{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T06:02:26Z","timestamp":1789624946134,"version":"build-2803163510"},"reference-count":66,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2026,9,16]],"date-time":"2026-09-16T00:00:00Z","timestamp":1789516800000},"content-version":"vor","delay-in-days":15,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100001700","name":"Ministry of Education, Culture, Sports, Science and Technology","doi-asserted-by":"publisher","award":["26H01084"],"award-info":[{"award-number":["26H01084"]}],"id":[{"id":"10.13039\/501100001700","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001700","name":"Ministry of Education, Culture, Sports, Science and Technology","doi-asserted-by":"publisher","award":["24H02271"],"award-info":[{"award-number":["24H02271"]}],"id":[{"id":"10.13039\/501100001700","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Physiologia Plantarum"],"published-print":{"date-parts":[[2026,9]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Ycf10 and DLDG1 are homologous proteins embedded in the chloroplast envelope membranes (EM) and encoded in the plastidial and nuclear genomes, respectively. We previously characterized Arabidopsis\n                    <jats:italic>dldg1<\/jats:italic>\n                    and tobacco\n                    <jats:italic>ycf10<\/jats:italic>\n                    mutants, and showed that these proteins mediate proton transport across the EM, thereby influencing ATP synthesis and the non\u2010photochemical quenching (NPQ) induction. However, the consequences of the simultaneous loss of Ycf10\/DLDG1 homologs have remained unclear. Here, we generated and analyzed Arabidopsis\n                    <jats:italic>ycf10<\/jats:italic>\n                    and\n                    <jats:italic>ycf10\u2010dldg1<\/jats:italic>\n                    double mutants. These mutants exhibited a pale\u2010green phenotype under continuous light conditions, which was complemented by the exogenous addition of NaCl, as previously observed in mutants lacking the plastid EM\u2010localized K\n                    <jats:sup>+<\/jats:sup>\n                    \/H\n                    <jats:sup>+<\/jats:sup>\n                    antiporters KEA1 and KEA2. Both single mutants (\n                    <jats:italic>ycf10<\/jats:italic>\n                    and\n                    <jats:italic>dldg1<\/jats:italic>\n                    ), as well as the\n                    <jats:italic>ycf10\u2010dldg1<\/jats:italic>\n                    double mutant, showed enhanced NPQ induction compared with wild\u2010type. Furthermore, the\n                    <jats:italic>ycf10\u2010dldg1<\/jats:italic>\n                    double mutant showed stronger NPQ induction than either single mutant upon transition from dark to light, suggesting that Ycf10 and DLDG1 function independently while playing redundant roles. These mutants also showed reduced stomatal conductance and sugar accumulation compared with the wild\u2010type. Together, these findings indicate that EM\u2010localized DLDG1 and Ycf10 act in concert to maintain chloroplast proton\/ion homeostasis, contributing to chloroplast pH homeostasis for sustaining plant growth and efficient photosynthesis.\n                  <\/jats:p>","DOI":"10.1111\/ppl.71114","type":"journal-article","created":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T05:13:18Z","timestamp":1789621998000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Impact of the Simultaneous Loss of the Plastid Envelope\u2010Localized Homologous Pair Ycf10 and\n                    <scp>DLDG1<\/scp>\n                    in Plant Growth and Photosynthetic Performance"],"prefix":"10.1111","volume":"178","author":[{"given":"Elham","family":"Esmailpourmoghadam","sequence":"first","affiliation":[{"name":"Department of Life Science and Technology Institute of Science Tokyo  Yokohama Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3428-3657","authenticated-orcid":false,"given":"Mai Duy Luu","family":"Trinh","sequence":"additional","affiliation":[{"name":"Department