was found to agree with these data, one in which antennae exist in two states, efficient in either energy transfer or energy dissipation, and in which those photosynthetic units in a dissipative state are unable to exchange energy with non-dissipative units.Abbreviations: Fo, Fm
room-temperature chlorophyll fluorescence yield with all centres open, closed
- Fv
variable fluorescence yield
- LHC II
light-harvesting chlorophyll-protein complex of PS II
- PS I, PS II
Photosystem I, II
- P700, P680
primary donor in Photosystem I, II
- QA
primary electron acceptor of PS II
- Pmax
maximum quantum yield of oxygen evolution
- qN
coefficient of non-photochemical quenching of variable fluorescence
- qNe, qNt, qNi
coefficient of non-photochemical quenching due to high-energy-state, state transition, photoinhibition
- qO
coefficient of quenching of dark level fluorescence
- qP
coefficient of photochemical quenching of variable fluorescence
- P
intrinsic quantum yield of open PS II reaction centres = s/qP
- PS 2
quantum yield of PS = qP × Fv/Fm
- S
quantum yield of oxygen evolution = rate of oxygen evolution/light intensity 相似文献
7.
Douglas Campbell Doug Bruce Christene Carpenter Petter Gustafsson Gunnar Öquist 《Photosynthesis research》1996,47(2):131-144
Synechococcus sp. PCC 7942 (Anacystis nidulans R2) contains two forms of the Photosystem II reaction centre protein D1, which differ in 25 of 360 amino acids. D1: 1 predominates under low light but is transiently replaced by D1:2 upon shifts to higher light. Mutant cells containing only D1:1 have lower photochemical energy capture efficiency and decreased resistance to photoinhibition, compared to cells containing D1:2. We show that when dark-adapted or under low to moderate light, cells with D1:1 have higher non-photochemical quenching of PS II fluorescence (higher qN) than do cells with D1:2. This is reflected in the 77 K chlorophyll emission spectra, with lower Photosystem II fluorescence at 697–698 nm in cells containing D1:1 than in cells with D1:2. This difference in quenching of Photosystem II fluorescence occurs upon excitation of both chlorophyll at 435 nm and phycobilisomes at 570 nm. Measurement of time-resolved room temperature fluorescence shows that Photosystem II fluorescence related to charge stabilization is quenched more rapidly in cells containing D1:1 than in those with D1:2. Cells containing D1:1 appear generally shifted towards State II, with PS II down-regulated, while cells with D1:2 tend towards State I. In these cyanobacteria electron transport away from PS II remains non-saturated even under photoinhibitory levels of light. Therefore, the higher activity of D1:2 Photosystem II centres may allow more rapid photochemical dissipation of excess energy into the electron transport chain. D1:1 confers capacity for extreme State II which may be of benefit under low and variable light.Abbreviations D1
the atrazine-binding 32 kDa protein of the PS II reaction centre core
- D1:1
the D1 protein constitutively expressed during acclimated growth in Synechococcus sp. PCC 7942
- D1:2
an alternate form of the D1 protein induced under excess excitation in Synechococcus sp. PCC 7942
- DCMU
3-(3,4-dichlorophenyl)-1,1-dimethyl urea
- Fo
minimal fluorescence in the dark-adapted state
- Fo
minimal fluorescence in a light-adapted state
- FM
maximum fluorescence with all quenching mechanisms at a minimum, measured in presence of DCMU
- FM
maximal fluorescence in a light-adapted state, measured with a saturating flash
- FMdark
maximal fluorescence in the dark-adapted state
- FV
variable fluorescence in a light-adapted state (FM-Fo)
- PAM
pulse amplitude modulated fluorometer
- qN
non-photochemical quenching of PS II fluorescence
- qN (dark)
qN in the dark adapted state
- qP
photochemical quenching of fluorescence 相似文献
8.
