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1.
2.
P700 is rapidly, but only transiently photooxidized upon illuminating dark-adapted leaves. Initial oxidation is followed by a reductive phase even under far-red illumination which excites predominantly photosystem (PS) I. In this phase, oxidized P700 is reduced by electrons coming from PSII. Charge separation in the reaction center of PSI is prevented by the unavailability of electron acceptors on the reducing side of PSI. It is subsequently made possible by the opening of an electron gate which is situated between PSI and the electron acceptor phosphoglycerate. Electron acceptors immediately available for reduction while the gate is closed corresponded to 10 nmol · (mg chlorophyll)–1 electrons in geranium leaves, 16 nmol · (mg chlorophyll)–1 in sunflower and 22 nmol · (mg chlorophyll)–1 in oleander. Reduction of NADP during the initial phase of P700 oxidation showed that the electron gate was not represented by ferredoxin-NADP reductase. Availability of ATP indicated that electron flow was not hindered by deactivation of the thylakoid ATP synthetase. It is concluded that NADP-dependent glyceraldehydephosphate dehydrogenase is completely deactivated in the dark and activated in the light. The rate of activation depends on the length of the preceding dark period. As chloroplasts contain both NAD- and NADP-dependent glyceraldehydephosphate dehydrogenases, deactivation of the NADP-dependent enzyme disconnects chloroplast NAD and NADP systems and prevents phosphoglycerate reduction in the dark at the expense of NADPH and ATP which are generated by glucose-6-phosphate oxidation and glycolytic starch breakdown, respectively.Abbreviations Chl
chlorophyll
- P700
electron donor pigment in the reaction center of photosystem I
Cooperation of the Institute of Botany of the University of Würzburg with the Institute of Astrophysics and Atmospheric Physics of the Estonian Academy of Sciences in Tartu was supported by the Deutsche Forschungsgemeinschaft and the Estonian Academy of Sciences. This work was performed within the Sonderforschungsbereich 251 of the University of Würzburg. 相似文献
3.
Oscillations in photosynthesis are initiated and supported by imbalances in the supply of ATP and NADPH to the Calvin cycle 总被引:4,自引:0,他引:4
Agu Laisk Katharina Siebke Ulvi Gerst Hillar Eichelmann Vello Oja Ulrich Heber 《Planta》1991,185(4):554-562
Oscillations in the rate of photosynthesis of sunflower (Helianthus annuus L.) leaves were induced by subjecting leaves, whose photosynthetic apparatus had been activated, to a sudden transition from darkness or low light to high-intensity illumination, or by transfering them in the light from air to an atmosphere containing saturating CO2. It was found that at the first maximum, light-and CO2-saturated photosynthesis can be much faster than steady-state photosynthesis. Both QA in the reaction center of PS II and P700 in the reaction center of PS I of the chloroplast electron-transport chain were more oxidized during the maxima of photosynthesis than during the minima. Maxima of P700 oxidation slightly preceded maxima in photosynthesis. During a transition from low to high irradiance, the assimilatory force FA, which was calculated from ratios of dihydroxyacetone phosphate to phosphoglycerate under the assumption that the reactions catalyzed by NADP-dependent glyceraldehydephosphate dehydrogenase, phosphoglycerate kinase and triosephosphate isomerase are close to equilibrium, oscillated in parallel with photosynthesis. However, only one of its components, the calculated phosphorylation potential (ATP)/(ADP)(Pi), paralleled photosynthesis, whereas calculated NADPH/NADP ratios exhibited antiparallel behaviour. When photosynthetic oscillations were initiated by a transition from low to high CO2, the assimilatory force FA declined, was very low at the first minimum of photosynthesis and increased as photosynthesis rose to its second maximum. The observations indicate that the minima in photosynthesis are caused by lack of ATP. This leads to overreduction of the electron-transport chain which is indicated by the reduction of P700. During photosynthetic oscillations the chloroplast thylakoid system is unable to adjust the supply of ATP and NADPH rapidly to demand at the stoichiometric relationship required by the carbonreduction cycle.Abbreviations PGA
3-phosphoglycerate
- DHAP
dihydroxyacetone phosphate
- P700
electron-donor pigment in the reaction enter of PS I
- QA
quinone acceptor in the reaction center of PS II
This work received support from the Estonian Academy of Sciences, the Bavarian Ministry of Science and Art and the Sonderforschungsbereich 251 of the University of Würzburg. We are grateful for criticism by D.A. Walker, Robert Hill Institute, University of Sheffield, U.K. and by Mark Stitt, Institute of Botany, University of Heidelberg, FRG. 相似文献
4.
Effects of freezing on biological membranes in vivo and in vitro 总被引:5,自引:0,他引:5
5.
Kenneth Heber 《BMJ (Clinical research ed.)》1965,2(5467):943-944
6.