of Plant and Environmental Sciences University of Copenhagen  Frederiksberg C Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryoichi","family":"Sato","sequence":"additional","affiliation":[{"name":"RIKEN Center for Sustainable Resource Science  Yokohama Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1543-937X","authenticated-orcid":false,"given":"Masaru","family":"Kono","sequence":"additional","affiliation":[{"name":"Astrobiology Center, Mitaka  Tokyo Japan"},{"name":"Laboratory for Chemistry and Life Science, Institute of Integrated Research Institute of Science Tokyo  Yokohama Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5747-6064","authenticated-orcid":false,"given":"Issei","family":"Nakazato","sequence":"additional","affiliation":[{"name":"Graduate School of Agricultural and Life Sciences The University of Tokyo  Tokyo Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9537-1626","authenticated-orcid":false,"given":"Shin\u2010ichi","family":"Arimura","sequence":"additional","affiliation":[{"name":"Graduate School of Agricultural and Life Sciences The University of Tokyo  Tokyo Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9982-6114","authenticated-orcid":false,"given":"Michael","family":"Palmgren","sequence":"additional","affiliation":[{"name":"Department of Plant and Environmental Sciences University of Copenhagen  Frederiksberg C Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5349-4927","authenticated-orcid":false,"given":"Shinji","family":"Masuda","sequence":"additional","affiliation":[{"name":"Department of Life Science and Technology Institute of Science Tokyo  Yokohama Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,16]]},"reference":[{"issue":"33","key":"e_1_2_10_2_1","doi-asserted-by":"crossref","first-page":"10231","DOI":"10.1073\/pnas.1500012112","article-title":"Why Chloroplasts and Mitochondria Retain Their Own Genomes and Genetic Systems: Colocation for Redox Regulation of Gene Expression","volume":"112","author":"Allen J. F.","year":"2015","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"e_1_2_10_3_1","doi-asserted-by":"crossref","first-page":"2080","DOI":"10.1111\/nph.17042","article-title":"Plastidial Transporters KEA1 and KEA2 at the Inner Envelope Membrane Adjust Stromal pH in the Dark","volume":"229","author":"Aranda Sicilia M. N.","year":"2021","journal-title":"New Phytologist"},{"issue":"19","key":"e_1_2_10_4_1","doi-asserted-by":"crossref","first-page":"R911","DOI":"10.1016\/j.cub.2015.07.055","article-title":"Endosymbiosis and Eukaryotic Cell Evolution","volume":"25","author":"Archibald J. M.","year":"2015","journal-title":"Current Biology"},{"key":"e_1_2_10_5_1","first-page":"1557","article-title":"Regulation and Levels of the Thylakoid K+\/H+ Antiporter KEA3 Shape the Dynamic Response of Photosynthesis in Fluctuating Light","volume":"57","author":"Armbruster U.","year":"2016","journal-title":"Plant and Cell Physiology"},{"key":"e_1_2_10_6_1","doi-asserted-by":"crossref","DOI":"10.1038\/ncomms6439","article-title":"Ion Antiport Accelerates Photosynthetic Acclimation in Fluctuating Light Environments","volume":"5","author":"Armbruster U.","year":"2014","journal-title":"Nature Communications"},{"issue":"2","key":"e_1_2_10_7_1","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1093\/dnares\/11.2.93","article-title":"Complete Nucleotide Sequence of the Sugarcane (\n                     Saccharum officinarum\n                  ) Chloroplast Genome: A Comparative Analysis of Four Monocot Chloroplast Genomes","volume":"11","author":"Asano T.","year":"2004","journal-title":"DNA Research"},{"issue":"6","key":"e_1_2_10_8_1","doi-asserted-by":"crossref","first-page":"1642","DOI":"10.1111\/tpj.16658","article-title":"WRKY48 Negatively Regulates Plant Acclimation to a Combination of High Light and Heat Stress","volume":"117","author":"Balfag\u00f3n D.","year":"2024","journal-title":"Plant Journal"},{"issue":"7","key":"e_1_2_10_9_1","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1016\/j.bbabio.2018.03.013","article-title":"The Peculiar NPQ Regulation in the Stramenopile phaeomonas sp. 