Abstract
This chapter presents an overview of structural properties of the cytochrome (Cyt) b 6 f complex and its functioning in chloroplasts. The Cyt b 6 f complex stands at the crossroad of photosynthetic electron transport pathways, providing connectivity between Photosystem (PSI) and Photosysten II (PSII) and pumping protons across the membrane into the thylakoid lumen. After a brief review of the chloroplast electron transport chain, the consideration is focused on the structural organization of the Cyt b 6 f complex and its interaction with plastoquinol (PQH2, reduced form of plastoquinone), a mediator of electron transfer from PSII to the Cyt b 6 f complex. The processes of PQH2 oxidation by the Cyt b 6 f complex have been considered within the framework of the Mitchell’s Q-cycle. The overall rate of the intersystem electron transport is determined by PQH2 turnover at the quinone-binding site Qo of the Cyt b 6 f complex. The rate of PQH2 oxidation is controlled by the intrathylakoid pHin, which value determines the protonation/deprotonation events in the Qo-center. Two other regulatory mechanisms associated with the Cyt b 6 f complex are briefly overviewed: (i) redistribution of electron fluxes between alternative (linear and cyclic) pathways, and (ii) “state transitions” related to redistribution of solar energy between PSI and PSII.
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- 1.
Here and below, the general terms QH2 and Q are used to denote the quinol and quinone species, regardless of their origin (plastoquinone, PQ, or ubiquinone, UQ). SQ designates the redox states of semiquinone species, either plastosemiquinone (in the b 6 f complex) or ubisemiquinone (in the bc 1 complex), regardless of their protonation state.
- 2.
It is likely that this species will appear in the form of the anion-radical \( {\mathrm{PQ}}^{\underset{\_}{\bullet }} \), because the pK values of semiquinones are usually fall in the range below the stromal pH establishes under the normal physiological conditions (pHout~7 − 8).
Abbreviations
- CBC:
-
Calvin-Benson cycle
- CEF1:
-
cyclic electron flow around photosystem I
- ETC:
-
electron transport chain
- Fd:
-
ferredoxin
- FNR:
-
ferredoxin-NADP-oxidoreductase
- ISP:
-
iron-sulfur protein
- LEF:
-
linear electron flow
- LHCI:
-
light-harvesting complex I
- LHCII:
-
light-harvesting complex II
- NDH:
-
NDH(P)H dehydrogenase complex
- NPQ:
-
non-photochemical quenching
- P680 :
-
special chlorophyll pair in PSII
- P700 :
-
special chlorophyll pair in PSI
- Pc:
-
plastocyanin
- PCET:
-
proton-coupled electron transfer
- PQ and PQH2 :
-
plastoquinone and plastoquinol, respectively
- PSA:
-
photosynthetic apparatus
- PSI and PSII:
-
photosystem I and photosystem II respectively
- Q, SQ and QH2 :
-
general notations for three accessible redox states of quinone species – quinone (Q), semiquinone (SQ) and quinol (QH2)
- TMQH2 :
-
trimethylbenzoquinol
- UQ and UQH2 :
-
ubiquinone and ubiquinol, respectively
- WOC:
-
water-oxidizing complex
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Acknowledgments
This work was supported in part by the Russian Foundation for Basic Research (RFBR Project 15-04-03790a).
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Tikhonov, A.N. (2018). The Cytochrome b 6 f Complex: Biophysical Aspects of Its Functioning in Chloroplasts. In: Harris, J., Boekema, E. (eds) Membrane Protein Complexes: Structure and Function. Subcellular Biochemistry, vol 87. Springer, Singapore. https://doi.org/10.1007/978-981-10-7757-9_10
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