The effect of high-energy-state excitation quenching on maximum and dark level chlorophyll fluorescence yield 总被引:3,自引:0,他引:3
The quenching of variable fluorescence yield (qN) and the quenching of dark level fluorescence yield (q0) directly atributable to high-energy-state fluorescence quenching (qE) was studied to distinguish between energy dissipation in the antenna and light harvesting complexes (antenna quenching) and energy dissipation at the reaction centres (reaction centre quenching). A consistent relationship was obtained between qN and q0 in barley leaves, the green alga Dunaliella C9AA and in pea thylakoids with 2,3,5,6-tetramethyl-p-phenylene diamine (DAD) as mediator of cyclic electron flow around PS 1. This correlated well with the relationship obtained using m-dinitrobenzene (DNB), a chemical model for antenna quenching, to quench fluorescence in Dunaliella C9AA or pea thylakoids. The results also correlated reasonably well with theoretical predictions by the Butler model for antenna quenching, but did not correlate with the predictions for reaction centre quenching. It is postulated that qE quenching therefore occures in the antenna and light harvesting complexes, and that the small deviation from the Butler prediction is due to PS 2 heterogeneity.Abbreviations 9-aa
9-aminoacridine
- DCMU
3-(3,4-dichlorophenyl)-1,1-dimethylurea
- EDTA
Ethylenediaminetetra-acetic acid
- Hepes
4-(2-hydroxyethyl)-1-piperazineethanesulphonic acid
- Mes
2-(N-morpholino) prophanesulfonate
- PS 1
photosystem 1
- PS 2
photosystem 2
- QA and QB
primary and secondary stable electron acceptors of photosystem 2
- qN
non-photochemical fluorescence quenching coefficient
- qE
high-energy-state fluorescence quenching coefficient
- q0
quenching coefficient for F0
- F0
dark level fluorescence yield
- Fm
maximum fluorescence yield
- Fv
variable fluorescence yield
- Fv/Fm
ratio of variable to total fluorescence yield
- DAD
2,3,5,6-tetramethyl-p-phenylene diamine
- DNB
m-dinitrobenzene 相似文献
9.
Twenty-five years ago, non-photochemical quenching of chlorophyll fluorescence by oxidised plastoquinone (PQ) was proposed to be responsible for the lowering of the maximum fluorescence yield reported to occur when leaves or chloroplasts were treated in the dark with 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU), an inhibitor of electron flow beyond the primary quinone electron acceptor (QA) of photosystem (PS) II [C. Vernotte, A.L. Etienne, J.-M. Briantais, Quenching of the system II chlorophyll fluorescence by the plastoquinone pool, Biochim. Biophys. Acta 545 (1979) 519-527]. Since then, the notion of PQ-quenching has received support but has also been put in doubt, due to inconsistent experimental findings. In the present study, the possible role of the native PQ-pool as a non-photochemical quencher was reinvestigated, employing measurements of the fast chlorophyll a fluorescence kinetics (from 50 μs to 5 s). The about 20% lowering of the maximum fluorescence yield FM, observed in osmotically broken spinach chloroplasts treated with DCMU, was eliminated when the oxidised PQ-pool was non-photochemically reduced to PQH2 by dark incubation of the samples in the presence of NAD(P)H, both under anaerobic and aerobic conditions. Incubation under anaerobic conditions in the absence of NAD(P)H had comparatively minor effects. In DCMU-treated samples incubated in the presence of NAD(P)H fluorescence quenching started to develop again after 20-30 ms of illumination, i.e., the time when PQH2 starts getting reoxidised by PS I activity. NAD(P)H-dependent restoration of FM was largely, if not completely, eliminated when the samples were briefly (5 s) pre-illuminated with red or far-red light. Addition to the incubation medium of HgCl2 that inhibits dark reduction of PQ by NAD(P)H also abolished NAD(P)H-dependent restoration of FM. Collectively, our results provide strong new evidence for the occurrence of PQ-quenching. The finding that DCMU alone did not affect the minimum fluorescence yield F0 allowed us to calculate, for different redox states of the native PQ-pool, the fractional quenching at the F0 level (Q0) and to compare it with the fractional quenching at the FM level (QM). The experimentally determined Q0/QM ratios were found to be equal to the corresponding F0/FM ratios, demonstrating that PQ-quenching is solely exerted on the excited state of antenna chlorophylls. 相似文献
10.
