{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,29]],"date-time":"2026-10-29T15:31:42Z","timestamp":1793287902034,"version":"4.5.0"},"reference-count":106,"publisher":"Springer Science and Business Media LLC","issue":"5","license":[{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T00:00:00Z","timestamp":1791936000000},"content-version":"vor","delay-in-days":13,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Photosynth Res"],"published-print":{"date-parts":[[2026,10]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Thylakoid membranes are highly organized yet dynamic systems in which Photosystem II (PSII) complexes undergo continual assembly, repair and lateral redistribution. Here a hypothesis is proposed that local lumenal pH, generated by the trans-thylakoidal proton gradient and potentially exhibiting transient spatial heterogeneity, may contribute to the regulation of protein movement and PSII maturation within the thylakoid membrane. Five mechanistically distinct but potentially coupled routes by which \u0394pH could influence protein dynamics are outlined: (i) pH-dependent modulation of lumenal protein-protein affinities; (ii) proton-sensitivity of lumenal proteases (e.g., CtpA) that gate processing and release of PSII subunits; (iii) proton-induced changes in membrane packing and stacking that alter diffusion pathways; (iv) formation of local proton microdomains that create chemically distinct niches; and (v) coupling of proton flux to electrostatic and lipid compositional shifts that bias protein localization. Together, these considerations suggest that \u0394pH may act as a co-regulatory parameter integrating biochemical and biophysical cues in thylakoid organization and PSII homeostasis.<\/jats:p>","DOI":"10.1007\/s11120-026-01240-2","type":"journal-article","created":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T09:02:29Z","timestamp":1791968549000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["From proton microdomains to protein migration: hypothesis for \u0394pH-controlled PSII relocation"],"prefix":"10.1007","volume":"164","author":[{"given":"Matthias","family":"Ostermeier","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,14]]},"reference":[{"issue":"1","key":"1240_CR1","doi-asserted-by":"publisher","first-page":"10067","DOI":"10.1038\/s41598-017-10700-8","volume":"7","author":"P Albanese","year":"2017","unstructured":"Albanese P, Melero R, Engel BD, Grinzato A, Berto P, Manfredi M, Chiodoni A, Vargas J, Sorzano C\u00d3S, Marengo E (2017) Pea PSII-LHCII supercomplexes form pairs by making connections across the stromal gap. Sci Rep 7(1):10067","journal-title":"Sci Rep"},{"issue":"2","key":"1240_CR2","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1016\/S0092-8674(03)00032-1","volume":"112","author":"NN Alder","year":"2003","unstructured":"Alder NN, Theg SM (2003) Energetics of protein transport across biological membranes: a study of the thylakoid \u2206pH-dependent\/cpTat pathway. Cell 112(2):231\u2013242","journal-title":"Cell"},{"issue":"17","key":"1240_CR3","doi-asserted-by":"crossref","first-page":"8082","DOI":"10.1073\/pnas.91.17.8082","volume":"91","author":"P Anbudurai","year":"1994","unstructured":"Anbudurai P, Mor TS, Ohad I, Shestakov SV, Pakrasi HB (1994) The ctpA gene encodes the C-terminal processing protease for the D1 protein of the photosystem II reaction center complex. Proc Natl Acad Sci 91(17):8082\u20138086","journal-title":"Proc Natl Acad Sci"},{"issue":"2","key":"1240_CR4","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1111\/j.1399-3054.1989.tb05640.x","volume":"76","author":"JM Anderson","year":"1989","unstructured":"Anderson JM (1989) The grana margins of plant thylakoid membranes. Physiol Plant 76(2):243\u2013248","journal-title":"Physiol Plant"},{"issue":"1608","key":"1240_CR5","doi-asserted-by":"crossref","first-page":"3384","DOI":"10.1098\/rstb.2012.0060","volume":"367","author":"JM Anderson","year":"2012","unstructured":"Anderson JM (2012) Lateral heterogeneity of plant thylakoid protein complexes: early reminiscences. Philosophical Trans Royal Soc B: Biol Sci 367(1608):3384\u20133388","journal-title":"Philosophical Trans Royal Soc B: Biol Sci"},{"issue":"2","key":"1240_CR6","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1016\/0005-2728(80)90078-X","volume":"593","author":"B Andersson","year":"1980","unstructured":"Andersson B, Anderson JM (1980) Lateral heterogeneity in the distribution of chlorophyll-protein complexes of the thylakoid membranes of spinach chloroplasts. Biochim et Biophys Acta (BBA)-Bioenergetics 593(2):427\u2013440","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"key":"1240_CR7","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.pbi.2017.03.012","volume":"37","author":"U Armbruster","year":"2017","unstructured":"Armbruster U, Galvis VC, Kunz H-H, Strand DD (2017) The regulation of the chloroplast proton motive force plays a key role for photosynthesis in fluctuating light. Curr Opin Plant Biol 37:56\u201362","journal-title":"Curr Opin Plant Biol"},{"issue":"14","key":"1240_CR8","doi-asserted-by":"crossref","first-page":"7334","DOI":"10.1074\/jbc.M115.704601","volume":"291","author":"M Ballottari","year":"2016","unstructured":"Ballottari M, Truong TB, De Re E, Erickson E, Stella GR, Fleming GR, Bassi R, Niyogi KK (2016) Identification of pH-sensing sites in the light harvesting complex stress-related 3 protein essential for triggering non-photochemical quenching in Chlamydomonas reinhardtii. J Biol Chem 291(14):7334\u20137346","journal-title":"J Biol Chem"},{"issue":"22","key":"1240_CR9","doi-asserted-by":"crossref","first-page":"15255","DOI":"10.1074\/jbc.M808625200","volume":"284","author":"N