{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T12:16:55Z","timestamp":1779365815856,"version":"3.53.0"},"reference-count":91,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2026,4,17]],"date-time":"2026-04-17T00:00:00Z","timestamp":1776384000000},"content-version":"vor","delay-in-days":1,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100005302","name":"University of Illinois","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100005302","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Realizing Increased Photosynthetic Efficiency","award":["OPP1172157"],"award-info":[{"award-number":["OPP1172157"]}]},{"name":"Gates Foundation, Foundation for Food and Agriculture Research"},{"name":"U.K. Government\u2019s Department for International Development"},{"name":"U.S. Department of Energy, Office of Science, Basic Energy Sciences, Chemical Sciences, Geosciences, and Biosciences Division","award":["449B"],"award-info":[{"award-number":["449B"]}]},{"DOI":"10.13039\/100000011","name":"Howard Hughes Medical Institute","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000011","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,5,6]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>CP26 is a monomeric minor light-harvesting complex of Photosystem II (LHCII) protein located at the interface between LHCII trimers and the PSII core in thylakoid membranes. Previous studies have proposed that CP26 plays a role in non-photochemical quenching (NPQ) in addition to light harvesting. Here, we utilized biophysical and pharmacological approaches to investigate this role using single- and higher-order Arabidopsis (Arabidopsis thaliana) cp26 mutants, examining its relationship to known NPQ regulators (Photosystem II subunit S, PsbS, violaxanthin de-epoxidase, and the pH gradient across the thylakoid membrane). cp26 mutants showed significantly reduced maximum PSII quantum efficiencies (Fv\/Fm) in darkness, indicating a constitutively quenched state, further confirmed by fluorescence lifetime measurements. Destabilized PSII-LHCII supercomplexes observed in native gel electrophoresis and tighter PSII supercomplex packing were potential causes, with no other antenna proteins capable of rescuing this phenotype. In addition, the cp26 mutants exhibited altered NPQ capacity\u2014modest in single mutants but substantial in double mutants\u2014independent of PsbS and violaxanthin de-epoxidase. Together, these results show that CP26 is not involved in qE or qZ but may primarily play an indirect role in apparent NPQ responses via PSII-LHCII supercomplex organization.<\/jats:p>","DOI":"10.1093\/plphys\/kiag207","type":"journal-article","created":{"date-parts":[[2026,4,13]],"date-time":"2026-04-13T11:24:16Z","timestamp":1776079456000},"source":"Crossref","is-referenced-by-count":0,"title":["CP26 is not involved in qE- or qZ-type non-photochemical quenching in Arabidopsis"],"prefix":"10.1093","volume":"201","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6230-9626","authenticated-orcid":false,"given":"Julia","family":"Walter","sequence":"first","affiliation":[{"name":"Department of Plant Sciences, University of Cambridge , CB2 3EA Cambridge ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0642-1485","authenticated-orcid":false,"given":"Dhruv","family":"Patel-Tupper","sequence":"additional","affiliation":[{"name":"Department of Plant and Microbial Biology, University of California , Berkeley, CA 94720 ,","place":["United States"]},{"name":"Howard Hughes Medical Institute, University of California , Berkeley, CA 94720 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7292-9463","authenticated-orcid":false,"given":"Lam","family":"Lam","sequence":"additional","affiliation":[{"name":"Graduate Group in Biophysics, University of California , Berkeley, CA 94720 ,","place":["United States"]},{"name":"Molecular Biophysics and Integrated Bioimaging Division, Lawrence Berkeley National Laboratory , Berkeley, CA 94720 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-7921-6535","authenticated-orcid":false,"given":"Alexa","family":"Ma","sequence":"additional","affiliation":[{"name":"Department of Chemistry, University of California , Berkeley, CA 94720 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-2204-6927","authenticated-orcid":false,"given":"Georgia","family":"Taylor","sequence":"additional","affiliation":[{"name":"Department of Plant Sciences, University of Cambridge , CB2 3EA Cambridge ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7064-1917","authenticated-orcid":false,"given":"Alistair","family":"Leverett","sequence":"additional","affiliation":[{"name":"Department of Plant Sciences, University of Cambridge , CB2 3EA Cambridge ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0847-1838","authenticated-orcid":false,"given":"Graham R","family":"Fleming","sequence":"additional","affiliation":[{"name":"Graduate Group in Biophysics, University of California , Berkeley, CA 94720 ,","place":["United States"]},{"name":"Molecular Biophysics and Integrated Bioimaging Division, Lawrence Berkeley National Laboratory , Berkeley, CA 94720 ,","place":["United States"]},{"name":"Department of Chemistry, University of California , Berkeley, CA 94720 ,","place":["United States"]},{"name":"Kavli Energy Nanoscience Institute , University of California, Berkeley, CA 94720 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7229-2071","authenticated-orcid":false,"given":"Krishna K","family":"Niyogi","sequence":"additional","affiliation":[{"name":"Department of Plant and Microbial Biology, University of California , Berkeley, CA 94720 ,","place":["United States"]},{"name":"Howard Hughes Medical Institute, University of California , Berkeley, CA 94720 ,","place":["United States"]},{"name":"Molecular Biophysics and Integrated Bioimaging Division, Lawrence Berkeley National Laboratory , Berkeley, CA 94720 ,","place":["United States"]},{"name":"Innovative Genomics Institute, University of California , Berkeley, CA 94720 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4423-4100","authenticated-orcid":false,"given":"Johannes","family":"Kromdijk","sequence":"additional","affiliation":[{"name":"Department of Plant Sciences, University of Cambridge , CB2 3EA Cambridge ,","place":["United Kingdom"]},{"name":"Carl R Woese Institute for Genomic Biology, University of Illinois, 1206 W Gregory Drive , Urbana, IL IL61801 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,4,16]]},"reference":[{"key":"2026052107515930400_kiag207-B1","doi-asserted-by":"publisher","first-page":"1075","DOI":"10.1016\/j.bbabio.2018.06.011","article-title":"Correlated fluorescence quenching and topographic mapping of light-harvesting Complex II within surface-assembled aggregates and lipid bilayers","volume":"1859","author":"Adams","year":"2018","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2026052107515930400_kiag207-B2","doi-asserted-by":"publisher","first-page":"794","DOI":"10.1126\/science.1154800","article-title":"Architecture of a charge-transfer state regulating light harvesting in a plant antenna protein","volume":"320","author":"Ahn","year":"2008","journal-title":"Science"},{"key":"2026052107515930400_kiag207-B3","doi-asserted-by":"publisher","first-page":"1193","DOI":"10.1105\/tpc.13.5.1193","article-title":"Antisense inhibition of the photosynthetic antenna proteins CP29 and CP26: implications for the mechanism of protective energy dissipation","volume":"13","author":"Andersson","year":"2001","journal-title":"Plant Cell"},{"key":"2026052107515930400_kiag207-B4","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1093\/jxb\/eri041","article-title":"Dynamics of photosystem II: a proteomic approach to thylakoid protein complexes","volume":"56","author":"Aro","year":"2005","journal-title":"J Exp Bot"},{"key":"2026052107515930400_kiag207-B5","doi-asserted-by":"publisher","first-page":"3550","DOI":"10.1074\/jbc.M705645200","article-title":"Zeaxanthin radical cation formation in minor light-harvesting complexes of higher plant antenna","volume":"283","author":"Avenson","year":"2008","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B6","doi-asserted-by":"publisher","first-page":"1107","DOI":"10.1111\/j.1365-3040.2007.01680.x","article-title":"Determining the limitations and regulation of photosynthetic energy transduction in leaves","volume":"30","author":"Baker","year":"2007","journal-title":"Plant Cell Environ"},{"key":"2026052107515930400_kiag207-B7","doi-asserted-by":"publisher","first-page":"8103","DOI":"10.1074\/jbc.M808326200","article-title":"Occupancy and functional architecture of the pigment binding sites of photosystem II antenna complex Lhcb5","volume":"284","author":"Ballottari","year":"2009","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B8","doi-asserted-by":"publisher","first-page":"11337","DOI":"10.1021\/jp402977y","article-title":"Chlorophyll triplet quenching and photoprotection in the higher plant monomeric antenna protein Lhcb5","volume":"117","author":"Ballottari","year":"2013","journal-title":"J Phys Chem B"},{"key":"2026052107515930400_kiag207-B9","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1146\/annurev-arplant-071720-015522","article-title":"Dissipation of light energy absorbed in excess: the molecular mechanisms","volume":"72","author":"Bassi","year":"2021","journal-title":"Annu Rev Plant Biol"},{"key":"2026052107515930400_kiag207-B10","doi-asserted-by":"publisher","first-page":"15265","DOI":"10.1073\/pnas.2533072100","article-title":"Light- and