Dissipation of absorbed excitation energy as heat, measured by its effect on the quenching of chlorophyll fluorescence, is induced under conditions of excess light in order to protect the photosynthetic apparatus of plants from light-dependent damage. The spectral characteristics of this quenching have been compared to that due to photochemistry in the Photosystem II reaction centre using leaves of Guzmania monostachia. This was achieved by making measurements at 77K when fluorescence emission bands from each type of chlorophyll protein complex can be distinguished. It was demonstrated that photochemistry and non-photochemical dissipation preferentially quench different emission bands and therefore occur by dissimilar mechanisms at separate sites. It was found that photochemistry was associated with a preferential quenching of emission at 688 nm whereas the spectrum for rapidly reversible non-photochemical quenching had maxima at 683 nm and 698 nm, suggesting selective quenching of the bands originating from the light harvesting complexes of Photosystem II. Further evidence that this was occurring in the light harvesting system was obtained from the fluorescence excitation spectra recorded in the quenched and relaxed states.Abbreviations pH
transthylakoid pH gradient
- Fo
minimum level of chlorophyll fluorescence when Photosystem II reaction centres are open
- Fm
maximum level of fluorescence when Photosystem II reaction centres are closed
- Fv
variable fluorescence Fm
minus Fo
- F'o
Fo in any quenched state
- Fm
Fm in any quenched state
- LHCII
light harvesting complexes of Photosystem II
- PSI
Photosystem I
- PS II
Photosystem II
- qN
non-photochemical quenching of chlorophyll fluorescence
- qE
non-photochemical quenching of chlorophyll fluorescence that occurs in the presence of a pH 相似文献
11.
The time courses of some Photosystem II (PS II) parameters have been monitored during in-vivo and in-vitro photoinhibition of spinach chloroplasts, at room temperature and at 10 °C or 0 °C. Exposing leaf discs of low-light grown spinach at 25 °C to high light led to photoinhibition of chloroplasts in-vivo as manifested by a parallel decrease in the number of functional PS II centres, the variable chlorophyll fluorescence at 77K (F
v
/F
m
), and the number of atrazine-binding sites. When the photoinhibitory treatment was given at 10 °C, the former two parameters declined in parallel but the loss of atrazine-binding sites occurred more slowly and to a lesser extent. During in-vitro photoinhibition of chloroplast thylakoids at 25 °C, the loss of functional PS II centres proceeded slightly more rapidly than the loss of atrazine-binding sites, and this difference in rate was further increased when the thylakoids were photoinhibited at 0 °C. During the recovery phase of leaf discs (up to 9 h) the increases in F
v
/F
m
preceded that of the number of functional PS II centres, while only a further decline in the number of atrazine-binding sites was observed. The recovery of variable chlorophyll fluorescence and the concentration of functional PS II centres occurred more rapidly at 25 °C than at 10 °C. These results suggest that the photoinhibition of PS II function is a relatively temperature-independent early photochemical event, whereas the changes in the concentration of herbicide-binding sites appear to be a more complex biochemical process which can occur with a delayed time course.Abbreviations BSA
bovine serum albumin
- Chl
chlorophyll
- D1
32kDa herbicide-binding polypeptide in photosystem II and product of the psbA gene
- D2
34kDa polypeptide in photosystem II which is the product of the psbD gene
- DCMU
3-(3,4-dichlorophenyl)-1,1-dimethylurea
- DCPIP
2,6-dichlorophenolin-dophenol
-
F
0, F
v
, F
m
chlorophyll fluorescence with reaction centres open, variable and maximum fluorescence, respectively
- LDS
lithium dodecyl sulfate
- MES
2-(N-morpholino) ethanesulfonic acid
- PSII
photosystem II
- QA, QB
first and second quinone-type PS II acceptor, respectively 相似文献
12.
