1. Asada, K. The water–water cycle as alternative photon and electron sinks. Philos. Trans. R. Soc. B Biol. Sci. 2000, 355, 1419–1431. [CrossRef] [PubMed]
2. Hamilton, T.L. The trouble with oxygen: The ecophysiology of extant phototrophs and implications for the evolution of oxygenic photosynthesis. Free Radic. Biol. Med. 2019, 140, 233–249. [CrossRef] [PubMed]
3. Mattila, H.; Khorobrykh, S.; Havurinne, V.; Tyystjärvi, E. Reactive oxygen species: Reactions and detection from photosynthetic tissues. J. Photochem. Photobiol. B Biol. 2015, 152, 176–214. [CrossRef]
4. Khorobrykh, S.; Havurinne, V.; Mattila, H.; Tyystjärvi, E. Oxygen and ROS in photosynthesis. Plants 2020, 9, 91. [CrossRef] [PubMed]
5. Cherepanov, D.A.; Milanovsky, G.E.; Petrova, A.A.; Tikhonov, A.N.; Semenov, A.Y. Electron transfer through the acceptor side of photosystem I: Interaction with exogenous acceptors and molecular oxygen. Biochemistry 2017, 82, 1249–1268. [CrossRef]
6. Mehler, A.H. Studies on reactions of illuminated chloroplasts. I. Mechanism of the reduction of oxygen and other Hill reagents. Arch. Biochem. Biophys. 1951, 33, 65–77. [CrossRef]
7. Mehler, A.H. Studies on reactions of illuminated chloroplasts. II. Stimulation and inhibition of the reaction with molecular oxygen. Arch. Biochem. Biophys. 1951, 34, 339–351. [CrossRef] [PubMed]
8. Mehler, A.H.; Brown, A.H. Studies on reactions of illuminated chloroplasts. III. Simultaneous photoproduction and consumption of oxygen studied with oxygen isotopes. Arch. Biochem. Biophys. 1952, 38, 365–370. [CrossRef]
9. Asada, K.; Kiso, K. The photo-oxidation of epinephrine by spinach chloroplasts and its inhibition by superoxide dismutase: Evidence for the formation of superoxide radicals in chloroplasts. Agri. Biol. Chem. 1973, 37, 453–454. [CrossRef]
10. Asada, K.; Kiso, K.; Yoshikawa, K. Univalent reduction of molecular oxygen by spinach chloroplasts on illumination. J. Biol. Chem. 1974, 249, 2175–2181. [CrossRef]
11. Takahashi, M.; Asada, K. Superoxide production in aprotic interior of chloroplast thylakoids. Arch. Biochem. Biophys. 1988, 267, 714–722. [CrossRef] [PubMed]
12. Kozuleva, M.; Petrova, A.; Milrad, Y.; Semenov, A.; Ivanov, B.; Redding, K.E.; Yacoby, I. Phylloquinone is the principal Mehler reaction site within photosystem I in high light. Plant Physiol. 2021, 186, 1848–1858. [CrossRef] [PubMed]
13. Kozuleva, M.A.; Petrova, A.A.; Mamedov, M.D.; Semenov, A.Y.; Ivanov, B.N. O2 reduction by photosystem I involves phylloquinone under steady-state illumination. FEBS Lett. 2014, 588, 4364–4368. [CrossRef] [PubMed]
14. Kruk, J.; Jemioła-Rzemi ´nska, M.; Burda, K.; Schmid, G.H.; Strzałka, K. Scavenging of superoxide generated in photosystem I by plastoquinol and other prenyllipids in thylakoid membranes. Biochemistry 2003, 42, 8501–8505. [CrossRef]
15. Sejima, T.; Takagi, D.; Fukayama, H.; Makino, A.; Miyake, C. Repetitive short-pulse light mainly inactivates photosystem I in sunflower leaves. Plant Cell Physiol. 2014, 55, 1184–1193. [CrossRef]
16. Zivcak, M.; Brestic, M.; Kunderlikova, K.; Olsovska, K.; Allakhverdiev, S.I. Effect of photosystem I inactivation on chlorophyll a fluorescence induction in wheat leaves: Does activity of photosystem I play any role in OJIP rise? J. Photochem. Photobiol. B Biol. 2015, 152, 318–324. [CrossRef]
17. Takahashi, M.; Asada, K. Dependence of oxygen affinity for Mehler reaction on photochemical activity of chloroplast thylakoids. Plant Cell Physiol. 1982, 23, 1457–1461.
