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Determining photosynthetic control, a probe for the balance between electron transport and Calvin–Benson cycle activity, with the DUAL-KLAS-NIR

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Abstract

Photosynthetic Control is defined as the control imposed on photosynthetic electron transport by the lumen-pH-sensitive re-oxidation of plastoquinol (PQH2) by cytochrome b6f. Photosynthetic Control leads at higher actinic light intensities to an electron transport chain with a (relatively) reduced photosystem (PS) II and PQ pool and a (relatively) oxidized PS I. Making Light Curves of more than 33 plant species with the recently introduced DUAL-KLAS-NIR (Chl a fluorescence + the redox states of plastocyanin (PC), P700, and ferredoxin (Fd)) the light intensity-dependent induction of Photosynthetic Control was probed and characterized. It was observed that PC became completely oxidized at light intensities ≤ 400 µmol photons m−2 s−1 (at lower light intensities in shade than in sun leaves). The relationship between qP and P700(red) was used to determine the extent of Photosynthetic Control. Instead of measuring the whole Light Curve, it was shown that a single moderate light intensity can be used to characterize the status of a leaf relative to that of other leaves. It was further found that in some shade-acclimated leaves Fd becomes again more oxidized at high light intensities indicating that electron transfer from the PQ pool to P700 cannot keep up with the outflow of electrons on the acceptor side of PS I. It was observed as well that for NPQ-induction a lower light intensity (less acidified lumen) was needed than for the induction of Photosynthetic Control. The measurements were also used to make a comparison between the parameters qP and qL, a comparison suggesting that qP was the more relevant parameter.

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Abbreviations

CCCP:

Carbonyl cyanide m-chlorophenylhydrazone

CET:

Cyclic electron transport

Cyt:

Cytochrome

DCMU:

3-(3,4-Dichlorophenyl)-1,1-dimethylurea

EPR:

Electron paramagnetic resonance

FCCP:

Carbonyl cyanide-p-trifluoromethoxyphenylhydrazone

Fd:

Ferredoxin

FeS cluster:

Iron-sulfur cluster of photosystem I

F M :

Maximum fluorescence yield

F 0, F 0′:

Minimum fluorescence yield of a dark and light acclimated sample, respectively

F V/F M :

Maximum quantum yield of photosystem II

MV:

Methylviologen

NIR:

Near infra-red

NPQ:

Non-photochemical quenching

O-I1-I2-P, O-J-I-P:

Fluorescence induction kinetics defined by their different kinetic steps

P700:

Reaction center of photosystem I

PC:

Plastocyanin

ΦPSI, ΦPSII :

Operational quantum yield of photosystem I and II, respectively

PQ, PQH2 :

Plastoquinone, plastoquinol

PS I, PS II:

Photosystem I and II, respectively

Q A :

First quinone electron acceptor of photosystem II

qE:

(Non-photochemical) energy quenching

qL:

Photochemical quenching (assuming a lake model)

qP:

Photochemical quenching (assuming a puddle model)

Y(I):

(Operational) quantum yield of photosystem I

Y(NA):

PS I quantum yield loss due to a PS I acceptor side limitation

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Acknowledgements

I am indebted to Drs. Christof Klughammer and Ulrich Schreiber, without their development of the DUAL-KLAS-NIR this study would not have been possible. I want to thank as well Dr. Erhard Pfündel for critical reading of the manuscript and Heinz Walz GmbH for enabling me to work on this paper.

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  1. Heinz Walz GmbH, Eichenring 6, 91090, Effeltrich, Germany

    Gert Schansker

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  1. Gert Schansker
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Correspondence to Gert Schansker.

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11120_2022_934_MOESM1_ESM.docx

Lack of effect of SP-intensity in the 3700–12000 µmol photons m-2 s-1 range on qP (A and C) and P700(red) determination (B and D) illustrated by measurements on Ribes sanguineum leaves. In each case data from a Light Curve consisting of 15 light steps of 2 min each are shown. In A, 5 experiments for which the F0′ values were calculated and in C, 5 experiments where the F0′ values were measured with a far-red routine following each saturation pulse. Supplementary file1 (DOCX 37 kb)

11120_2022_934_MOESM2_ESM.docx

Four examples of the relationship between P700(red) and qP and qL, respectively. In each case 5 Light Curves measured with 5 different SP-intensities between 3700 and 12000 µmol photons m-2 s-1 were averaged and the standard error determined. Leaves of the Phalaenopsis orchid had the smallest pool of PC and gave the best linear regression for both qP and qL versus P700(red). The calculated qP and qL values are in three out of four cases systematically lower than the measured values. Not in all cases qL yields a good correlation with P700(red). Supplementary file2 (DOCX 68 kb)

11120_2022_934_MOESM3_ESM.xlsx

Set of light curves that were used for the determination of the datapoints in Fig. 6. The figure represents light curves of 22 plant species of the initially measured dataset. Supplementary file3 (XLSX 72 kb)

11120_2022_934_MOESM4_ESM.docx

Effect of changes in the PC-redox state on qP. The impact is visualized for 4 different plants by showing the light intensity dependence of P700(red), PC(red) and qP simultaneously. At the lowest light intensities P700(red) remained nearly completely reduced and all changes in qP were due to PC redox changes. The standard deviation is indicated (n = 7-10). A line was drawn through the point where P700(red) starts to decline. Supplementary file4 (DOCX 42 kb)

11120_2022_934_MOESM5_ESM.docx

Same experiment as in Fig. 9, showing data on four additional plant species to illustrate the variability in the relationship between NPQ and Y(I), P700(red) and P700(red)+PC(red). In panels A, C, E and G the NPQ was determined on the basis of fluorescence measured through the leaf using green measuring light and in B, D, F and H the NPQ was determined on the basis of blue measuring light applied to the top of the leaf. In the inset of A, the qN is plotted as a function of qP. Five Light Curves measured with 5 different SP-intensities between 3700 and 12000 µmol photons m-2 s-1 were averaged and the standard error determined. The light intensity dependence is derived from light curves with 2 min per light step. Supplementary file5 (DOCX 100 kb)

11120_2022_934_MOESM6_ESM.pdf

ΦPSII and qP as a function of P700(red) for three plants taken from 3 different classes out of Table 1 (bean = class 1, Begonia = 4 and Rumex = 5). The light intensity dependence is derived from light curves with 2 min per light step. N = 3-7. Supplementary file6 (PDF 16 kb)

Supplementary file7 (DOCX 18 kb)

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Schansker, G. Determining photosynthetic control, a probe for the balance between electron transport and Calvin–Benson cycle activity, with the DUAL-KLAS-NIR. Photosynth Res 153, 191–204 (2022). https://doi.org/10.1007/s11120-022-00934-7

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