Betterle","year":"2009","unstructured":"Betterle N, Ballottari M, Zorzan S, De Bianchi S, Cazzaniga S, Dall\u2019Osto L, Morosinotto T, Bassi R (2009) Light-induced dissociation of an antenna hetero-oligomer is needed for non-photochemical quenching induction. J Biol Chem 284(22):15255\u201315266","journal-title":"J Biol Chem"},{"issue":"12","key":"1240_CR10","doi-asserted-by":"crossref","first-page":"184037","DOI":"10.1016\/j.bbamem.2022.184037","volume":"1864","author":"S Bolik","year":"2022","unstructured":"Bolik S, Albrieux C, Schneck E, Dem\u00e9 B, Jouhet J (2022) Sulfoquinovosyldiacylglycerol and phosphatidylglycerol bilayers share biophysical properties and are good mutual substitutes in photosynthetic membranes. Biochim et Biophys Acta (BBA)-Biomembranes 1864(12):184037","journal-title":"Biochim et Biophys Acta (BBA)-Biomembranes"},{"issue":"12","key":"1240_CR11","doi-asserted-by":"crossref","first-page":"184052","DOI":"10.1016\/j.bbamem.2022.184052","volume":"1864","author":"A-N Bondar","year":"2022","unstructured":"Bondar A-N (2022) Interplay between local protein interactions and water bridging of a proton antenna carboxylate cluster. Biochim et Biophys Acta (BBA)-Biomembranes 1864(12):184052","journal-title":"Biochim et Biophys Acta (BBA)-Biomembranes"},{"issue":"1","key":"1240_CR12","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1093\/plphys\/kiad526","volume":"194","author":"PR Bos","year":"2024","unstructured":"Bos PR, Berentsen J, Wientjes E (2024) Expansion microscopy resolves the thylakoid structure of spinach. Plant Physiol 194(1):347\u2013358","journal-title":"Plant Physiol"},{"issue":"49","key":"1240_CR13","doi-asserted-by":"crossref","first-page":"9782","DOI":"10.1021\/bi301332v","volume":"51","author":"MF Brown","year":"2012","unstructured":"Brown MF (2012) Curvature forces in membrane lipid\u2013protein interactions. Biochemistry 51(49):9782\u20139795","journal-title":"Biochemistry"},{"key":"1240_CR14","doi-asserted-by":"crossref","first-page":"676036","DOI":"10.3389\/fpls.2021.676036","volume":"12","author":"W Chang","year":"2021","unstructured":"Chang W, Li C, Cui Z, Li W, Song H, Chang H, Fu W, Wang C, Huang T, Luo Y (2021) Diverged early from CtpB and CtpC, CtpA has evolved to process D1 precursor in oxygenic photosynthetic organisms. Front Plant Sci 12:676036","journal-title":"Front Plant Sci"},{"issue":"4","key":"1240_CR15","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1021\/acs.jpclett.2c03760","volume":"14","author":"MG Chiariello","year":"2023","unstructured":"Chiariello MG, Gr\u00fcnewald F, Zarmiento-Garcia R, Marrink SJ (2023) pH-dependent conformational switch impacts stability of the PsbS dimer. J Phys Chem Lett 14(4):905\u2013911","journal-title":"J Phys Chem Lett"},{"issue":"11","key":"1240_CR16","doi-asserted-by":"crossref","first-page":"e70008","DOI":"10.1002\/pld3.70008","volume":"8","author":"P Chotewutmontri","year":"2024","unstructured":"Chotewutmontri P, Barkan A (2024) Localization of proteins involved in the biogenesis and repair of the photosynthetic apparatus to thylakoid subdomains in Arabidopsis. Plant Direct 8(11):e70008","journal-title":"Plant Direct"},{"issue":"1\u20132","key":"1240_CR17","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.bbabio.2004.09.009","volume":"1706","author":"JP Dekker","year":"2005","unstructured":"Dekker JP, Boekema EJ (2005) Supramolecular organization of thylakoid membrane proteins in green plants. Biochim et Biophys Acta (BBA)-Bioenergetics 1706(1\u20132):12\u201339","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"issue":"8","key":"1240_CR18","doi-asserted-by":"crossref","first-page":"3373","DOI":"10.1096\/fj.13-247395","volume":"28","author":"B Dem\u00e9","year":"2014","unstructured":"Dem\u00e9 B, Cataye C, Block MA, Mar\u00e9chal E, Jouhet J (2014) Contribution of galactoglycerolipids to the 3-dimensional architecture of thylakoids. FASEB J 28(8):3373\u20133383","journal-title":"FASEB J"},{"issue":"1","key":"1240_CR19","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1146\/annurev.pp.38.060187.002023","volume":"38","author":"RA Dilley","year":"1987","unstructured":"Dilley RA, Theg SM, Beard WA (1987) Membrane-proton interactions in chloroplast bioenergetics: Localized proton domains. Annu Rev Plant Physiol 38(1):347\u2013389","journal-title":"Annu Rev Plant Physiol"},{"key":"1240_CR20","doi-asserted-by":"crossref","first-page":"e04889","DOI":"10.7554\/eLife.04889","volume":"4","author":"BD Engel","year":"2015","unstructured":"Engel BD, Schaffer M, Cuellar LK, Villa E, Plitzko JM, Baumeister W (2015) Native architecture of the Chlamydomonas chloroplast revealed by in situ cryo-electron tomography. elife 4:e04889","journal-title":"elife"},{"issue":"23","key":"1240_CR21","doi-asserted-by":"crossref","first-page":"2898","DOI":"10.1039\/C7CC08740C","volume":"54","author":"Y Ermakova","year":"2018","unstructured":"Ermakova Y, Pak V, Bogdanova Y, Kotlobay A, Yampolsky I, Shokhina A, Panova A, Marygin R, Staroverov D, Bilan D (2018) SypHer3s: a genetically encoded fluorescent ratiometric probe with enhanced brightness and an improved dynamic range. Chem Commun 54(23):2898\u20132901","journal-title":"Chem Commun"},{"key":"1240_CR22","doi-asserted-by":"crossref","first-page":"1310167","DOI":"10.3389\/fphgy.2023.1310167","volume":"1","author":"D Farci","year":"2023","unstructured":"Farci D, Schr\u00f6der WP (2023) Thylakoid Lumen; from proton bag to photosynthetic functionally important compartment. Front Plant Physiol 1:1310167","journal-title":"Front Plant Physiol"},{"key":"1240_CR23","doi-asserted-by":"publisher","unstructured":"Fujii S, Nagata N, Kobayashi K (2025) Membrane lipid-driven thylakoid biogenesis coordinating chlorophyll synthesis and expression of photosynthetic proteins. Plant and Cell Physiology, Volume 66, Issue 11, November 2025, Pages 1645\u20131665, https:\/\/doi.org\/10.1093\/pcp\/pcaf130","DOI":"10.1093\/pcp\/pcaf130"},{"issue":"1","key":"1240_CR24","doi-asserted-by":"crossref","first-page":"13343","DOI":"10.1038\/s41598-017-13574-y","volume":"7","author":"G Garab","year":"2017","unstructured":"Garab G, Ughy B, Waard Pd, Akhtar P, Javornik U, Kotakis C, \u0160ket P, Karlick\u00fd V, Materov\u00e1 Z, \u0160punda V (2017) Lipid polymorphism in chloroplast thylakoid membranes\u2013as revealed by 31P-NMR and time-resolved merocyanine fluorescence spectroscopy. Sci Rep 7(1):13343","journal-title":"Sci Rep"},{"issue":"11","key":"1240_CR25","doi-asserted-by":"crossref","first-page":"1597","DOI":"10.1002\/cbic.201900739","volume":"21","author":"L Gerland","year":"2020","unstructured":"Gerland L, Friedrich D, Hopf L, Donovan EJ, Wallmann A, Erdmann N, Diehl A, Bommer M, Buzar K, Ibrahim M (2020) pH-Dependent Protonation of Surface Carboxylate Groups in PsbO Enables Local Buffering and Triggers Structural Changes. ChemBioChem 21(11):1597\u20131604","journal-title":"ChemBioChem"},{"issue":"1","key":"1240_CR26","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/0005-2728(83)90174-3","volume":"722","author":"YQ Hong","year":"1983","unstructured":"Hong YQ, Junge W (1983) Localized or delocalized protons in photophosphorylation? On the accessibility of the thylakoid lumen for ions and buffers. Biochim et Biophys Acta (BBA)-Bioenergetics 722(1):197\u2013208","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"issue":"5","key":"1240_CR27","doi-asserted-by":"crossref","first-page":"e2018053118","DOI":"10.1073\/pnas.2018053118","volume":"118","author":"G Huang","year":"2021","unstructured":"Huang G, Xiao Y, Pi X, Zhao L, Zhu Q, Wang W, Kuang T, Han G, Sui S-F, Shen J-R (2021) Structural insights into a dimeric Psb27-photosystem II complex from a cyanobacterium Thermosynechococcus vulcanus. Proc Natl Acad Sci 118(5):e2018053118","journal-title":"Proc Natl Acad Sci"},{"issue":"1","key":"1240_CR28","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41467-021-23680-1","volume":"12","author":"T Huokko","year":"2021","unstructured":"Huokko T, Ni T, Dykes GF, Simpson DM, Brownridge P, Conradi FD, Beynon RJ, Nixon PJ, Mullineaux CW, Zhang P (2021) Probing the biogenesis pathway and dynamics of thylakoid membranes. Nat Commun 12(1):1\u201314","journal-title":"Nat Commun"},{"issue":"32","key":"1240_CR29","doi-asserted-by":"crossref","first-page":"30099","DOI":"10.1074\/jbc.M102600200","volume":"276","author":"N Inagaki","year":"2001","unstructured":"Inagaki N, Maitra R, Satoh K, Pakrasi HB (2001) Amino acid residues that are critical for in vivoCatalytic activity of CtpA, the Carboxyl-terminal processing protease for the D1 protein of photosystem II. J Biol Chem 276(32):30099\u201330105","journal-title":"J Biol Chem"},{"key":"1240_CR30","doi-asserted-by":"crossref","first-page":"434","DOI":"10.3389\/fpls.2013.00434","volume":"4","author":"S J\u00e4rvi","year":"2013","unstructured":"J\u00e4rvi S, Gollan PJ, Aro E-M (2013) Understanding the roles of the thylakoid lumen in photosynthesis regulation. Front Plant Sci 4:434","journal-title":"Front Plant Sci"},{"issue":"4","key":"1240_CR31","doi-asserted-by":"crossref","first-page":"148039","DOI":"10.1016\/j.bbabio.2019.06.011","volume":"1861","author":"MP Johnson","year":"2020","unstructured":"Johnson MP, Wientjes E (2020) The relevance of dynamic thylakoid organisation to photosynthetic regulation. Biochim et Biophys Acta (BBA)-Bioenergetics 1861(4):148039","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"key":"1240_CR32","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1111\/j.1749-6632.1989.tb25164.x","volume":"574","author":"W Junge","year":"1989","unstructured":"Junge W (1989) Protons, the thylakoid membrane, and the chloroplast ATP synthase. Ann N Y Acad Sci 574:268\u2013286","journal-title":"Ann N Y Acad Sci"},{"key":"1240_CR33","doi-asserted-by":"crossref","first-page":"e1","DOI":"10.1017\/S0033583518000112","volume":"52","author":"W Junge","year":"2019","unstructured":"Junge W (2019) Oxygenic photosynthesis: history, status and perspective. Q Rev Biophys 52:e1","journal-title":"Q Rev Biophys"},{"issue":"3","key":"1240_CR35","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1016\/0005-2728(86)90200-8","volume":"848","author":"W Junge","year":"1986","unstructured":"Junge W, Polle A (1986) Theory of proton flow along appressed thylakoid membranes under both non-stationary and stationary conditions. Biochim et Biophys Acta (BBA)-Bioenergetics 848(3):265\u2013273","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"issue":"1","key":"1240_CR34","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/0005-2728(79)90175-0","volume":"546","author":"W Junge","year":"1979","unstructured":"Junge W, Ausl\u00e4nder W, McGeer AJ, Runge T (1979) The buffering capacity of the internal phase of thylakoids and the magnitude of the pH changes inside under flashing light. Biochim et Biophys Acta (BBA)-Bioenergetics 546(1):121\u2013141","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"issue":"4","key":"1240_CR36","doi-asserted-by":"crossref","first-page":"483","DOI":"10.32615\/ps.2023.043","volume":"61","author":"R Ka\u0148a","year":"2023","unstructured":"Ka\u0148a R, \u0160ediv\u00e1 B, Pr\u00e1\u0161il O (2023) Microdomains heterogeneity in the thylakoid membrane proteins visualized by super-resolution microscopy. Photosynthetica 