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Structure, regulation and assembly of the photosynthetic electron transport chain

Nature Reviews Molecular Cell Biology volume 26, pages 667–690 (2025)Cite this article

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An Author Correction to this article was published on 16 July 2025

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Abstract

The electron transfer chain of chloroplast thylakoid membranes uses solar energy to split water into electrons and protons, creating energetic gradients that drive the formation of photosynthetic fuel in the form of NADPH and ATP. These metabolites are then used to power the fixation of carbon dioxide into biomass through the Calvin–Benson–Bassham cycle in the chloroplast stroma. Recent advances in molecular genetics, structural biology and spectroscopy have provided an unprecedented understanding of the molecular events involved in photosynthetic electron transfer from photon capture to ATP production. Specifically, we have gained insights into the assembly of the photosynthetic complexes into larger supercomplexes, thylakoid membrane organization and the mechanisms underpinning efficient light harvesting, photoprotection and oxygen evolution. In this Review, I focus on the angiosperm plant thylakoid system, outlining our current knowledge on the structure, function, regulation and assembly of each component of the photosynthetic chain. I explain how solar energy is harvested and converted into chemical energy by the photosynthetic electron transfer chain, how its components are integrated into a complex membrane macrostructure and how this organization contributes to regulation and photoprotection.

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Fig. 1: Overview of the photosynthetic electron transfer chain and Calvin–Benson–Bassham cycle.
Fig. 2: Light-harvesting antenna structure of the photosystems.
Fig. 3: Catalytic mechanisms of electron transfer chain components.
Fig. 4: Lateral heterogeneity in thylakoid organization.
Fig. 5: Regulation of the photosynthetic electron transfer chain.
Fig. 6: Assembly pathway for the four core electron transfer chain components.

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Acknowledgements

M.P.J. acknowledges funding from the Leverhulme Trust grants RPG-2019-045 and RPG-2021-345 and Biotechnology and Biological Sciences Research Council (BBSRC) UK grants BB/V006630/1 and BB/W015269/1. M.P.J. would also like to thank B. Engel and W. Wietrzynski for collaboration in producing the spinach chloroplast tomograms, and A. Ruban, B. Rutherford, P. Nixon, N. Hunter and A. Hitchcock for useful discussions.

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  1. Plants, Photosynthesis and Soil, School of Biosciences, University of Sheffield, Sheffield, UK

    Matthew P. Johnson

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Glossary

C4 type photosynthesis

This is a more efficient type of photosynthesis that concentrates carbon dioxide around the enzyme Rubisco.

Jablonski diagram

A diagram that illustrates the electronic states and often the vibrational levels of a molecule.

Serial femtosecond crystallography

A form of X-ray crystallography developed for use at X-ray free electron lasers (XFELs). Diffraction patterns from sub-micrometre crystals can be generated via single pulses of XFELs. However, the pulses are so intense that the crystals are destroyed, meaning data must be collected from many crystals in series.

Signal recognition particle pathway

A cellular mechanism that directs nascent proteins with a signal sequence to the chloroplast thylakoid membrane by binding to the signal sequence as it emerges from the ribosome and then docking the ribosome-nascent chain complex with the thylakoid membrane through a receptor, allowing for co-translational protein targeting and translocation across the membrane.

Twin-arginine translocation pathway

A protein transport system that moves folded proteins across the thylakoid membrane.

X-ray free electron lasers

This approach combines the properties of X-ray sources with the properties of lasers, utilized for obtaining high-resolution structural and chemical information on isolated molecules.

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Johnson, M.P. Structure, regulation and assembly of the photosynthetic electron transport chain. Nat Rev Mol Cell Biol 26, 667–690 (2025). https://doi.org/10.1038/s41580-025-00847-y

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