Adam M. Gilmore Theodore L. Hazlett Peter G. Debrunner Govindjee 《Photosynthesis research》1996,48(1-2):171-187
Photosystem II (PS II) chlorophyll (Chl) a fluorescence lifetimes were measured in thylakoids and leaves of barley wild-type and chlorina f104 and f2 mutants to determine the effects of the PS II Chl a+b antenna size on the deexcitation of absorbed light energy. These barley chlorina mutants have drastically reduced levels of PS II light-harvesting Chls and pigment-proteins when compared to wild-type plants. However, the mutant and wild-type PS II Chl a fluorescence lifetimes and intensity parameters were remarkably similar and thus independent of the PS II light-harvesting antenna size for both maximal (at minimum Chl fluorescence level, Fo) and minimal rates of PS II photochemistry (at maximum Chl fluorescence level, Fm). Further, the fluorescence lifetimes and intensity parameters, as affected by the trans-thylakoid membrane pH gradient (pH) and the carotenoid pigments of the xanthophyll cycle, were also similar and independent of the antenna size differences. In the presence of a pH, the xanthophyll cycle-dependent processes increased the fractional intensity of a Chl a fluorescence lifetime distribution centered around 0.4–0.5 ns, at the expense of a 1.6 ns lifetime distribution (see Gilmore et al. (1995) Proc Natl Acad Sci USA 92: 2273–2277). When the zeaxanthin and antheraxanthin concentrations were measured relative to the number of PS II reaction center units, the ratios of fluorescence quenching to [xanthophyll] were similar between the wild-type and chlorina f104. However, the chlorina f104, compared to the wild-type, required around 2.5 times higher concentrations of these xanthophylls relative to Chl a+b to obtain the same levels of xanthophyll cycle-dependent fluorescence quenching. We thus suggest that, at a constant pH, the fraction of the short lifetime distribution is determined by the concentration and thus binding frequency of the xanthophylls in the PS II inner antenna. The pH also affected both the widths and centers of the lifetime distributions independent of the xanthophyll cycle. We suggest that the combined effects of the xanthophyll cycle and pH cause major conformational changes in the pigment-protein complexes of the PS II inner or core antennae that switch a normal PS II unit to an increased rate constant of heat dissipation. We discuss a model of the PS II photochemical apparatus where PS II photochemistry and xanthophyll cycle-dependent energy dissipation are independent of the Peripheral antenna size.Abbreviations Ax
antheraxanthin
- BSA
bovine serum albumin
- cx
lifetime center of fluorescence decay component x
- CP
chlorophyll binding protein of PS II inner antenna
- DCMU
3-(3,4-dichlorophenyl)-1,1-dimethylurea
- DTT
dithiothreitol
- fx
fractional intensity of fluorescence lifetime component x
- Fm, Fm
maximal PS II Chl a fluorescence intensity with all QA reduced in the absence, presence of thylakoid membrane energization
- Fo
minimal PS II Chl a fluorescence intensity with all QA oxidized
- Fv=Fm–Fo
variable level of PS II Chl a fluorescence
- HPLC
high performance liquid chromatography
- kA
rate constant of all combined energy dissipation pathways in PS II except photochemistry and fluorescence
- kF
rate constant of PS II Chl a fluorescence
- LHCIIb
main light harvesting pigment-protein complex (of PS II)
- Npig
mols Chl a+b per PS II
- NPQ=(Fm/Fm–1)
nonphotochemical quenching of PS II Chl a fluorescence