18. Takahashi, M.; Asada, K. Superoxide anion permeability of phospholipid membranes and chloroplast thylakoids. Arch. Biochem. Biophys. 1983, 226, 558–566. [CrossRef]
19. Munekage, Y.; Hojo, M.; Meurer, J.; Endo, T.; Tasaka, M.; Shikanai, T. PGR5 is involved in cyclic electron flow around photosystem I and is essential for photoprotection in Arabidopsis. Cell 2002, 110, 361–371. [CrossRef]
20. Wada, S.; Amako, K.; Miyake, C. Identification of a novel mutation exacerbated the PSI photoinhibition in pgr5/pgrl1 mutants; Caution for overestimation of the phenotypes in Arabidopsis pgr5-1 mutant. Cells 2021, 10, 2884. [CrossRef]
21. Furutani, R.; Ohnishi, M.; Mori, Y.; Wada, S.; Miyake, C. The difficulty of estimating the electron transport rate at photosystem I. J. Plant Res. 2021, 135, 565–577. [CrossRef] [PubMed]
22. Klughammer, C.; Schreiber, U. Deconvolution of ferredoxin, plastocyanin, and P700 transmittance changes in intact leaves with a new type of kinetic LED array spectrophotometer. Photosynth. Res. 2016, 128, 195–214. [CrossRef] [PubMed]
23. Sétif, P.; Boussac, A.; Krieger-Liszkay, A. Near-infrared in vitro measurements of photosystem I cofactors and electron-transfer partners with a recently developed spectrophotometer. Photosynth. Res. 2019, 142, 307–319. [CrossRef] [PubMed]
24. Baker, N.R.; Harbinson, J.; Kramer, D. Determining the limitations and regulation of photosynthetic energy transduction in leaves. Plant Cell Environ. 2007, 30, 1107–1125. [CrossRef]
25. Shimakawa, G.; Miyake, C. Oxidation of P700 ensures robust photosynthesis. Front. Plant Sci. 2018, 9, 1617. [CrossRef]
26. Takagi, D.; Ishizaki, K.; Hanawa, H.; Mabuchi, T.; Shimakawa, G.; Yamamoto, H.; Miyake, C. Diversity of strategies for escaping reactive oxygen species production within photosystem I among land plants: P700 oxidation system is prerequisite for alleviating photoinhibition in photosystem I. Physiol. Plant. 2017, 161, 56–74. [CrossRef]
27. Kadota, K.; Furutani, R.; Makino, A.; Suzuki, Y.; Wada, S.; Miyake, C. Oxidation of P700 induces alternative electron flow in Photosystem I in wheat leaves. Plants 2019, 8, 152. [CrossRef]
28. Sétif, P.; Shimakawa, G.; Krieger-Liszkay, A.; Miyake, C. Identification of the electron donor to flavodiiron proteins in Synechocystis sp. PCC 6803 by in vivo spectroscopy. Biochim. Biophys. Acta BBA Bioenerg. 2020, 1861, 148256. [CrossRef]
29. Johnson, M.P.; Ruban, A.V. Rethinking the existence of a steady-state ∆ψ component of the proton motive force across plant thylakoid membranes. Photosynth. Res. 2013, 119, 233–242. [CrossRef]
30. Shikanai, T.; Müller-Moulé, P.; Munekage, Y.; Niyogi, K.K.; Pilon, M. PAA1, a P-Type ATPase of Arabidopsis, functions in copper transport in chloroplasts. Plant Cell 2003, 15, 1333–1346. [CrossRef]
31. Tikkanen, M.; Grieco, M.; Nurmi, M.; Rantala, M.; Suorsa, M.; Aro, E.-M. Regulation of the photosynthetic apparatus under fluctuating growth light. Philos. Trans. R. Soc. B Biol. Sci. 2012, 367, 3486–3493. [CrossRef] [PubMed]
32. Tikkanen, M.; Rantala, S.; Aro, E.-M. Electron flow from PSII to PSI under high light is controlled by PGR5 but not by PSBS. Front. Plant Sci. 2015, 6, 521. [CrossRef] [PubMed]
33. Satoh, K. Mechanism of photoinactivation in photosynthetic systems II. The occurrence and properties of two different types of photoinactivation. Plant Cell Physiol. 1970, 11, 29–38. [CrossRef]
34. Satoh, K. Mechanism of photoinactivation in photosynthetic systems. III. The site and mode of photoinactivation in photosystem I. Plant Cell Physiol. 1970, 11, 187–197. [CrossRef]
35. Inoue, K.; Fujii, T.; Yokoyama, E.; Matsuura, K.; Hiyama, T.; Sakurai, H. The Photoinhibition site of photosystem I in isolated chloroplasts under extremely reducing conditions. Plant Cell Physiol. 1989, 30, 65–71. [CrossRef]
36. Inoue, K.; Sakurai, H.; Hiyama, T. Photoinactivation sites of photosystem I in isolated chloroplasts. Plant Cell Physiol. 1986, 27, 961–968.
