Control of photosynthesis in leaves as revealed by rapid gas exchange and measurements of the assimilatory force FA |
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Authors: | K Siebke A Laisk V Oja O Kiirats K Raschke U Heber |
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Institution: | 1. Institut für Botanik und Pharmazeutische Biologie der Universit?t, W-8700, Würzburg, Federal Republic of Germany 2. Astrofüüsika Ja Atmosf??rifüüsika Instutuut, Eesti NSV Teaduste Akadeemia, 202444, Toravere, Tartu, Eesti, USSR 3. Pflanzenphysiologisches Institut der Universit?t, W-3400, G?ttingen, Federal Republic of Germany
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Abstract: | 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. |
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Keywords: | Assimilatory force Enzyme regulation Gas exchange Helianthus (photosynthesis) Photosynthesis Proton gradient Stroma alkalization |
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