61(4):483","journal-title":"Photosynthetica"},{"issue":"4","key":"1240_CR37","doi-asserted-by":"crossref","first-page":"149504","DOI":"10.1016\/j.bbabio.2024.149504","volume":"1865","author":"DB Kell","year":"2024","unstructured":"Kell DB (2024) A protet-based model that can account for energy coupling in oxidative and photosynthetic phosphorylation. Biochim et Biophys Acta (BBA)-Bioenergetics 1865(4):149504","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"issue":"2","key":"1240_CR38","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1007\/s11120-007-9201-1","volume":"94","author":"J Kern","year":"2007","unstructured":"Kern J, Renger G (2007) Photosystem II: Structure and mechanism of the water: plastoquinone oxidoreductase. Photosynth Res 94(2):183\u2013202","journal-title":"Photosynth Res"},{"issue":"4","key":"1240_CR39","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1016\/j.bbabio.2013.11.003","volume":"1837","author":"H Kirchhoff","year":"2014","unstructured":"Kirchhoff H (2014a) Diffusion of molecules and macromolecules in thylakoid membranes. Biochim et Biophys Acta (BBA)-Bioenergetics 1837(4):495\u2013502","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"issue":"1640","key":"1240_CR40","doi-asserted-by":"crossref","first-page":"20130225","DOI":"10.1098\/rstb.2013.0225","volume":"369","author":"H Kirchhoff","year":"2014","unstructured":"Kirchhoff H (2014b) Structural changes of the thylakoid membrane network induced by high light stress in plant chloroplasts. Philosophical Trans Royal Soc B: Biol Sci 369(1640):20130225","journal-title":"Philosophical Trans Royal Soc B: Biol Sci"},{"issue":"9","key":"1240_CR41","doi-asserted-by":"crossref","first-page":"1180","DOI":"10.1016\/j.bbabio.2007.05.009","volume":"1767","author":"H Kirchhoff","year":"2007","unstructured":"Kirchhoff H, Haase W, Haferkamp S, Schott T, Borinski M, Kubitscheck U, R\u00f6gner M (2007) Structural and functional self-organization of Photosystem II in grana thylakoids. Biochim et Biophys Acta (BBA)-Bioenergetics 1767(9):1180\u20131188","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"issue":"4","key":"1240_CR42","doi-asserted-by":"publisher","first-page":"1571","DOI":"10.1104\/pp.107.115170","volume":"146","author":"H Kirchhoff","year":"2008","unstructured":"Kirchhoff H, Haferkamp S, Allen JF, Epstein DB, Mullineaux CW (2008) Protein diffusion and macromolecular crowding in thylakoid membranes. Plant Physiol 146(4):1571\u20131578. https:\/\/doi.org\/10.1104\/pp.107.115170","journal-title":"Plant Physiol"},{"key":"1240_CR43","doi-asserted-by":"publisher","unstructured":"Komenda J, Sobotka R, Nixon PJ (2024) The biogenesis and maintenance of PSII: recent advances and current challenges. The Plant Cell, Volume 36, Issue 10, October 2024, Pages 3997\u20134013, https:\/\/doi.org\/10.1093\/plcell\/koae082","DOI":"10.1093\/plcell\/koae082"},{"issue":"3","key":"1240_CR44","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1034\/j.1600-0854.2003.00086.x","volume":"4","author":"EE Kooijman","year":"2003","unstructured":"Kooijman EE, Chupin V, de Kruijff B, Burger KN (2003) Modulation of membrane curvature by phosphatidic acid and lysophosphatidic acid. Traffic 4(3):162\u2013174","journal-title":"Traffic"},{"issue":"1","key":"1240_CR45","doi-asserted-by":"publisher","first-page":"2291","DOI":"10.1038\/s41467-021-22530-4","volume":"12","author":"M Krishnan-Schmieden","year":"2021","unstructured":"Krishnan-Schmieden M, Konold PE, Kennis JT, Pandit A (2021) The molecular pH-response mechanism of the plant light-stress sensor PsbS. Nat Commun 12(1):2291","journal-title":"Nat Commun"},{"key":"1240_CR46","doi-asserted-by":"publisher","unstructured":"Krysiak M, Oung HMO, Kirchhoff H (2025) What are grana in chloroplasts of vascular plants good for? Annals of Botany, Volume 137, Issue 3, March 2026, Pages 571\u2013590, https:\/\/doi.org\/10.1093\/aob\/mcaf229","DOI":"10.1093\/aob\/mcaf229"},{"issue":"7","key":"1240_CR48","doi-asserted-by":"publisher","first-page":"1047","DOI":"10.1038\/nn.3126","volume":"15","author":"Y Li","year":"2012","unstructured":"Li Y, Tsien RW (2012) pHTomato, a red, genetically encoded indicator that enables multiplex interrogation of synaptic activity. Nat Neurosci 15(7):1047\u20131053","journal-title":"Nat Neurosci"},{"issue":"11","key":"1240_CR47","doi-asserted-by":"crossref","first-page":"e00280","DOI":"10.1002\/pld3.280","volume":"4","author":"M Li","year":"2020","unstructured":"Li M, Mukhopadhyay R, Svoboda V, Oung HMO, Mullendore DL, Kirchhoff H (2020) Measuring the dynamic response of the thylakoid architecture in plant leaves by electron microscopy. Plant Direct 4(11):e00280","journal-title":"Plant Direct"},{"issue":"8","key":"1240_CR49","doi-asserted-by":"crossref","first-page":"1737","DOI":"10.1021\/acs.jpclett.9b00437","volume":"10","author":"N Liguori","year":"2019","unstructured":"Liguori N, Campos SR, Baptista AnM, Croce R (2019) Molecular anatomy of plant photoprotective switches: the sensitivity of PsbS to the environment, residue by residue. J Phys Chem Lett 10(8):1737\u20131742","journal-title":"J Phys Chem Lett"},{"key":"1240_CR50","doi-asserted-by":"crossref","unstructured":"L\u00fcbben MK, Klingl A, Nickelsen J, Ostermeier M (2024) CLEM, a universal tool for analyzing structural organization in thylakoid membranes. Physiol Plant 176(4):e14417","DOI":"10.1111\/ppl.14417"},{"issue":"1","key":"1240_CR51","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1007\/s11120-019-00648-3","volume":"142","author":"W Lubitz","year":"2019","unstructured":"Lubitz W, Chrysina M, Cox N (2019) Water oxidation in photosystem II. Photosynth Res 142(1):105\u2013125","journal-title":"Photosynth