pH-dependent structural changes in the PsbS subunit of photosystem II","volume":"100","author":"Bergantino","year":"2003","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026052107515930400_kiag207-B11","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1104\/pp.114.252379","article-title":"High light-dependent phosphorylation of photosystem II inner antenna CP29 in monocots is STN7 independent and enhances nonphotochemical quenching","volume":"167","author":"Betterle","year":"2015","journal-title":"Plant Physiol"},{"key":"2026052107515930400_kiag207-B12","doi-asserted-by":"publisher","first-page":"15255","DOI":"10.1074\/jbc.M808625200","article-title":"Light-induced dissociation of an antenna hetero-oligomer is needed for non-photochemical quenching induction","volume":"284","author":"Betterle","year":"2009","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B13","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1111\/tpj.13412","article-title":"The STN8 kinase-PBCP phosphatase system is responsible for high-light-induced reversible phosphorylation of the PSII inner antenna subunit CP29 in rice","volume":"89","author":"Betterle","year":"2017","journal-title":"Plant J"},{"key":"2026052107515930400_kiag207-B14","doi-asserted-by":"publisher","first-page":"3052","DOI":"10.1038\/emboj.2009.232","article-title":"Functional architecture of higher plant photosystem II supercomplexes","volume":"28","author":"Caffarri","year":"2009","journal-title":"EMBO J"},{"key":"2026052107515930400_kiag207-B15","doi-asserted-by":"publisher","first-page":"496","DOI":"10.1111\/pce.13680","article-title":"Photosystem II antenna complexes CP26 and CP29 are essential for nonphotochemical quenching in Chlamydomonas reinhardtii","volume":"43","author":"Cazzaniga","year":"2020","journal-title":"Plant Cell Environ"},{"key":"2026052107515930400_kiag207-B16","doi-asserted-by":"publisher","first-page":"1365","DOI":"10.1093\/plphys\/kiad391","article-title":"Photosystem II monomeric antenna CP26 plays a key role in nonphotochemical quenching in Chlamydomonas","volume":"193","author":"Cazzaniga","year":"2023","journal-title":"Plant Physiol"},{"key":"2026052107515930400_kiag207-B17","doi-asserted-by":"publisher","DOI":"10.1016\/j.xplc.2024.101179","article-title":"Distinct features of PsbS essential for mediating plant photoprotection","volume":"6","author":"Chen","year":"2025","journal-title":"Plant Commun"},{"key":"2026052107515930400_kiag207-B18","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1016\/j.plaphy.2018.07.014","article-title":"The roles of Arabidopsis proteins of Lhcb4, Lhcb5 and Lhcb6 in oxidative stress under natural light conditions","volume":"130","author":"Chen","year":"2018","journal-title":"Plant Physiol Biochem"},{"key":"2026052107515930400_kiag207-B19","doi-asserted-by":"publisher","first-page":"7152","DOI":"10.1038\/s41467-021-27526-8","article-title":"A different perspective for nonphotochemical quenching in plant antenna complexes","volume":"12","author":"Cignoni","year":"2021","journal-title":"Nat Commun"},{"key":"2026052107515930400_kiag207-B20","doi-asserted-by":"publisher","first-page":"15225","DOI":"10.1038\/nplants.2015.225","article-title":"Psbs interactions involved in the activation of energy dissipation in Arabidopsis","volume":"2","author":"Correa-Galvis","year":"2016","journal-title":"Nat Plants"},{"key":"2026052107515930400_kiag207-B21","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1016\/j.jphotobiol.2011.02.015","article-title":"Light-harvesting and structural organization of photosystem II: from individual complexes to thylakoid membrane","volume":"104","author":"Croce","year":"2011","journal-title":"J Photochem Photobiol B"},{"key":"2026052107515930400_kiag207-B22","doi-asserted-by":"publisher","first-page":"1226","DOI":"10.1021\/bi0018741","article-title":"Contribution of electric field (Delta psi) to steady-state transthylakoid proton motive force (pmf) in vitro and in vivo. Control of pmf parsing into Delta psi and Delta pH by ionic strength","volume":"40","author":"Cruz","year":"2001","journal-title":"Biochemistry"},{"key":"2026052107515930400_kiag207-B23","doi-asserted-by":"publisher","first-page":"1217","DOI":"10.1105\/tpc.104.030601","article-title":"A mechanism of nonphotochemical energy dissipation, independent from PsbS, revealed by a conformational change in the antenna protein CP26","volume":"17","author":"Dall\u2019Osto","year":"2005","journal-title":"Plant Cell"},{"key":"2026052107515930400_kiag207-B24","doi-asserted-by":"publisher","first-page":"17033","DOI":"10.1038\/nplants.2017.33","article-title":"Two mechanisms for dissipation of excess light in monomeric and trimeric light-harvesting