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Electron Transport in Chloroplasts: Regulation and Alternative Pathways of Electron Transfer

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Abstract

This work represents an overview of electron transport regulation in chloroplasts as considered in the context of structure-function organization of photosynthetic apparatus in plants. Main focus of the article is on bifurcated oxidation of plastoquinol by the cytochrome b6f complex, which represents the rate-limiting step of electron transfer between photosystems II and I. Electron transport along the chains of non-cyclic, cyclic, and pseudocyclic electron flow, their relationships to generation of the trans-thylakoid difference in electrochemical potentials of protons in chloroplasts, and pH-dependent mechanisms of regulation of the cytochrome b6f complex are considered. Redox reactions with participation of molecular oxygen and ascorbate, alternative mediators of electron transport in chloroplasts, have also been discussed.

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Abbreviations

Asc, MDHA, DHA:

three redox forms of ascorbate (fully reduced, semiquinone, and completely oxidized)

CBC:

Calvin–Benson cycle

CET:

cyclic electron transport

EPR:

electron paramagnetic resonance

ETC:

electron transport chain

ISP:

iron-sulfur protein, part of PSI

Fd:

ferredoxin

FNR:

ferredoxin-NADP reductase

NDH-1:

NAD(P)H-dehydrogenase of chloroplasts type 1

P700 and P680 :

primary electron donors in PSI and PSII

Pc:

plastocyanin

PGR5 and PGRL1:

proteins involved in cyclic electron transfer around PSI

PSI and PSII:

photosystem I and photosystem II

PQ:

plastoquinone

PQH2 :

plastoquinol

PTOX:

plastid (plastoquinol) terminal oxidase

ROS:

reactive oxygen species

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Acknowledgments

This article is dedicated to the memory of L. A. Drachev – one of the greatest Russian experimental biophysicists, whose scientific and technical developments and pioneering research in the field of photosynthesis contributed to a deeper understanding of the processes of charge separation and transfer in the reaction centers of photosynthetic systems.

The author is grateful to E. K. Ruuge, G. B. Khomutov, and L. Yu. Ustynyuk, together with whom the main results of the experimental and theoretical study of the regulation of electron transport in chloroplasts were previously obtained, to which the author refers in this review. The author thanks the anonymous reviewers for their helpful comments and recommendations.

Funding

This work was carried out within the framework of the research topic of the Faculty of Physics of Lomonosov Moscow State University “Physical Foundations of the Structure, Functioning and Regulation of Biological Systems” (State registration no. 012004 085 35) and with partial financial support from the Russian Foundation for Basic Research (grant no. 21-04-20047).

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    Alexander N. Tikhonov

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Tikhonov, A.N. Electron Transport in Chloroplasts: Regulation and Alternative Pathways of Electron Transfer. Biochemistry Moscow 88, 1438–1454 (2023). https://doi.org/10.1134/S0006297923100036

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