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Relative contributions of PGR5- and NDH-dependent photosystem I cyclic electron flow in the generation of a proton gradient in Arabidopsis chloroplasts

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Main conclusion

Respective contributions of PGR5- and NDH-dependent cyclic electron flows around photosystem I for generating the proton gradient across the thylakoid membrane are ~30 and ~5%.

The proton concentration gradient across the thylakoid membrane (ΔpH) produced by photosynthetic electron transport is the driving force of ATP synthesis and non-photochemical quenching. Two types of electron transfer contribute to ΔpH formation: linear electron flow (LEF) and cyclic electron flow (CEF, divided into PGR5- and NDH-dependent pathways). However, the respective contributions of LEF and CEF to ΔpH formation are largely unknown. We employed fluorescence quenching analysis with the pH indicator 9-aminoacridine to directly monitor ΔpH formation in isolated chloroplasts of Arabidopsis mutants lacking PGR5- and/or NDH-dependent CEF. The results indicate that ΔpH formation is mostly due to LEF, with the contributions of PGR5- and NDH-dependent CEF estimated as only ~30 and ~5%, respectively.

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Fig. 1
Fig. 2
Fig. 3



Cyclic electron flow


Proton concentration gradient across the inside and outside of the thylakoid membrane


Membrane potential


Linear electron flow


NAD(P)H dehydrogenase


Non-photochemical quenching


Proton gradient regulation 5


Proton motive force


Photosystem I


Photosystem II




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We thank Professor Toshiharu Shikanai at Kyoto University for providing mutant seeds. This work was supported by a Grant-in-Aid for Scientific Research, KAKENHI (17H05719, 16K14694 and 16H03280) to SM.

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Correspondence to Shinji Masuda.

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Kawashima, R., Sato, R., Harada, K. et al. Relative contributions of PGR5- and NDH-dependent photosystem I cyclic electron flow in the generation of a proton gradient in Arabidopsis chloroplasts. Planta 246, 1045–1050 (2017). https://doi.org/10.1007/s00425-017-2761-1

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  • Arabidopsis
  • Cyclic electron transfer
  • Non-photochemical quenching
  • PGR5
  • Photosynthesis
  • ΔpH