- PAM
pulse-amplitude modulation fluorometer
- PFD
photon-flux density, mols photons m–2 s–1
- PS II
Photosystem II
- P680
special-pair Chls of PS II reaction center
- QA
primary quinone electron acceptor of PS II
- Vx
violaxanthin
- wx
width at half maximum of Lorentzian fluorescence lifetime distribution x
- Zx
zeaxanthin
- pH
trans-thylakoid proton gradient
- % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXafv3ySLgzGmvETj2BSbqef0uAJj3BZ9Mz0bYu% H52CGmvzYLMzaerbd9wDYLwzYbItLDharqqr1ngBPrgifHhDYfgasa% acOqpw0xe9v8qqaqFD0xXdHaVhbbf9v8qqaqFr0xc9pk0xbba9q8Wq% Ffea0-yr0RYxir-Jbba9q8aq0-yq-He9q8qqQ8frFve9Fve9Ff0dme% GabaqaaiGacaGaamqadaabaeaafiaakeaacqGH8aapcqaHepaDcqGH% +aGpdaWgaaWcbaGaamOraiaad2gaaeqaaaaa!4989!\[< \tau > _{Fm}\],% MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXafv3ySLgzGmvETj2BSbqef0uAJj3BZ9Mz0bYu% H52CGmvzYLMzaerbd9wDYLwzYbItLDharqqr1ngBPrgifHhDYfgasa% acOqpw0xe9v8qqaqFD0xXdHaVhbbf9v8qqaqFr0xc9pk0xbba9q8Wq% Ffea0-yr0RYxir-Jbba9q8aq0-yq-He9q8qqQ8frFve9Fve9Ff0dme% GabaqaaiGacaGaamqadaabaeaafiaakeaacqGH8aapcqaHepaDcqGH% +aGpdaWgaaWcbaGaamOraiaad+gaaeqaaOGaeyypa0Zaaabqaeaaca% WGMbWaaSbaaSqaaiaadIhaaeqaaOGaam4yamaaBaaaleaacaWG4baa% beaaaeqabeqdcqGHris5aaaa!50D3!\[< \tau > _{Fo} = \sum {f_x c_x }\]
average lifetime of Chl a fluorescence calculated from a multi-exponential model under Fm, Fo conditions 相似文献
13.
The hypothesis that the chlorophyll fluorescence decline due to membrane phosphorylation is caused principally by the detachment and removal of LHCP from the LHCP-PS II matrix is examined. It is demonstrated that when membranes are phosphorylated in the dark (a) the fluorescence decline is greater when excited by light enriched in wavelengths absorbed mainly by LHCP (475 nm) than when excited by light absorbed to a large extent also by the PS II complex (435 nm), (b) titration with different artificial quenchers of chlorophyll fluorescence is unchanged after the phosphorylation-induced fluorescence decline, and (c) the Fv/Fm ratio does not change after the phosphorylation-induced fluorescence decline. These data indicate that it is indeed principally LHCP that interacts with the quencher (PS I presumably). This interaction involves a small fraction of the total PS II-coupled LHCP, which becomes functionally detached from the LHCP-PS II matrix. 相似文献
14.
A model is presented describing the relationship between chlorophyll fluorescence quenching and photoinhibition of Photosystem (PS) II-dependent electron transport in chloroplasts. The model is based on the hypothesis that excess light creates a population of inhibited PS II units in the thylakoids. Those units are supposed to posses photochemically inactive reaction centers which convert excitation energy to heat and thereby quench variable fluorescence. If predominant photoinhibition of PS II and cooperativity in energy transfer between inhibited and active units are presumed, a quasi-linear correlation between PS II activity and the ratio of variable to maximum fluorescence, FVFM, is obtained. However, the simulation does not result in an inherent linearity of the relationship between quantum yield of PS II and FVFM ratio. The model is used to fit experimental data on photoinhibited isolated chloroplasts. Results are discussed in view of current hypotheses of photoinhibition.Abbreviations FM
maximum total fluorescence
- F0
initial fluorescence
- FV
maximum variable fluorescence
- PS
Photosystem
- QA, QB
primary and secondary electron acceptors of Photosystem II 相似文献
15.