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H.","year":"1984","journal-title":"Photosynthesis Research"},{"issue":"20","key":"e_1_2_10_29_1","doi-asserted-by":"crossref","first-page":"7480","DOI":"10.1073\/pnas.1323899111","article-title":"Plastidial Transporters KEA1, \u22122, and \u22123 Are Essential for Chloroplast Osmoregulation, Integrity, and pH Regulation in Arabidopsis","volume":"111","author":"Kunz H. H.","year":"2014","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"20","key":"e_1_2_10_30_1","first-page":"4547","article-title":"Age\u2010Associated Growth Control Modifies Leaf Proximodistal Symmetry and Enabled Leaf Shape Diversification","volume":"34","author":"Li X. 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Y.","year":"2020","journal-title":"Nature"},{"issue":"4","key":"e_1_2_10_34_1","doi-asserted-by":"crossref","first-page":"872","DOI":"10.1111\/tpj.17027","article-title":"Genome Editing in Angiosperm Chloroplasts: Targeted DNA Double\u2010Strand Break and Base Editing","volume":"120","author":"Nakazato I.","year":"2024","journal-title":"Plant Journal: For Cell and Molecular Biology"},{"issue":"1","key":"e_1_2_10_35_1","doi-asserted-by":"crossref","DOI":"10.1002\/cpz1.70075","article-title":"Targeted C\u2010To\u2010T Base Editing in the Arabidopsis Plastid Genome","volume":"5","author":"Nakazato I.","year":"2025","journal-title":"Current Protocols"},{"issue":"4","key":"e_1_2_10_36_1","doi-asserted-by":"crossref","first-page":"1151","DOI":"10.1111\/tpj.16311","article-title":"Characterization and Development of a Plastid Genome Base Editor, ptpTALECD","volume":"115","author":"Nakazato I.","year":"2023","journal-title":"Plant Journal"},{"issue":"7","key":"e_1_2_10_37_1","doi-asserted-by":"crossref","first-page":"906","DOI":"10.1038\/s41477-021-00954-6","article-title":"Targeted Base Editing in the Plastid Genome of \n                     Arabidopsis thaliana","volume":"7","author":"Nakazato I.","year":"2021","journal-title":"Nature Plants"},{"issue":"24","key":"e_1_2_10_38_1","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2121177119","article-title":"Targeted Base Editing in the Mitochondrial Genome of \n                     Arabidopsis thaliana","volume":"119","author":"Nakazato I.","year":"2022","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"1","key":"e_1_2_10_39_1","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1111\/pbi.14490","article-title":"Resistance to the Herbicide Metribuzin Conferred to \n                     Arabidopsis thaliana\n                   by Targeted Base Editing of the Chloroplast Genome","volume":"23","author":"Nakazato I.","year":"2025","journal-title":"Plant Biotechnology Journal"},{"issue":"1","key":"e_1_2_10_40_1","doi-asserted-by":"crossref","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\u2010Harvesting Complex of Plants LHCII","volume":"11","author":"Nicol L.","year":"2021","journal-title":"Scientific Reports"},{"issue":"7","key":"e_1_2_10_41_1","doi-asserted-by":"crossref","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 K. 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V.","year":"2021","journal-title":"Plant & Cell Physiology"},{"issue":"1","key":"e_1_2_10_47_1","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/0014-5793(93)81743-J","article-title":"Chloroplast Envelope Protein Encoded by Chloroplast Genome","volume":"316","author":"Sasaki Y.","year":"1993","journal-title":"FEBS Letters"},{"key":"e_1_2_10_48_1","first-page":"1","article-title":"New Accessory for the DUAL\u2010PAM\u2010100: The P515\/535 Module and Examples of Its Application","volume":"1","author":"Schreiber U.","year":"2008","journal-title":"PAM Applications Notes"},{"issue":"15","key":"e_1_2_10_49_1","doi-asserted-by":"crossref","first-page":"3799","DOI":"10.1128\/JB.180.15.3799-3803.1998","article-title":"Photosynthetic Electron Transport Involved in PxcA\u2010Dependent Proton Extrusion in Synechocystis sp. 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