Inhibition of zeaxanthin formation and of rapid changes in radiationless energy dissipation by dithiothreitol in spinach leaves and chloroplasts 总被引:22,自引:11,他引:11
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Demmig-Adams B Adams WW Heber U Neimanis S Winter K Krüger A Czygan FC Bilger W Björkman O 《Plant physiology》1990,92(2):293-301
Dithiothreitol, which completely inhibits the de-epoxidation of violaxanthin to zeaxanthin, was used to obtain evidence for a causal relationship between zeaxanthin and the dissipation of excess excitation energy in the photochemical apparatus in Spinicia oleracea L. In both leaves and chloroplasts, inhibition of zeaxanthin formation by dithiothreitol was accompanied by inhibition of a component of nonphotochemical fluorescence quenching. This component was characterized by a quenching of instantaneous fluorescence (Fo) and a linear relationship between the calculated rate constant for radiationless energy dissipation in the antenna chlorophyll and the zeaxanthin content. In leaves, this zeaxanthin-associated quenching, which relaxed within a few minutes upon darkening, was the major component of nonphotochemical fluorescence quenching determined in the light, i.e. it represented the `high-energy-state' quenching. In isolated chloroplasts, the zeaxanthin-associated quenching was a smaller component of total nonphotochemical quenching and there was a second, rapidly reversible high-energy-state component of fluorescence quenching which occurred in the absence of zeaxanthin and was not accompanied by Fo quenching. Leaves, but not chloroplasts, were capable of maintaining the electron acceptor, Q, of photosystem II in a low reduction state up to high degrees of excessive light and thus high degrees of nonphotochemical fluorescence quenching. When ascorbate, which serves as the reductant for violaxanthin de-epoxidation, was added to chloroplast suspensions, zeaxanthin formation at low photon flux densities was stimulated and the relationship between nonphotochemical fluorescence quenching and the reduction state in chloroplasts then became more similar to that found in leaves. We conclude that the inhibition of zeaxanthin-associated fluorescence quenching by dithiothreitol provides further evidence that there exists a close relationship between zeaxanthin and potentially photoprotective dissipation of excess excitation energy in the antenna chlorophyll. 相似文献
7.
The rapid transients of CO2 gas exchange have been measured in leaves ofHelianthus annuus L. In parallel experiments the assimilatory force FA, which is the product of the phosphorylation potential and the redox ratio NADPH/NADP, has been calculated from measured
ratios of dihydroxyacetone phosphate to phosphoglycerate in the chloroplast stroma and in leaves. The following results were
obtained: (i) When the light-dependent stroma alkalization was measured under steady-state conditions for photosynthesis in
air containing 2000 μl · l-1 CO2, alkalization increased with photosynthesis as the quantum flux density (irradiance) was increased. This contrasts to the
light-dependent stroma alkalisation measured in dark-adapted leaves during the dark-light transient (Laisk et al. 1989, Planta177, 350–358) which reached a maximum at a quantum flux density far below that necessary to saturate photosynthesis. This maximum
was about three times higher than the maximum stroma alkalization at light- and CO2-saturated photosynthesis. (ii) Accurate calculations of the assimilatory force FA require a consideration of the stromal pH. However, under many conditions, changes in the stromal pH resulting from changes
in photosynthetic flux can be neglected because they are small. (iii) Stromal ratios of dihydroxyacetone phosphate to phosphoglycerate
are generally lower than ratios measured in leaf extracts. The value of FA calculated from stromal metabolites was about 30% lower than FA calculated from cellular metabolites. Still, it appears sufficient for many purposes to calculate FA from metabolite measurements in leaf extracts. (iv) In the light, the catalytic capacity of the photosynthetic apparatus
is adjusted to the level of irradiance. The response of carbon assimilation to large increases in irradiance is slow because
it requires enzyme activation. Deactivation of the Calvin cycle induced by decreases in irradiance is slower than activation.
(v) Changes in catalytic capacity and in the availability or level of substrates such as CO2 alter the flux resistance of the Calvin cycle. A decrease in flux resistance explains why FA often does not increase by much and may actually decrease when carbon flux is increased. Adjustments of flux resistances
in the Calvin cycle and of photosystem-II activity in the electron-transport chain permit varying rates of photosynthesis
at low levels of ATP and NADPH. As NADP remains available, the danger of over-reduction which leads to photoinactivation of
electron transport is minimized.
K.R. und U.H. were guests of the Estonian Academy of Sciences. Support by the Estonian Academy of Sciences, the Sonderforschungsbereich
251 of the University of Würzburg and the Fonds der Chemischen Industrie is gratefully acknowledged. 相似文献
8.