Photosynthetic electron transfer was studied in thylakoids isolated from control and DCMU-grown wheat (Triticum aestivum L.) seedlings. When exposed to high temperature (HT) and high iradiance (HI), thylakoids showed large variations in the photosynthetic electron transport activities and thylakoid membrane proteins. A drastic reduction in the rate of whole electron transport chain (H2O MV) was envisaged in control thylakoids when exposed to HT and HI. Such reduction was mainly due to the loss of photosystem 2, PS2 (H2O DCBQ) activity. The thylakoids isolated from seedlings grown in the presence of DCMU showed greater resistance to HT and HI treatment. The artificial exogenous electron donors MnCl2, DPC, and NH2OH failed to restore the HI induced loss of PS2 activity in both control and DCMU thylakoids. In contrast, addition of DPC and NH2OH significantly restored the HT induced loss of PS2 activity in control thylakoids and partially in DCMU thylakoids. Similar results were obtained when Fv/Fm was evaluated by chlorophyll fluorescence measurements. The marked loss of PS2 activity in control thylakoids was evidently due to the loss of 33, 23, and 17 kDa extrinsic polypeptides and 28-25 kDa LHCP polypeptides. 相似文献
16.
Effect of preheating of beet spinach leaves on chlorophyll a fluorescence yield was analyzed with the help of additional high intensity illumination pulses using a pulse modulated fluorometer. Preheating at mildly elevated temperature (35–45°C) causes a shift in the redox state of secondary donor of photosystem II, possibly due to uncoupling of phosphorylation because of thermal induced membrane disorganization and associated alkalinization of intra thylakoid space. Also, at these preheating temperatures, a rise in photosystem I catalyzed electron transfer has been shown to occur. These two effects induce rapid quenching of Chi a fluorescence, which drops even in the presence of actinic light, below the level of initial fluorescence (Fo′ monitored by the weak modulated probing light. Preheating of leaf segments induces an increase in fluorescence in the presence of dluron, which blocks electron flow between two photosystems, and thus this increases in fluorescence yield (Fo′ as monitored by weak modulated light, is not solely due to disorganization of light harvesting Chi-protein complex but also due to a shift in the redox equilibrium of the donor at the oxidizing side of photosystem II resulting in rapid reduction of QA the stable primary acceptor of photosystem II. In 50°C preheated DCMU treated samples, the fluorescence yield increases in weak modulated light and it approaches that of maximal steady state (Fmax) level. At preheating temperature of 48°–50°C, the inactivation of enzymes in the reducing side of photosystem I, causes an impairment of the reoxidation of QA and under this condition, a strong illumination causes quenching of Chi a fluorescence. This quenching seems to arise because of accumulation of the P680+, the oxidized physiological donor of photosystem which is a quencher of Chi a fluorescence. This quenching depended on the pulse intensity and duration which saturates P680+ accumulation and is greatly manifested when water oxidation complex is damaged. 相似文献
17.
Plants respond to excess light by a photoprotective reduction of the light harvesting efficiency. The notion that the non-photochemical quenching of chlorophyll fluorescence can be reliably used as an indicator of the photoprotection is put to a test here. The technique of the repetitive flash fluorescence induction is employed to measure in parallel the non-photochemical quenching of the maximum fluorescence and the functional cross-section (sigma(PS II)) which is a product of the photosystem II optical cross-section a(PS II) and of its photochemical yield Phi(PS II) (sigma (PS II) = a(PS II) Phi(PS II)). The quenching is measured for both, the maximum fluorescence found in a single-turnover flash (F(M) (ST)) and in a multiple turnover light pulse (F(M) (MT)). The experiment with the diatom Phaeodactylum tricornutum confirmed that, in line with the prevalent model, the PS II functional cross-section sigma (PS II) is reduced in high light and restored in the dark with kinetics and amplitude that are closely matching the changes of the F(M) (ST) and F(M) (MT) quenching. In contrast, a poor correlation between the light-induced changes in the PS II functional cross-section sigma (PS II) and the quenching of the multiple-turnover F(M) (MT) fluorescence was found in the green alga Scenedesmus quadricauda. The non-photochemical quenching in Scenedesmus quadricauda was further investigated using series of single-turnover flashes given with different frequencies. Several mechanisms that modulate the fluorescence emission in parallel to the Q(A) redox state and to the membrane energization were resolved and classified in relation to the light harvesting capacity of Photosystem II. 相似文献
18.