Res"},{"key":"1240_CR52","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1016\/j.plaphy.2014.02.013","volume":"81","author":"PD Mabbitt","year":"2014","unstructured":"Mabbitt PD, Wilbanks SM, Eaton-Rye JJ (2014) Structure and function of the hydrophilic Photosystem II assembly proteins: Psb27, Psb28 and Ycf48. Plant Physiol Biochem 81:96\u2013107","journal-title":"Plant Physiol Biochem"},{"issue":"5","key":"1240_CR53","doi-asserted-by":"crossref","first-page":"1119","DOI":"10.1105\/tpc.17.00071","volume":"29","author":"C MacGregor-Chatwin","year":"2017","unstructured":"MacGregor-Chatwin C, Sener M, Barnett SF, Hitchcock A, Barnhart-Dailey MC, Maghlaoui K, Barber J, Timlin JA, Schulten K, Hunter CN (2017) Lateral segregation of photosystem I in cyanobacterial thylakoids. Plant Cell 29(5):1119\u20131136","journal-title":"Plant Cell"},{"issue":"5","key":"1240_CR54","doi-asserted-by":"crossref","first-page":"1589","DOI":"10.1105\/tpc.19.00797","volume":"32","author":"LJ McKinnon","year":"2020","unstructured":"McKinnon LJ, Fukushima J, Endow JK, Inoue K, Theg SM (2020) Membrane chaperoning of a thylakoid protease whose structural stability is modified by the protonmotive force. Plant Cell 32(5):1589\u20131609","journal-title":"Plant Cell"},{"issue":"6","key":"1240_CR55","doi-asserted-by":"crossref","first-page":"811","DOI":"10.3390\/plants13060811","volume":"13","author":"HS Mehra","year":"2024","unstructured":"Mehra HS, Wang X, Russell BP, Kulkarni N, Ferrari N, Larson B, Vinyard DJ (2024) Assembly and Repair of Photosystem II in Chlamydomonas reinhardtii. Plants 13(6):811","journal-title":"Plants"},{"issue":"400","key":"1240_CR57","doi-asserted-by":"crossref","first-page":"1207","DOI":"10.1093\/jxb\/erh106","volume":"55","author":"CW Mullineaux","year":"2004","unstructured":"Mullineaux CW (2004) FRAP analysis of photosynthetic membranes. J Exp Bot 55(400):1207\u20131211","journal-title":"J Exp Bot"},{"issue":"6","key":"1240_CR58","doi-asserted-by":"crossref","first-page":"1310","DOI":"10.1111\/j.1751-1097.2008.00420.x","volume":"84","author":"CW Mullineaux","year":"2008","unstructured":"Mullineaux CW (2008) Factors controlling the mobility of photosynthetic proteins. Photochem Photobiol 84(6):1310\u20131316","journal-title":"Photochem Photobiol"},{"key":"1240_CR56","doi-asserted-by":"publisher","unstructured":"Mullineaux C, Mahbub M (2023) Locations of membrane protein production in a cyanobacterium. J Bacteriol 205:e00209-23. https:\/\/doi.org\/10.1128\/jb.00209-23","DOI":"10.1128\/jb.00209-23"},{"issue":"21","key":"1240_CR59","doi-asserted-by":"crossref","first-page":"3372","DOI":"10.1016\/j.febslet.2013.09.015","volume":"587","author":"K Nath","year":"2013","unstructured":"Nath K, Jajoo A, Poudyal RS, Timilsina R, Park YS, Aro E-M, Nam HG, Lee C-H (2013) Towards a critical understanding of the photosystem II repair mechanism and its regulation during stress conditions. FEBS Lett 587(21):3372\u20133381","journal-title":"FEBS Lett"},{"issue":"1","key":"1240_CR60","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1007\/s11120-010-9576-2","volume":"106","author":"JA Neilson","year":"2010","unstructured":"Neilson JA, Durnford DG (2010) Structural and functional diversification of the light-harvesting complexes in photosynthetic eukaryotes. Photosynth Res 106(1):57\u201371","journal-title":"Photosynth Res"},{"key":"1240_CR61","doi-asserted-by":"publisher","first-page":"609","DOI":"10.1146\/annurev-arplant-050312-120124","volume":"64","author":"J Nickelsen","year":"2013","unstructured":"Nickelsen J, Rengstl B (2013) Photosystem II assembly: from cyanobacteria to plants. Annu Rev Plant Biol 64:609\u2013635. https:\/\/doi.org\/10.1146\/annurev-arplant-050312-120124","journal-title":"Annu Rev Plant Biol"},{"issue":"1","key":"1240_CR62","doi-asserted-by":"publisher","first-page":"7415","DOI":"10.1038\/s41598-021-86975-9","volume":"11","author":"L Nicol","year":"2021","unstructured":"Nicol L, Croce R (2021) The PsbS protein and low pH are necessary and sufficient to induce quenching in the light-harvesting complex of plants LHCII. Sci Rep 11(1):7415","journal-title":"Sci Rep"},{"issue":"411","key":"1240_CR63","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1093\/jxb\/eri056","volume":"56","author":"KK Niyogi","year":"2005","unstructured":"Niyogi KK, Li X-P, Rosenberg V, Jung H-S (2005) Is PsbS the site of non-photochemical quenching in photosynthesis? J Exp Bot 56(411):375\u2013382","journal-title":"J Exp Bot"},{"issue":"8","key":"1240_CR64","doi-asserted-by":"crossref","first-page":"1876","DOI":"10.1016\/j.bpj.2020.02.029","volume":"118","author":"B Onoa","year":"2020","unstructured":"Onoa B, Fukuda S, Iwai M, Bustamante C, Niyogi KK (2020) Atomic force microscopy visualizes mobility of photosynthetic proteins in grana thylakoid membranes. Biophys J 118(8):1876\u20131886","journal-title":"Biophys J"},{"issue":"2","key":"1240_CR65","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1017\/S0033583513000024","volume":"46","author":"AV Onufriev","year":"2013","unstructured":"Onufriev AV, Alexov E (2013) Protonation and pK changes in protein\u2013ligand binding. Q Rev Biophys 46(2):181\u2013209","journal-title":"Q Rev Biophys"},{"issue":"10","key":"1240_CR67","doi-asserted-by":"crossref","first-page":"4014","DOI":"10.1093\/plcell\/koae102","volume":"36","author":"M Ostermeier","year":"2024","unstructured":"Ostermeier M, Garibay-Hern\u00e1ndez A, Holzer VJ, Schroda M, Nickelsen J (2024) Structure, biogenesis, and evolution of thylakoid membranes. Plant Cell 36(10):4014\u20134035","journal-title":"Plant Cell"},{"issue":"2","key":"1240_CR66","doi-asserted-by":"crossref","first-page":"e70234","DOI":"10.1111\/ppl.70234","volume":"177","author":"M