{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,8]],"date-time":"2026-03-08T03:32:26Z","timestamp":1772940746034,"version":"3.50.1"},"reference-count":47,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2025,12,1]],"date-time":"2025-12-01T00:00:00Z","timestamp":1764547200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2025,12,1]],"date-time":"2025-12-01T00:00:00Z","timestamp":1764547200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2025,12,1]],"date-time":"2025-12-01T00:00:00Z","timestamp":1764547200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2025,12,1]],"date-time":"2025-12-01T00:00:00Z","timestamp":1764547200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2025,12,1]],"date-time":"2025-12-01T00:00:00Z","timestamp":1764547200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2025,12,1]],"date-time":"2025-12-01T00:00:00Z","timestamp":1764547200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2025,12,1]],"date-time":"2025-12-01T00:00:00Z","timestamp":1764547200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/501100004543","name":"China Scholarship Council","doi-asserted-by":"publisher","award":["201808330245"],"award-info":[{"award-number":["201808330245"]}],"id":[{"id":"10.13039\/501100004543","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["31670303"],"award-info":[{"award-number":["31670303"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004731","name":"Zhejiang Province Natural Science Foundation","doi-asserted-by":"publisher","award":["LZ21C020001"],"award-info":[{"award-number":["LZ21C020001"]}],"id":[{"id":"10.13039\/501100004731","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Plant Physiology and Biochemistry"],"published-print":{"date-parts":[[2025,12]]},"DOI":"10.1016\/j.plaphy.2025.110394","type":"journal-article","created":{"date-parts":[[2025,8,18]],"date-time":"2025-08-18T07:50:01Z","timestamp":1755503401000},"page":"110394","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"special_numbering":"PA","title":["Thermal protection of the plastid-encoded RNA polymerase complex by PAP9 ensures chloroplast function in Arabidopsis"],"prefix":"10.1016","volume":"229","author":[{"given":"Lei","family":"Chen","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mengjie","family":"Zhao","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Luqi","family":"Tang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wenjun","family":"Hu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tianbao","family":"Lin","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Peigang","family":"Liu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuan","family":"Huang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yan","family":"Zhu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Zhiqiang","family":"Lv","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jia","family":"Wei","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"78","reference":[{"key":"10.1016\/j.plaphy.2025.110394_bib1","doi-asserted-by":"crossref","first-page":"2802","DOI":"10.1002\/j.1460-2075.1996.tb00640.x","article-title":"Deletion of rpoB reveals a second distinct transcription system in plastids of higher plants","volume":"15","author":"Allison","year":"1996","journal-title":"EMBO J."},{"key":"10.1016\/j.plaphy.2025.110394_bib3","first-page":"393","article-title":"Plastid development in leaves during growth and senescence","volume":"36","author":"Biswal","year":"2013","journal-title":"Advances in photosynthesis and"},{"key":"10.1016\/j.plaphy.2025.110394_bib4","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1016\/j.bbabio.2015.02.004","article-title":"Chloroplast RNA polymerases: role in chloroplast biogenesis","volume":"1847","author":"B\u00f6rner","year":"2015","journal-title":"Biochimica Et Biophysica Acta - Bioenergetics"},{"key":"10.1016\/j.plaphy.2025.110394_bib5","doi-asserted-by":"crossref","DOI":"10.1016\/j.plantsci.2022.111359","article-title":"The role of PAP4\/FSD3 and PAP9\/FSD2 in heat stress responses of chloroplast genes","volume":"322","author":"Bychkov","year":"2022","journal-title":"Plant Sci."