complexes","volume":"3","author":"Dall\u2019Osto","year":"2017","journal-title":"Nat Plants"},{"key":"2026052107515930400_kiag207-B25","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2019.183059","article-title":"Fine tuning of the photosystem II major antenna mobility within the thylakoid membrane of higher plants","volume":"1861","author":"Daskalakis","year":"2019","journal-title":"Biochim Biophys Acta Biomembr"},{"key":"2026052107515930400_kiag207-B26","doi-asserted-by":"publisher","first-page":"1012","DOI":"10.1105\/tpc.107.055749","article-title":"Minor antenna proteins CP24 and CP26 affect the interactions between photosystem II subunits and the electron transport rate in grana membranes of Arabidopsis","volume":"20","author":"De Bianchi","year":"2008","journal-title":"Plant Cell"},{"key":"2026052107515930400_kiag207-B27","doi-asserted-by":"publisher","first-page":"2659","DOI":"10.1105\/tpc.111.087320","article-title":"Arabidopsis mutants deleted in the light-harvesting protein Lhcb4 have a disrupted photosystem II macrostructure and are defective in photoprotection","volume":"23","author":"De Bianchi","year":"2011","journal-title":"Plant Cell"},{"key":"2026052107515930400_kiag207-B28","doi-asserted-by":"publisher","first-page":"1279","DOI":"10.1007\/s43630-023-00368-7","article-title":"The origin of pigment-binding differences in CP29 and LHCII: the role of protein structure and dynamics","volume":"22","author":"Elias","year":"2023","journal-title":"Photochem Photobiol Sci"},{"key":"2026052107515930400_kiag207-B29","doi-asserted-by":"publisher","first-page":"6585","DOI":"10.1021\/jp4004328","article-title":"Spectroscopic study of the light-harvesting CP29 antenna complex of photosystem II","volume":"117","author":"Feng","year":"2013","journal-title":"J Phys Chem B"},{"key":"2026052107515930400_kiag207-B30","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1007\/s11120-019-00676-z","article-title":"Carotenoid dark state to chlorophyll energy transfer in isolated light-harvesting complexes CP24 and CP29","volume":"143","author":"Gacek","year":"2020","journal-title":"Photosynth Res"},{"key":"2026052107515930400_kiag207-B31","doi-asserted-by":"publisher","first-page":"2273","DOI":"10.1073\/pnas.92.6.2273","article-title":"Xanthophyll cycle-dependent quenching of photosystem II chlorophyll a fluorescence: formation of a quenching complex with a short fluorescence lifetime","volume":"92","author":"Gilmore","year":"1995","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026052107515930400_kiag207-B32","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1111\/j.1365-313X.2011.04790.x","article-title":"Light-harvesting antenna composition controls the macrostructure and dynamics of thylakoid membranes in Arabidopsis","volume":"69","author":"Goral","year":"2012","journal-title":"Plant J"},{"key":"2026052107515930400_kiag207-B33","doi-asserted-by":"publisher","first-page":"649","DOI":"10.1111\/pce.13686","article-title":"Photosystem II 22\u2005kDa protein level \u2013 a prerequisite for excess light-inducible memory, cross-tolerance to UV-C and regulation of electrical signalling","volume":"43","author":"G\u00f3recka","year":"2020","journal-title":"Plant Cell Environ"},{"key":"2026052107515930400_kiag207-B34","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1038\/s41477-020-0612-8","article-title":"Identification of a pigment cluster catalysing fast photoprotective quenching response in CP29","volume":"6","author":"Guardini","year":"2020","journal-title":"Nat Plants"},{"key":"2026052107515930400_kiag207-B35","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbabio.2022.148555","article-title":"Loss of a single chlorophyll in CP29 triggers re-organization of the photosystem II supramolecular assembly","volume":"1863","author":"Guardini","year":"2022","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2026052107515930400_kiag207-B36","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1126\/science.1105833","article-title":"Carotenoid cation formation and the regulation of photosynthetic light harvesting","volume":"307","author":"Holt","year":"2005","journal-title":"Science"},{"key":"2026052107515930400_kiag207-B37","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/0014-5793(91)80819-O","article-title":"Control of the light-harvesting function of chloroplast membranes by aggregation of the LHCII chlorophyll-protein complex","volume":"292","author":"Horton","year":"1991","journal-title":"FEBS Lett"},{"key":"2026052107515930400_kiag207-B38","doi-asserted-by":"publisher","first-page":"2691","DOI":"10.1093\/plphys\/kiab396","article-title":"Towards spruce-type photosystem II: consequences of the loss of light-harvesting proteins LHCB3 and LHCB6 in Arabidopsis","volume":"187","author":"Il\u00edkov\u00e1","year":"2021","journal-title":"Plant Physiol"},{"key":"2026052107515930400_kiag207-B39","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1042\/BJ20102155","article-title":"Optimized native gel systems for separation of thylakoid protein complexes: novel super- and mega-complexes","volume":"439","author":"J\u00e4rvi","year":"2011","journal-title":"Biochem J"},{"key":"2026052107515930400_kiag207-B40","doi-asserted-by":"publisher","first-page":"3972","DOI":"10.1074\/jbc.M707410200","article-title":"The PsbS protein controls the organization of the photosystem II antenna in higher plant thylakoid membranes","volume":"283","author":"Kiss","year":"2008","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B41","doi-asserted-by":"publisher","first-page":"3106","DOI":"10.1105\/tpc.106.045641","article-title":"Lack of the light-harvesting complex CP24 affects the structure and function of the grana membranes of higher plant chloroplasts","volume":"18","author":"Kov\u00e1cs","year":"2006","journal-title":"Plant Cell"},{"key":"2026052107515930400_kiag207-B42","doi-asserted-by":"publisher","first-page":"15200","DOI":"10.1038\/s41598-017-15068-3","article-title":"Large-scale in vitro production, refolding and dimerization of PsbS in different microenvironments","volume":"7","author":"Krishnan","year":"2017","journal-title":"Sci Rep"},{"key":"2026052107515930400_kiag207-B43","doi-asserted-by":"publisher","first-page":"E7009","DOI":"10.1073\/pnas.1704502114","article-title":"Dissecting and modeling zeaxanthin- and lutein-dependent nonphotochemical quenching in Arabidopsis thaliana","volume":"114","author":"Leuenberger","year":"2017","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026052107515930400_kiag207-B44","doi-asserted-by":"publisher","first-page":"15222","DOI":"10.1073\/pnas.232447699","article-title":"PsbS-dependent enhancement of feedback de-excitation protects photosystem II from photoinhibition","volume":"99","author":"Li","year":"2002","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026052107515930400_kiag207-B45","doi-asserted-by":"publisher","first-page":"1131","DOI":"10.1071\/FP02065","article-title":"Structure-function analysis of photosystem II subunit S (PsbS) in vivo","volume":"29","author":"Li","year":"2002","journal-title":"Funct Plant Biol"},{"key":"2026052107515930400_kiag207-B46","doi-asserted-by":"publisher","first-page":"391","DOI":"10.1038\/35000131","article-title":"A pigment-binding protein essential for regulation of photosynthetic light harvesting","volume":"403","author":"Li","year":"2000","journal-title":"Nature"},{"key":"2026052107515930400_kiag207-B47","doi-asserted-by":"publisher","first-page":"22866","DOI":"10.1074\/jbc.M402461200","article-title":"Regulation of photosynthetic light harvesting involves intrathylakoid lumen pH sensing by the PsbS protein","volume":"279","author":"Li","year":"2004","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B48","doi-asserted-by":"publisher","first-page":"1798","DOI":"10.1105\/tpc.109.066571","article-title":"Lutein accumulation in the absence of zeaxanthin restores nonphotochemical quenching in the Arabidopsis thaliana npq1 mutant","volume":"21","author":"Li","year":"2009","journal-title":"Plant Cell"},{"key":"2026052107515930400_kiag207-B49","doi-asserted-by":"publisher","first-page":"1737","DOI":"10.1021\/acs.jpclett.9b00437","article-title":"Molecular anatomy of plant photoprotective switches: the sensitivity of PsbS to the environment, residue by residue","volume":"10","author":"Liguori","year":"2019","journal-title":"J Phys Chem Lett"},{"key":"2026052107515930400_kiag207-B50","doi-asserted-by":"publisher","DOI":"10.1016\/j.jmb.2023.168407","article-title":"Photosystem II subunit S (PsbS): a nano regulator of plant photosynthesis","volume":"436","author":"Marulanda Valencia","year":"2024","journal-title":"J Mol Biol"},{"key":"2026052107515930400_kiag207-B51","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbabio.2020.148156","article-title":"Design principles of solar light harvesting in plants: functional architecture of the monomeric antenna CP29","volume":"1861","author":"Mascoli","year":"2020","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2026052107515930400_kiag207-B52","doi-asserted-by":"publisher","first-page":"1795","DOI":"10.1093\/jxb\/erj196","article-title":"Controlled levels of salicylic acid are required for optimal photosynthesis and redox homeostasis","volume":"57","author":"Mateo","year":"2006","journal-title":"J Exp Bot"},{"key":"2026052107515930400_kiag207-B53","doi-asserted-by":"publisher","first-page":"36830","DOI":"10.1074\/jbc.M111.273227","article-title":"Quenching in Arabidopsis thaliana mutants lacking monomeric antenna proteins of photosystem