We have measured thermoluminescence (TL) and chlorophyll fluorescence from leaves of peas grown under an intermittent light regime (IML) and followed changes in those leaves during greening. IML peas show low variable fluorescence and a certain capacity for reversible non-photochemical quenching. It has been suggested that reversible quenching may be caused by pH-dependent release of Ca2+ from Photosystem II (PS II) (Krieger and Weis (1992) Photosynthetica 27: 89–98). Under conditions in which reversible non-photochemical quenching occurs, a TL band at around 50 °C is observed, in the presence of DCMU, in IML leaves. A band in this temperature range has previously been observed in PS II depleted of Ca2+ (Ono and Inoue (1989) Biochimica et Biophysica Acta 973: 443–449). The 50 °C band disappears upon dark adaptation. In mature leaves, no significant band is seen at 50 °C. It is concluded that, in IML leaves, reversible quenching may be related to the release of Ca2+ from Photosystem II. However, it seems that in the mature system, under most conditions, such release does not contribute significantly to quenching 相似文献
19.
《BBA》1986,848(2):239-246
Analyses of room-temperature chlorophyll fluorescence curves from DCMU-inhibited thylakoids were used to investigate the proposed PS II structural heterogeneity of α and β centres. The kinetics of the area growth curves, representative of QA photoreduction, could be modified in the presence of DCMU by exogenous electron acceptors and by added reductants of the PQ pool. The effect of altered DCMU levels (range 0.2–100 μM) on the induction curve kinetics was to modify preferentially the slow-β component, while having only a very small effect on the total variable fluorescence yield. Over the DCMU concentration range used, the unnormalized area of the induction curve (Amax) decreased with increasing herbicide concentration by approx. 45%, indicating that less quanta were required to reduce QA. It was found that the dark reoxidation of QA in the presence of DCMU and Ant 2p after a light pretreatment regenerated the slow kinetic component. When chlorophyll fluorescence emission at 685 and 731 nm was measured, no difference was observed in the kinetics of the induction curve. The analysis of PS II-enriched, oxygen-evolving membranes indicated the presence of both the fast and slow kinetic components, although this type of preparation showed a modified fast phase. The above observations led to the conclusion that several of the previously proposed characteristics of PS IIα and PS IIβ centres do not hold and that a type of PS II heterogeneity involving different degrees of DCMU inhibition is sufficient to explain many of the observations made. 相似文献
20.
Thylakoids isolated from peas (Pisum sativum cv. Kelvedon Wonder) and phosphorylated by incubation with ATP have been compared with non-phosphorylated thylakoids in their sensitivity to photoinhibition by exposure to illumination in vitro. Assays of the kinetics of fluorescence induction at 20° C and the fluorescence emission spectra at-196° C indicate a proportionally larger decrease in fluorescence as a result of photoinhibitory treatment of non-phosphorylated compared with phosphorylated thylakoids. It is concluded that protein phosphorylation can afford partial protection to thylakoids exposed to photoinhibitory conditions.Abbreviations and symbols DCMU
3-(3,4-dichlorophenyl)-1,1-dimethylurea
-
F
0
Level of chlorophyll fluorescence when photosystem 2 traps are open
-
F
m
Level of chlorphyll fluorescence when photosystem 2 traps are closed
- P
Maximum level of fluorescence reached in the absence of DCMU
- PSI (II)
photosystem I(II) 相似文献