37. Havaux, M.; Davaud, A. Photoinhibition of photosynthesis in chilled potato leaves is not correlated with a loss of Photosystem-II activity. Photosynth. Res. 1994, 40, 75–92. [CrossRef]
38. Sonoike, K.; Terashima, I. Mechanism of photosystem-I photoinhibition in leaves of Cucumis sativus L. Planta 1994, 194, 287–293. [CrossRef]
39. Sonoike, K.; Terashima, I.; Iwaki, M.; Itoh, S. Destruction of photosystem I iron-sulfur centers in leaves of Cucumis sativus L. by weak illumination at chilling temperatures. FEBS Lett. 1995, 362, 235–238. [CrossRef]
40. Terashima, I.; Funayama, S.; Sonoike, K. The site of photoinhibition in leaves of Cucumis sativus L. at low temperatures is photosystem I, not photosystem II. Planta 1994, 193, 300–306. [CrossRef]
41. Ivanov, A.G.; Morgan, R.M.; Gray, G.R.; Velitchkova, M.Y.; Huner, N.P. Temperature/light dependent development of selective resistance to photoinhibition of photosystem I. FEBS Lett. 1998, 430, 288–292. [CrossRef] [PubMed]
42. Takeuchi, K.; Che, Y.; Nakano, T.; Miyake, C.; Ifuku, K. The ability of P700 oxidation in photosystem I reflects chilling stress tolerance in cucumber. J. Plant Res. 2022, 135, 681–692. [CrossRef] [PubMed]
43. Scheller, H.V.; Haldrup, A. Photoinhibition of photosystem I. Planta 2005, 221, 5–8. [CrossRef] [PubMed]
44. Tjus, S.E.; Møller, B.L.; Scheller, H. Photosystem I is an early target of photoinhibition in barley illuminated at chilling Temperatures. Plant Physiol. 1998, 116, 755–764. [CrossRef]
45. Zhang, S.; Scheller, H. Photoinhibition of Photosystem I at chilling temperature and Subsequent Recovery in Arabidopsis thaliana. Plant Cell Physiol. 2004, 45, 1595–1602. [CrossRef]
46. Furutani, R.; Ifuku, K.; Suzuki, Y.; Noguchi, K.; Shimakawa, G.; Wada, S.; Makino, A.; Sohtome, T.; Miyake, C. P700 Oxidation Suppresses the Production of Reactive Oxygen Species in Photosystem I; Hisabori, T., Ed.; Academic Press: Cambridge, MA, USA, 2000; Volume 96, p. 26.
47. Furutani, R.; Makino, A.; Suzuki, Y.; Wada, S.; Shimakawa, G.; Miyake, C. Intrinsic fluctuations in transpiration induce photorespiration to oxidize P700 in photosystem I. Plants 2020, 9, 1761. [CrossRef]
48. Miyake, C. Molecular mechanism of oxidation of P700 and suppression of ROS production in photosystem I in response to electron-sink limitations in C3 Plants. Antioxidants 2020, 9, 230. [CrossRef]
49. Miyake, C.; Schreiber, U.; Hormann, H.; Sano, S.; Asada, K. The FAD-enzyme monodehydroascorbate radical reductase mediates photoproduction of superoxide radicals in spinach thylakoid membranes. Plant Cell Physiol. 1998, 39, 821–829. [CrossRef]
50. Ruuska, S.A.; Badger, M.R.; Andrews, T.J.; von Caemmerer, S. Photosynthetic electron sinks in transgenic tobacco with reduced amounts of Rubisco: Little evidence for significant Mehler reaction. J. Exp. Bot. 2000, 51, 357–368. [CrossRef]
51. Driever, S.M.; Baker, N.R. The water-water cycle in leaves is not a major alternative electron sink for dissipation of excess excitation energy when CO2 assimilation is restricted. Plant Cell Environ. 2011, 34, 837–846. [CrossRef]
52. Sonoike, K. Photoinhibition of photosystem I. Physiol. Plant. 2011, 142, 56–64. [CrossRef] [PubMed]
53. Suorsa, M.; Järvi, S.; Grieco, M.; Nurmi, M.; Pietrzykowska, M.; Rantala, M.; Kangasjärvi, S.; Paakkarinen, V.; Tikkanen, M.; Jansson, S.; et al. PROTON GRADIENT REGULATION5 is essential for proper acclimation of Arabidopsis photosystem I to naturally and artificially fluctuating light conditions. Plant Cell 2012, 24, 2934–2948. [CrossRef] [PubMed]