II","volume":"286","author":"Miloslavina","year":"2011","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B54","doi-asserted-by":"publisher","first-page":"3625","DOI":"10.1016\/j.febslet.2008.09.044","article-title":"Far-red fluorescence: a direct spectroscopic marker for LHCII oligomer formation in non-photochemical quenching","volume":"582","author":"Miloslavina","year":"2008","journal-title":"FEBS Lett"},{"key":"2026052107515930400_kiag207-B55","doi-asserted-by":"publisher","first-page":"36913","DOI":"10.1074\/jbc.M205339200","article-title":"Dynamics of chromophore binding to Lhc proteins in vivo and in vitro during operation of the xanthophyll cycle","volume":"277","author":"Morosinotto","year":"2002","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B56","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1016\/0005-2728(93)90184-H","article-title":"Excitation-energy quenching in aggregates of the LHC II chlorophyll-protein complex: a time-resolved fluorescence study","volume":"1141","author":"Mullineaux","year":"1993","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2026052107515930400_kiag207-B57","doi-asserted-by":"publisher","first-page":"16730","DOI":"10.1074\/jbc.M116.730101","article-title":"Light-harvesting complexes (LHCs) cluster spontaneously in membrane environment leading to shortening of their excited state lifetimes","volume":"291","author":"Natali","year":"2016","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B58","doi-asserted-by":"publisher","first-page":"7415","DOI":"10.1038\/s41598-021-86975-9","article-title":"The PsbS protein and low pH are necessary and sufficient to induce quenching in the light-harvesting complex of plants LHCII","volume":"11","author":"Nicol","year":"2021","journal-title":"Sci Rep"},{"key":"2026052107515930400_kiag207-B59","doi-asserted-by":"publisher","first-page":"1177","DOI":"10.1038\/s41477-019-0526-5","article-title":"Disentangling the sites of non-photochemical quenching in vascular plants","volume":"5","author":"Nicol","year":"2019","journal-title":"Nat Plants"},{"key":"2026052107515930400_kiag207-B60","doi-asserted-by":"publisher","first-page":"466","DOI":"10.1016\/j.bbabio.2010.01.001","article-title":"Identification of a slowly inducible zeaxanthin-dependent component of non-photochemical quenching of chlorophyll fluorescence generated under steady-state conditions in Arabidopsis","volume":"1797","author":"Nilkens","year":"2010","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2026052107515930400_kiag207-B61","doi-asserted-by":"publisher","first-page":"1121","DOI":"10.1105\/tpc.10.7.1121","article-title":"Arabidopsis mutants define a central role for the xanthophyll cycle in the regulation of photosynthetic energy conversion","volume":"10","author":"Niyogi","year":"1998","journal-title":"Plant Cell"},{"key":"2026052107515930400_kiag207-B62","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1023\/A:1005936823310","article-title":"Resolving chlorophyll a fluorescence images of photosynthetic efficiency into photochemical and non-photochemical components \u2013 calculation of qP and Fv-\/Fm- without measuring Fo","volume":"54","author":"Oxborough","year":"1997","journal-title":"Photosynth Res"},{"key":"2026052107515930400_kiag207-B63","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1007\/s11120-020-00740-z","article-title":"On the PsbS-induced quenching in the plant major light-harvesting complex LHCII studied in proteoliposomes","volume":"144","author":"Pawlak","year":"2020","journal-title":"Photosynth Res"},{"key":"2026052107515930400_kiag207-B64","doi-asserted-by":"publisher","first-page":"1870","DOI":"10.1016\/j.bbabio.2016.09.001","article-title":"Electron transfer between carotenoid and chlorophyll contributes to quenching in the LHCSR1 protein from Physcomitrella patens","volume":"1857","author":"Pinnola","year":"2016","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2026052107515930400_kiag207-B65","doi-asserted-by":"publisher","first-page":"384","DOI":"10.1016\/S0005-2728(89)80347-0","article-title":"Determination of accurate extinction coefficients and simultaneous equations for assaying chlorophylls a and b extracted with four different solvents: verification of the concentration of chlorophyll standards by atomic absorption spectroscopy","volume":"975","author":"Porra","year":"1989","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2026052107515930400_kiag207-B66","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1016\/0014-5793(92)81089-5","article-title":"The molecular mechanism of the control of excitation energy dissipation in chloroplast membranes: inhibition of delta pH-dependent quenching of chlorophyll fluorescence by dicyclohexylcarbodiimide","volume":"309","author":"Ruban","year":"1992","journal-title":"FEBS