Ostermeier","year":"2025","unstructured":"Ostermeier M, Buschmann I, Heinz S, Nickelsen J (2025) The subcellular localisation of early photosystem II assembly in Synechocystis sp. PCC 6803. Physiol Plant 177(2):e70234","journal-title":"Physiol Plant"},{"issue":"44","key":"1240_CR68","doi-asserted-by":"crossref","first-page":"15839","DOI":"10.1073\/pnas.1413739111","volume":"111","author":"S Puthiyaveetil","year":"2014","unstructured":"Puthiyaveetil S, Tsabari O, Lowry T, Lenhert S, Lewis RR, Reich Z, Kirchhoff H (2014) Compartmentalization of the protein repair machinery in photosynthetic membranes. Proc Natl Acad Sci 111(44):15839\u201315844","journal-title":"Proc Natl Acad Sci"},{"issue":"9","key":"1240_CR69","doi-asserted-by":"publisher","first-page":"821","DOI":"10.1016\/j.bbabio.2015.01.007","volume":"1847","author":"A Rast","year":"2015","unstructured":"Rast A, Heinz S, Nickelsen J (2015) Biogenesis of thylakoid membranes. Biochim Biophys Acta 1847(9):821\u2013830. https:\/\/doi.org\/10.1016\/j.bbabio.2015.01.007","journal-title":"Biochim Biophys Acta"},{"issue":"4","key":"1240_CR70","doi-asserted-by":"publisher","first-page":"436","DOI":"10.1038\/s41477-019-0399-7","volume":"5","author":"A Rast","year":"2019","unstructured":"Rast A, Schaffer M, Albert S, Wan W, Pfeffer S, Beck F, Plitzko JM, Nickelsen J, Engel BD (2019) Biogenic regions of cyanobacterial thylakoids form contact sites with the plasma membrane. Nat Plants 5(4):436\u2013446. https:\/\/doi.org\/10.1038\/s41477-019-0399-7","journal-title":"Nat Plants"},{"issue":"24","key":"1240_CR71","doi-asserted-by":"publisher","first-page":"21944","DOI":"10.1074\/jbc.M111.237867","volume":"286","author":"B Rengstl","year":"2011","unstructured":"Rengstl B, Oster U, Stengel A, Nickelsen J (2011) An intermediate membrane subfraction in cyanobacteria is involved in an assembly network for Photosystem II biogenesis. J Biol Chem 286(24):21944\u201321951. https:\/\/doi.org\/10.1074\/jbc.M111.237867","journal-title":"J Biol Chem"},{"issue":"1","key":"1240_CR73","doi-asserted-by":"crossref","first-page":"3103","DOI":"10.1038\/ncomms4103","volume":"5","author":"B Rieger","year":"2014","unstructured":"Rieger B, Junge W, Busch KB (2014) Lateral pH gradient between OXPHOS complex IV and F0F1 ATP-synthase in folded mitochondrial membranes. Nat Commun 5(1):3103","journal-title":"Nat Commun"},{"key":"1240_CR72","first-page":"2011430746","volume":"20212002","author":"B Rieger","year":"2021","unstructured":"Rieger B, Arroum T, Villalta J, Busch KB (2021) pH profiling of mitochondrial sub-compartments reveals that IF1 is essential to prevent ATP hydrolysis under OXPHOS conditions. bioRxiv 20212002:2011430746","journal-title":"bioRxiv"},{"key":"1240_CR74","doi-asserted-by":"crossref","first-page":"1234","DOI":"10.3389\/fpls.2015.01234","volume":"6","author":"T R\u00fchle","year":"2016","unstructured":"R\u00fchle T, Leister D (2016) Photosystem II assembly from scratch. Front Plant Sci 6:1234","journal-title":"Front Plant Sci"},{"key":"1240_CR75","doi-asserted-by":"publisher","unstructured":"Sakamoto W (2025) Thylakostasis: key factors in thylakoid membrane organization with emphasis on biogenesis and remodeling proteins in vascular plants. Plant and Cell Physiology, Volume 66, Issue 11, November 2025, Pages 1602\u20131618, https:\/\/doi.org\/10.1093\/pcp\/pcaf098","DOI":"10.1093\/pcp\/pcaf098"},{"key":"1240_CR76","unstructured":"Sarcina M, Mullineaux C, Murata N (2001) Thylakoid membrane fluidity and its crucial importance in photoinhibition. In: Proceedings of the 12th International Congress on Photosynthesis; 2001; Brisbane, Australia. Collingwood (VIC): CSIRO Publishing 3(1)"},{"issue":"3","key":"1240_CR77","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1038\/s41566-025-01625-0","volume":"19","author":"L Scheiderer","year":"2025","unstructured":"Scheiderer L, Marin Z, Ries J (2025) MINFLUX achieves molecular resolution with minimal photons. Nat Photonics 19(3):238\u2013247","journal-title":"Nat Photonics"},{"issue":"47","key":"1240_CR78","doi-asserted-by":"crossref","first-page":"19286","DOI":"10.1073\/pnas.1209860109","volume":"109","author":"M Schottkowski","year":"2012","unstructured":"Schottkowski M, Peters M, Zhan Y, Rifai O, Zhang Y, Zerges W (2012) Biogenic membranes of the chloroplast in Chlamydomonas reinhardtii. Proc Natl Acad Sci 109(47):19286\u201319291","journal-title":"Proc Natl Acad Sci"},{"issue":"9","key":"1240_CR79","doi-asserted-by":"crossref","first-page":"2373","DOI":"10.1093\/jxb\/eru495","volume":"66","author":"MA Sch\u00f6ttler","year":"2015","unstructured":"Sch\u00f6ttler MA, T\u00f3th SZ, Boulouis A, Kahlau S (2015) Photosynthetic complex stoichiometry dynamics in higher plants: biogenesis, function, and turnover of ATP synthase and the cytochrome b 6 f complex. J Exp Bot 66(9):2373\u20132400","journal-title":"J Exp Bot"},{"key":"1240_CR80","doi-asserted-by":"crossref","unstructured":"Sekiguchi T, Yoshida K, Wakabayashi K-i, Hisabori T (2024) Proton gradient across the chloroplast thylakoid membrane governs the redox regulatory function of ATP synthase. J Biol Chem 300(9)","DOI":"10.1016\/j.jbc.2024.107659"},{"issue":"5","key":"1240_CR81","doi-asserted-by":"publisher","first-page":"782","DOI":"10.1038\/emboj.2008.12","volume":"27","author":"T Shutova","year":"2008","unstructured":"Shutova T, Kenneweg H, Buchta J, Nikitina J, Terentyev V, Chernyshov S, Andersson B, Allakhverdiev SI, Klimov VV, Dau H (2008) The photosystem II-associated Cah3 in Chlamydomonas enhances the O2 evolution rate by proton removal. EMBO J 27(5):782\u2013791","journal-title":"EMBO J"},{"issue":"4023","key":"1240_CR82","doi-asserted-by":"crossref","first-page":"720","DOI":"10.1126\/science.175.4023.720","volume":"175","author":"SJ Singer","year":"1972","unstructured":"Singer SJ, Nicolson GL (1972) The Fluid Mosaic Model of the Structure of Cell Membranes: Cell membranes are viewed as two-dimensional solutions of oriented globular proteins and lipids. Science 175(4023):720\u2013731","journal-title":"Science"},{"issue":"2","key":"1240_CR83","doi-asserted-by":"publisher","first-page":"660","DOI":"10.1105\/tpc.111.093914","volume":"24","author":"A Stengel","year":"2012","unstructured":"Stengel A, Gugel IL, Hilger D, Rengstl B, Jung H, Nickelsen J (2012) Initial steps of photosystem II de novo assembly and preloading with manganese take place in biogenesis centers in Synechocystis. Plant Cell 24(2):660\u2013675. https:\/\/doi.org\/10.1105\/tpc.111.093914","journal-title":"Plant Cell"},{"issue":"2","key":"1240_CR86","doi-asserted-by":"crossref","first-page":"294","DOI":"10.1016\/0005-2728(83)90129-9","volume":"723","author":"SM Theg","year":"1983","unstructured":"Theg SM, Junge W (1983) The effect of low concentrations of uncouplers on the detectability of proton deposition in thylakoids. Evidence for subcompartmentation and preexisting pH differences in the dark. Biochim et Biophys Acta (BBA)-Bioenergetics 723(2):294\u2013307","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"key":"1240_CR87","doi-asserted-by":"publisher","unstructured":"Theg SM, Tom C (2011) Measurement of the \u0394pH and electric field developed across Arabidopsis thylakoids in the light. In: Jarvis, R. (eds) Chloroplast Research in Arabidopsis. Methods in Molecular Biology, vol 775. Humana Press, Totowa, NJ. https:\/\/doi.org\/10.1007\/978-1-61779-237-3_18","DOI":"10.1007\/978-1-61779-237-3_18"},{"issue":"1","key":"1240_CR85","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/0014-5793(82)81199-X","volume":"145","author":"SM Theg","year":"1982","unstructured":"Theg SM, Johnson JD, Homann PH (1982) Proton efflux from thylakoids induced in darkness and its effect on photosystem II. FEBS Lett 145(1):25\u201329","journal-title":"FEBS Lett"},{"issue":"2","key":"1240_CR84","doi-asserted-by":"crossref","first-page":"673","DOI":"10.1016\/S0021-9258(19)35405-5","volume":"263","author":"S Theg","year":"1988","unstructured":"Theg S, Chiang G, Dilley R (1988) Protons in the thylakoid membrane-sequestered domains can directly pass through the coupling factor during ATP synthesis in flashing light. J Biol Chem 263(2):673\u2013681","journal-title":"J Biol Chem"},{"issue":"9","key":"1240_CR88","doi-asserted-by":"crossref","first-page":"e1218587","DOI":"10.1080\/15592324.2016.1218587","volume":"11","author":"J Theis","year":"2016","unstructured":"Theis J, Schroda M (2016) Revisiting the photosystem II repair cycle. Plant Signal Behav 11(9):e1218587","journal-title":"Plant Signal Behav"},{"issue":"8","key":"1240_CR89","doi-asserted-by":"crossref","first-page":"2147","DOI":"10.1074\/mcp.M114.040923","volume":"13","author":"M Tomizioli","year":"2014","unstructured":"Tomizioli M, Lazar C, Brugi\u00e8re S, Burger T, Salvi D, Gatto L, Moyet L, Breckels LM, Hesse A-M, Lilley KS (2014) Deciphering thylakoid sub-compartments using a mass spectrometry-based approach. Mol Cell Proteom 13(8):2147\u20132167","journal-title":"Mol Cell Proteom"},{"issue":"1","key":"1240_CR90","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1016\/S0005-2728(00)00069-4","volume":"1458","author":"C Tommos","year":"2000","unstructured":"Tommos C, Babcock GT (2000) Proton and hydrogen currents in photosynthetic water oxidation. Biochim et Biophys Acta (BBA)-Bioenergetics 1458(1):199\u2013219","journal-title":"Biochim et Biophys Acta (BBA)-Bioenergetics"},{"key":"1240_CR91","doi-asserted-by":"crossref","first-page":"919896","DOI":"10.3389\/fpls.2022.919896","volume":"13","author":"MDL Trinh","year":"2022","unstructured":"Trinh MDL, Masuda S (2022) Chloroplast pH homeostasis for the regulation of photosynthesis. Front Plant Sci 13:919896","journal-title":"Front Plant Sci"},{"issue":"11","key":"1240_CR92","doi-asserted-by":"publisher","first-page":"3640","DOI":"10.1105\/tpc.107.054882","volume":"19","author":"J Uniacke","year":"2007","unstructured":"Uniacke J, Zerges W (2007) Photosystem II assembly and repair are differentially localized in Chlamydomonas. Plant Cell 19(11):3640\u20133654. https:\/\/doi.org\/10.1105\/tpc.107.054882","journal-title":"Plant Cell"},{"issue":"9","key":"1240_CR93","doi-asserted-by":"crossref","first-page":"200144","DOI":"10.1098\/rsob.200144","volume":"10","author":"R \u00dcnnep","year":"2020","unstructured":"\u00dcnnep R, Paul S, Zsiros O, Kov\u00e1cs L, Sz\u00e9kely NK, Steinbach G, Appavou M-S, Porcar L, Holzwarth AR, Garab G (2020) Thylakoid membrane reorganizations revealed by small-angle neutron scattering of Monstera deliciosa leaves associated with non-photochemical quenching. Open Biology 10(9):200144","journal-title":"Open Biology"},{"issue":"15","key":"1240_CR94","doi-asserted-by":"crossref","first-page":"3237","DOI":"10.1021\/acs.jpcb.6b06865","volume":"121","author":"FJ Van Eerden","year":"2017","unstructured":"Van Eerden FJ, Van Den Berg T, Frederix PW, De Jong DH, Periole X, Marrink SJ (2017) Molecular dynamics of photosystem II embedded in the thylakoid membrane. J Phys Chem B 121(15):3237\u20133249","journal-title":"J Phys Chem B"},{"issue":"3","key":"1240_CR95","doi-asserted-by":"crossref","first-page":"388","DOI":"10.1016\/j.bbamem.2016.11.016","volume":"1859","author":"AV Vershubskii","year":"2017","unstructured":"Vershubskii AV, Trubitsin BV, Priklonskii VI, Tikhonov AN (2017) Lateral heterogeneity of the proton potential along the thylakoid membranes of chloroplasts. Biochim et Biophys Acta (BBA)-Biomembranes 1859(3):388\u2013401","journal-title":"Biochim et Biophys Acta (BBA)-Biomembranes"},{"issue":"2","key":"1240_CR96","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1007\/s11120-013-9905-3","volume":"116","author":"M Watanabe","year":"2013","unstructured":"Watanabe M, Ikeuchi M (2013) Phycobilisome: architecture of a light-harvesting supercomplex. Photosynth Res 116(2):265\u2013276","journal-title":"Photosynth Res"},{"issue":"28","key":"1240_CR97","doi-asserted-by":"crossref","first-page":"21391","DOI":"10.1074\/jbc.M110.105064","volume":"285","author":"L Wei","year":"2010","unstructured":"Wei L, Guo J, Ouyang M, Sun X, Ma J, Chi W, Lu C, Zhang L (2010) LPA19, a Psb27 homolog in Arabidopsis thaliana, facilitates D1 protein precursor processing during PSII biogenesis. J Biol Chem 285(28):21391\u201321398","journal-title":"J Biol Chem"},{"key":"1240_CR98","doi-asserted-by":"crossref","first-page":"153246","DOI":"10.1016\/j.jplph.2020.153246","volume":"252","author":"C Wilhelm","year":"2020","unstructured":"Wilhelm C, Goss R, Garab G (2020) The fluid-mosaic membrane theory in the context of photosynthetic membranes: Is the thylakoid membrane more like a mixed crystal or like a fluid? J Plant Physiol 252:153246","journal-title":"J Plant Physiol"},{"key":"1240_CR99","doi-asserted-by":"crossref","unstructured":"Wu J, Wu C, Chen S, Huang C, Wen Q, Lin W, Wang C, Han D, Lu D, Xu X (2025) Assembly mechanism of PSII-LHCII array from higher plants. J. Integr. Plant Biol. 67: 3152\u20133166","DOI":"10.1111\/jipb.70045"},{"key":"1240_CR100","doi-asserted-by":"crossref","first-page":"417","DOI":"10.3389\/fpls.2013.00417","volume":"4","author":"T Wunder","year":"2013","unstructured":"Wunder T, Xu W, Liu Q, Wanner G, Leister D, Pribil M (2013) The major thylakoid protein kinases STN7 and STN8 revisited: effects of altered STN8 levels and regulatory specificities of the STN kinases. Front Plant Sci 4:417","journal-title":"Front Plant Sci"},{"issue":"10","key":"1240_CR101","doi-asserted-by":"crossref","first-page":"7518","DOI":"10.1074\/jbc.M008877200","volume":"276","author":"Y Yamamoto","year":"2001","unstructured":"Yamamoto Y, Inagaki N, Satoh K (2001) Overexpression and characterization of carboxyl-terminal processing protease for precursor D1 protein: regulation of enzyme-substrate interaction by molecular environments. J Biol Chem 276(10):7518\u20137525","journal-title":"J Biol Chem"},{"issue":"1","key":"1240_CR102","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.bbrc.2016.12.032","volume":"486","author":"H Yang","year":"2017","unstructured":"Yang H, Pu X, Wang L, Liu L, Theg SM (2017) A new fluorescence-based method to monitor the pH in the thylakoid lumen using GFP variants. Biochem Biophys Res Commun 486(1):1\u20135","journal-title":"Biochem Biophys Res Commun"},{"issue":"9","key":"1240_CR103","doi-asserted-by":"crossref","first-page":"4860","DOI":"10.3390\/ijms22094860","volume":"22","author":"A Yoshihara","year":"2021","unstructured":"Yoshihara A, Nagata N, Wada H, Kobayashi K (2021) Plastid anionic lipids are essential for the development of both photosynthetic and non-photosynthetic organs in Arabidopsis thaliana. Int J Mol Sci 22(9):4860","journal-title":"Int J Mol Sci"},{"key":"1240_CR104","doi-asserted-by":"crossref","unstructured":"Zabret J, Bohn S, Schuller SK, Arnolds O, M\u00f6ller M, Meier-Credo J, Liauw P, Chan A, Tajkhorshid E, Langer JD (2020) How to build a water-splitting machine: structural insights into photosystem II assembly. bioRxiv","DOI":"10.21203\/rs.3.rs-88039\/v1"},{"issue":"4","key":"1240_CR105","doi-asserted-by":"publisher","first-page":"524","DOI":"10.1038\/s41477-021-00895-0","volume":"7","author":"J Zabret","year":"2021","unstructured":"Zabret J, Bohn S, Schuller SK, Arnolds O, M\u00f6ller M, Meier-Credo J, Liauw P, Chan A, Tajkhorshid E, Langer JD, Stoll R, Krieger-Liszkay A, Engel BD, Rudack T, Schuller JM, Nowaczyk MM (2021) Structural insights into photosystem II assembly. Nat plants 7(4):524\u2013538","journal-title":"Nat plants"},{"issue":"9","key":"1240_CR106","doi-asserted-by":"crossref","first-page":"1769","DOI":"10.1105\/tpc.12.9.1769","volume":"12","author":"L Zhang","year":"2000","unstructured":"Zhang L, Paakkarinen V, van Wijk KJ, Aro E-M (2000) Biogenesis of the chloroplast-encoded D1 protein: regulation of translation elongation, insertion, and assembly into photosystem II. Plant Cell 12(9):1769\u20131781","journal-title":"Plant Cell"}],"container-title":["Photosynthesis Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11120-026-01240-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s11120-026-01240-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11120-026-01240-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,29]],"date-time":"2026-10-29T14:38:25Z","timestamp":1793284705000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s11120-026-01240-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10]]},"references-count":106,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2026,10]]}},"alternative-id":["1240"],"URL":"https:\/\/doi.org\/10.1007\/s11120-026-01240-2","relation":{},"ISSN":["0166-8595","1573-5079"],"issn-type":[{"value":"0166-8595","type":"print"},{"value":"1573-5079","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10]]},"assertion":[{"value":"21 August 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 October 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 October 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","label":"Ethical approval","group":{"name":"EthicsHeading","label":"Declaration"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declaration"}}],"article-number":"51"}}