54. Hormann, H.; Neubauer, C.; Asada, K.; Schreiber, U. Intact chloroplasts display pH 5 optimum of O2 -reduction in the absence of methyl viologen: Indirect evidence for a regulatory role of superoxide protonation. Photosynth. Res. 1993, 37, 69–80. [CrossRef]
55. Hormann, H.; Neubauer, C.; Schreiber, U. An active Mehler-peroxidase reaction sequence can prevent cyclic PS I electron transport in the presence of dioxygen in intact spinach chloroplasts. Photosynth. Res. 1994, 41, 429–437. [CrossRef] [PubMed]
56. Flint, D.; Tuminello, J.; Emptage, M. The inactivation of Fe-S cluster containing hydro-lyases by superoxide. J. Biol. Chem. 1993, 268, 22369–22376. [CrossRef] [PubMed]
57. Holden, H.M.; Jacobson, B.L.; Hurley, J.K.; Tollin, G.; Oh, B.-H.; Skjeldal, L.; Chae, Y.K.; Cheng, H.; Xia, B.; Markley, J.L. Structure-function studies of [2Fe-2S] ferredoxins. J. Bioenerg. Biomembr. 1994, 26, 67–88. [CrossRef] [PubMed]
58. Golding, A.J.; Johnson, G.N. Down-regulation of linear and activation of cyclic electron transport during drought. Planta 2003, 218, 107–114. [CrossRef] [PubMed]
59. Harbinson, J.; Genty, B.; Foyer, C.H. Relationship between photosynthetic electron transport and stromal enzyme activity in pea leaves: Toward an understanding of the nature of photosynthetic control. Plant Physiol. 1990, 94, 545–553. [CrossRef]
60. Harbinson, J.; Foyer, C.H. Relationships between the efficiencies of photosystems I and II and stromal redox state in CO2 -free air: Evidence for cyclic electron flow in vivo. Plant Physiol. 1991, 97, 41–49. [CrossRef]
61. Harbinson, J.; Hedley, C.L. Changes in P-700 oxidation during the early stages of the induction of photosynthesis. Plant Physiol. 1993, 103, 649–660. [CrossRef]
62. Kono, M.; Noguchi, K.; Terashima, I. Roles of the cyclic electron flow around PSI (CEF-PSI) and O2 -dependent alternative pathways in regulation of the photosynthetic electron flow in short-term fluctuating light in Arabidopsis thaliana. Plant Cell Physiol. 2014, 55, 990–1004. [CrossRef] [PubMed]
63. Miyake, C.; Miyata, M.; Shinzaki, Y.; Tomizawa, K.-I. CO2 response of cyclic electron flow around PSI (CEF-PSI) in Tobacco leaves—Relative electron fluxes through PSI and PSII determine the magnitude of non-photochemical quenching (NPQ) of Chl fluorescence. Plant Cell Physiol. 2005, 46, 629–637. [CrossRef] [PubMed]
64. Wada, S.; Miyake, C.; Makino, A.; Suzuki, Y. Photorespiration coupled with CO2 assimilation protects photosystem I from photoinhibition under moderate poly(ethylene glycol)-induced osmotic stress in Rice. Front. Plant Sci. 2020, 11, 1121. [CrossRef] [PubMed]
65. Wada, S.; Takagi, D.; Miyake, C.; Makino, A.; Suzuki, Y. Responses of the photosynthetic electron transport reactions stimulate the oxidation of the reaction center chlorophyll of photosystem I, P700, under drought and high temperatures in Rice. Int. J. Mol. Sci. 2019, 20, 2068. [CrossRef] [PubMed]
66. Shi, Q.; Wang, X.; Zeng, Z.; Huang, W. Photoinhibition of photosystem I induced by different intensities of fluctuating light is determined by the kinetics of ∆pH formation rather than linear electron flow. Antioxidants 2022, 11, 2325. [CrossRef]
67. Tikhonov, A.N. The cytochrome b6f complex at the crossroad of photosynthetic electron transport pathways. Plant Physiol. Biochem. 2014, 81, 163–183. [CrossRef]