Lett"},{"key":"2026052107515930400_kiag207-B67","doi-asserted-by":"publisher","first-page":"1063","DOI":"10.1093\/pcp\/pcaa155","article-title":"The mechanism of non-photochemical quenching in plants: localization and driving forces","volume":"62","author":"Ruban","year":"2021","journal-title":"Plant Cell Physiol"},{"key":"2026052107515930400_kiag207-B68","doi-asserted-by":"publisher","first-page":"3626","DOI":"10.1093\/jxb\/eraa126","article-title":"Rapid regulation of photosynthetic light harvesting in the absence of minor antenna and reaction centre complexes","volume":"71","author":"Saccon","year":"2020","journal-title":"J Exp Bot"},{"key":"2026052107515930400_kiag207-B69","doi-asserted-by":"publisher","first-page":"16225","DOI":"10.1038\/nplants.2016.225","article-title":"The xanthophyll cycle affects reversible interactions between PsbS and light-harvesting complex II to control non-photochemical quenching","volume":"3","author":"Sacharz","year":"2017","journal-title":"Nat Plants"},{"key":"2026052107515930400_kiag207-B70","doi-asserted-by":"publisher","DOI":"10.1063\/5.0087898","article-title":"Molecular mechanisms of light harvesting in the minor antenna CP29 in near-native membrane lipidic environment","volume":"156","author":"Sardar","year":"2022","journal-title":"J Chem Phys"},{"key":"2026052107515930400_kiag207-B71","doi-asserted-by":"publisher","first-page":"3149","DOI":"10.1021\/acs.jpclett.3c02900","article-title":"Site-directed mutagenesis of the chlorophyll-binding sites modulates excited-state lifetime and chlorophyll-xanthophyll energy transfer in the monomeric light-harvesting complex CP29","volume":"15","author":"Sardar","year":"2024","journal-title":"J Phys Chem Lett"},{"key":"2026052107515930400_kiag207-B72","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1186\/s13007-022-00858-2","article-title":"Qualitative and quantitative evaluation of thylakoid complexes separated by blue native PAGE","volume":"18","author":"S\u00e1rv\u00e1ri","year":"2022","journal-title":"Plant Methods"},{"key":"2026052107515930400_kiag207-B73","doi-asserted-by":"publisher","first-page":"17816","DOI":"10.1074\/jbc.RA120.016181","article-title":"A novel method produces native light-harvesting complex II aggregates from the photosynthetic membrane revealing their role in nonphotochemical quenching","volume":"295","author":"Shukla","year":"2020","journal-title":"J Biol Chem"},{"key":"2026052107515930400_kiag207-B74","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1385\/1-59259-413-1:241","article-title":"High-throughput TAIL-PCR as a tool to identify DNA flanking insertions","volume":"236","author":"Singer","year":"2003","journal-title":"Methods Mol Biol"},{"key":"2026052107515930400_kiag207-B75","doi-asserted-by":"publisher","first-page":"10311","DOI":"10.1021\/acs.jpcb.0c06265","article-title":"Complex roles of PsbS and xanthophylls in the regulation of nonphotochemical quenching in Arabidopsis thaliana under fluctuating light","volume":"124","author":"Steen","year":"2020","journal-title":"J Phys Chem B"},{"key":"2026052107515930400_kiag207-B76","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1007\/s11120-015-0104-2","article-title":"Characterizing non-photochemical quenching in leaves through fluorescence lifetime snapshots","volume":"127","author":"Sylak-Glassman","year":"2016","journal-title":"Photosynth Res"},{"key":"2026052107515930400_kiag207-B77","doi-asserted-by":"publisher","first-page":"2339","DOI":"10.1021\/acs.jpclett.5b00944","article-title":"LHCII populations in different quenching states are present in the thylakoid membranes in a ratio that depends on the light conditions","volume":"6","author":"Tian","year":"2015","journal-title":"J Phys Chem Lett"},{"key":"2026052107515930400_kiag207-B78","doi-asserted-by":"publisher","first-page":"1243","DOI":"10.1111\/pce.12924","article-title":"NPQ(T): a chlorophyll fluorescence parameter for rapid estimation and imaging of non-photochemical quenching of excitons in photosystem-II-associated antenna complexes","volume":"40","author":"Tietz","year":"2017","journal-title":"Plant Cell Environ"},{"key":"2026052107515930400_kiag207-B79","doi-asserted-by":"publisher","first-page":"666","DOI":"10.1016\/j.bbabio.2018.03.005","article-title":"The causes of altered chlorophyll fluorescence quenching induction in the Arabidopsis mutant lacking all minor antenna complexes","volume":"1859","author":"Townsend","year":"2018","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2026052107515930400_kiag207-B80","doi-asserted-by":"publisher","DOI":"10.1016\/j.jphotobiol.2021.112174","article-title":"Aggregation-related quenching of LHCII fluorescence in liposomes revealed by single-molecule spectroscopy","volume":"218","author":"Tutkus","year":"2021","journal-title":"J Photochem Photobiol B"},{"key":"2026052107515930400_kiag207-B81","doi-asserted-by":"publisher","DOI":"10.1093\/plcell\/koaf240","article-title":"Nonphotochemical quenching in plants: mechanisms and mysteries","volume":"37","author":"Van Amerongen","year":"2025","journal-title":"Plant Cell."