Nikolai G. Bukhov Christian Wiese Spidola Neimanis Ulrich Heber 《Photosynthesis research》1996,50(2):181-191
The light-induced induction of components of non-photochemical quenching of chlorophyll fluorescence which are distinguished by different rates of dark relaxation (qNf, rapidly relaxing and qNs, slowly relaxing or not relaxing at all in the presence brief saturating light pulses which interrupt darkness at low frequencies) was studied in leaves of spinach.After dark adaptation of the leaves, a fast relaxing component developed in low light only after a lag phase. Quenching increased towards a maximum with increasing photon flux density. This fast component of quenching was identified as energy-dependent quenching qE. It required formation of an appreciable transthylakoid pH and was insignificant when darkened spinach leaves received 1 s pulses of light every 30 s even though zeaxanthin was formed from violaxanthin under these conditions.Another quenching component termed qNs developed in low light without a lag phase. It was not dependent on a transthylakoid pH gradient, decayed exponentially with a long half time of relaxation and was about 20% of total quenching irrespective of light intensity. When darkened leaves were flashed at frequencies higher than 0.004 Hz with 1 s light pulses, this quenching also appeared. Its extent was very considerable, and it did not require formation of zeaxanthin. Relaxation was accelerated by far-red light, and this acceleration was abolished by NaF.We suggest that qNs is the result of a so-called state transition, in which LHC II moves after its phosphorylation from fluorescent PS II to nonfluorescent PS I. This state transition was capable of decreasing in darkened leaves the potential maximum quantum efficiency of electron flow through Photosystem II by about 20%.Abbreviations PFD
photon flux density
- PS
photosystem 相似文献
9.
Atmospheric ammonia (NH3) from various anthropogenic sources has become a serious problem for natural vegetation. Ammonia not only causes changes in plant nitrogen metabolism, but also affects the acid-base balance of plants. Using the pH-sensitive fluorescent dyes pyranine and esculin, cytosolic and vacuolar pH changes were measured in leaves of C3 and C4 plants exposed for brief periods to concentrations of NH3 in air ranging from 1.33 to 8.29 mol NH3 · mol-1 gas (0.94–5.86 mg · m-3). After a lag phase, uptake of NH3 from air at a rate of 200 nmol NH3 · m - 2 leaf area · s- 1 into leaves of Zea mays L. increased pyranine fluorescence indicating cytosolic alkalinisation. The increase was much larger in the dark than in the light. In illuminated leaves of the C3 plant Pelargonium zonale L. and the C4 plants Z. mays and Amaranthus caudatus L., NH3-dependent cytosolic alkalinisation was particularly pronounced when CO2 was supplied at very low levels (16 or 20 mol CO2 · mol- 1 gas, containing 210 mmol O2 · mol- 1 gas). An increase in esculin fluorescence, which was smaller than that of pyranine, was indicative of trapping of some of the NH3 in the vacuoles of leaves of Spinacia oleracea L. and Z. mays. Photosynthesis and transpiration remained unchanged during exposure of illuminated leaves to NH3, yielding an influx of 200 nmol NH3 · m-2 leaf area · s-1 for up to 30 min, the longest exposure time used. Both CO2 and O2 influenced the extent of cytosolic alkalinisation. At 500 mol CO2 · mol-1 gas the cytosolic alkalinisation was suppressed more than at 16 or 20 mol CO2 · mol-1 gas. The suppressing effect of CO2 on the NH3induced alkalinisation was larger in illuminated leaves of the C4 plants Z. mays and A. caudatus than in leaves of the C3 plant P. zonale. A reduction of the O2 concentration from 210 to 10 mmol O2 · mol -1 gas, which inhibits photorespiration, increased the NH3induced cytosolic alkalinisation in C3 plants. Suppression by CO2 or O2 of the alkaline pH shift caused by the dissolution and protonation of NH3 in queous leaf compartments, and possibly by the production of organic compounds synthesised from atmospheric NH3, indicates that NH3 which enters leaves is rapidly assimilated if photosynthesis or photorespiration provide nitrogen acceptor molecules.This work was supported by the Biotechnology and Biological Sciences Research Council and the Deutsche Forschungsgemein-schaft within the framework of the research of Sonderforschun-gsbreich 251 of the University of Würzburg. We are grateful to Dr. B. Wollenweber (The Royal Veterinary and Agricultural University, Denmark) for discussions. 相似文献
10.
The fat on women's thighs is more difficult to mobilize due to increased α-2 adrenergic receptor activity induced by estrogen. Lipolysis can be initiated through adipocyte receptor stimulation (β adrenergic) or inhibition (adenosine or α-2 adrenergic) or by inhibition of phosphodiesterase. Since many women desire regional thigh fat loss, a series of clinical trials were initiated using one thigh as a double-blinded control. Trial #1: Five overweight women had injections of isoproterenol at intervals around the thigh three times a week for 4 weeks with diet and walking. Trial #2: Five overweight woman had ointment containing forskolin, yohimbine and aminophylline applied to the thigh five times a week for 4 weeks after hypertonic warm soaks with a diet and walking. Trial #3: Eighteen overweight women were divided into three groups of six and trial #2 was repeated with each agent alone vs. placebo using forskolin, yohimbine or aminophylline in separate ointments. Trial #4: Thirty overweight women had 10% aminophylline ointment applied to the thigh five times a week for 6 weeks with diet and walking. Chemistry panel, theophylline level and patch testing wereperformed. Trial#5: Twelve women had trial #4 repeated with 2% aminophylline cream without a diet or walking. Trial #6: Trial #5 was repeated with 0.5% aminophylline cream. All trials except yohimbine ointment gave significantly more girth loss from the treated thigh (p < 0.05 to p < 0.001). Chemistry panel showed no toxicity. Theophylline was undetectable and patch testing was negative. We conclude that topical fat reduction for women's thighs can be achieved without diet or exercise. 相似文献