},{"key":"2026052107515930400_kiag207-B82","doi-asserted-by":"publisher","first-page":"922","DOI":"10.1016\/j.bpj.2009.11.012","article-title":"Effect of antenna-depletion in photosystem II on excitation energy transfer in Arabidopsis thaliana","volume":"98","author":"Van Oort","year":"2010","journal-title":"Biophys J"},{"key":"2026052107515930400_kiag207-B83","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.70167","article-title":"Photosystem II supercomplexes lacking light-harvesting antenna protein LHCB5 and their organization in the thylakoid membrane","volume":"177","author":"V\u00e1nsk\u00e1","year":"2025","journal-title":"Physiol Plant"},{"key":"2026052107515930400_kiag207-B84","doi-asserted-by":"publisher","first-page":"14204","DOI":"10.1073\/pnas.93.24.14204","article-title":"Identification of proton-active residues in a higher plant light-harvesting complex","volume":"93","author":"Walters","year":"1996","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026052107515930400_kiag207-B85","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1016\/j.jphotobiol.2015.07.016","article-title":"Psbs protein modulates non-photochemical chlorophyll fluorescence quenching in membranes depleted of photosystems","volume":"152","author":"Ware","year":"2015","journal-title":"J Photochem Photobiol B"},{"key":"2026052107515930400_kiag207-B86","doi-asserted-by":"publisher","first-page":"5452","DOI":"10.1073\/pnas.1205561110","article-title":"Direct interaction of the major light-harvesting complex II and PsbS in nonphotochemical quenching","volume":"110","author":"Wilk","year":"2013","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026052107515930400_kiag207-B87","doi-asserted-by":"publisher","first-page":"5683","DOI":"10.1021\/acs.jpclett.2c01019","article-title":"The structural and spectral features of light-harvesting complex II proteoliposomes mimic those of native thylakoid membranes","volume":"13","author":"Wilson","year":"2022","journal-title":"J Phys Chem Lett"},{"key":"2026052107515930400_kiag207-B88","doi-asserted-by":"publisher","first-page":"1795","DOI":"10.1104\/pp.112.208116","article-title":"LESION SIMULATING DISEASE1, ENHANCED DISEASE SUSCEPTIBILITY1, and PHYTOALEXIN DEFICIENT4 conditionally regulate cellular signaling homeostasis, photosynthesis, water use efficiency, and seed yield in Arabidopsis","volume":"161","author":"Wituszy\u0144ska","year":"2013","journal-title":"Plant Physiol"},{"key":"2026052107515930400_kiag207-B89","doi-asserted-by":"publisher","first-page":"548","DOI":"10.1016\/j.jplph.2012.11.016","article-title":"Multivariable environmental conditions promote photosynthetic adaptation potential in Arabidopsis thaliana","volume":"170","author":"Wituszy\u0144ska","year":"2013","journal-title":"J Plant Physiol"},{"key":"2026052107515930400_kiag207-B90","doi-asserted-by":"publisher","first-page":"13679","DOI":"10.1038\/srep13679","article-title":"Molecular insights into zeaxanthin-dependent quenching in higher plants","volume":"5","author":"Xu","year":"2015","journal-title":"Sci Rep"},{"key":"2026052107515930400_kiag207-B91","doi-asserted-by":"publisher","first-page":"608","DOI":"10.1021\/bi027109z","article-title":"The structure of photosystem II in Arabidopsis: localization of the CP26 and CP29 antenna complexes","volume":"42","author":"Yakushevska","year":"2003","journal-title":"Biochemistry"}],"container-title":["Plant Physiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/plphys\/advance-article-pdf\/doi\/10.1093\/plphys\/kiag207\/68098531\/kiag207.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/plphys\/article-pdf\/201\/1\/kiag207\/68098531\/kiag207.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/plphys\/article-pdf\/201\/1\/kiag207\/68098531\/kiag207.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T11:52:05Z","timestamp":1779364325000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/plphys\/article\/doi\/10.1093\/plphys\/kiag207\/8656904"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,16]]},"references-count":91,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,5,6]]}},"URL":"https:\/\/doi.org\/10.1093\/plphys\/kiag207","relation":{},"ISSN":["0032-0889","1532-2548"],"issn-type":[{"value":"0032-0889","type":"print"},{"value":"1532-2548","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,5]]},"published":{"date-parts":[[2026,4,16]]},"article-number":"kiag207"}}