},{"key":"10.1016\/j.plaphy.2025.110394_bib6","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1104\/pp.17.00248","article-title":"pTAC10, a key subunit of plastid-encoded RNA polymerase, promotes chloroplast development","volume":"174","author":"Chang","year":"2017","journal-title":"Plant Physiology"},{"key":"10.1016\/j.plaphy.2025.110394_bib7","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1111\/j.1365-313X.2008.03634.x","article-title":"CLB19, a pentatricopeptide repeat protein required for editing ofrpoAandclpPchloroplast transcripts","volume":"56","author":"Chateigner-Boutin","year":"2008","journal-title":"Plant J."},{"key":"10.1016\/j.plaphy.2025.110394_bib8","doi-asserted-by":"crossref","first-page":"770","DOI":"10.1016\/j.bbabio.2015.01.001","article-title":"Plastid sigma factors: their individual functions and regulation in transcription","volume":"1847","author":"Chi","year":"2015","journal-title":"Biochimica Et Biophysica Acta - Bioenergetics"},{"key":"10.1016\/j.plaphy.2025.110394_bib9","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.plaphy.2018.05.023","article-title":"Differential impact of heat stress on the expression of chloroplast-encoded genes","volume":"29","author":"Danilova","year":"2018","journal-title":"Plant Physiol. Biochem."},{"key":"10.1016\/j.plaphy.2025.110394_bib10","doi-asserted-by":"crossref","first-page":"1643","DOI":"10.1042\/BST0351643","article-title":"Pentatricopeptide repeat (PPR) proteins as sequence-specificity factors in post-transcriptional processes in organelles","volume":"35","author":"Delannoy","year":"2007","journal-title":"Biochem. Soc. Trans."},{"key":"10.1016\/j.plaphy.2025.110394_bib11","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0220078","article-title":"Chloroplast-localized iron superoxide dismutases FSD2 and FSD3 are functionally distinct in arabidopsis","volume":"14","author":"Gallie","year":"2019","journal-title":"PLoS One"},{"key":"10.1016\/j.plaphy.2025.110394_bib12","doi-asserted-by":"crossref","first-page":"1354","DOI":"10.4161\/psb.21618","article-title":"Regulatory role of arabidopsis pTAC14 in chloroplast development and plastid gene expression","volume":"7","author":"Gao","year":"2012","journal-title":"Plant Signal. Behav."},{"key":"10.1016\/j.plaphy.2025.110394_bib13","doi-asserted-by":"crossref","first-page":"1733","DOI":"10.1104\/pp.111.184762","article-title":"A functional component of the transcriptionally active chromosome complex, arabidopsis pTAC14, interacts with pTAC12\/HEMERA and regulates plastid gene expression","volume":"157","author":"Gao","year":"2011","journal-title":"Plant Physiology"},{"key":"10.1016\/j.plaphy.2025.110394_bib14","doi-asserted-by":"crossref","first-page":"924","DOI":"10.1111\/j.1365-313X.2007.03379.x","article-title":"An arabidopsis homolog of the bacterial peptidoglycan synthesis enzyme MurE has an essential role in chloroplast development","volume":"53","author":"Garcia","year":"2007","journal-title":"Plant J."},{"key":"10.1016\/j.plaphy.2025.110394_bib16","doi-asserted-by":"crossref","first-page":"4041","DOI":"10.1093\/emboj\/16.13.4041","article-title":"The two RNA polymerases encoded by the nuclear and the plastid compartments transcribe distinct groups of genes in tobacco plastids","volume":"16","author":"Hajdukiewicz","year":"1997","journal-title":"EMBO J."},{"key":"10.1016\/j.plaphy.2025.110394_bib17","doi-asserted-by":"crossref","first-page":"7090","DOI":"10.1093\/nar\/gkg935","article-title":"Molecular genetic analysis of chloroplast gene promoters dependent on SIG2, a nucleus\u2010encoded sigma factor for the plastid\u2010encoded RNA polymerase, in Arabidopsis thaliana","volume":"31","author":"Hanaoka","year":"2003","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.plaphy.2025.110394_bib18","doi-asserted-by":"crossref","first-page":"1149","DOI":"10.1093\/pcp\/pci112","article-title":"A resource of 5,814 dissociation transposon-tagged and sequence-indexed lines of arabidopsis transposed from start loci on chromosome 5","volume":"46","author":"Ito","year":"2005","journal-title":"Plant Cell Physiol."},{"key":"10.1016\/j.plaphy.2025.110394_bib19","doi-asserted-by":"crossref","first-page":"3780","DOI":"10.1093\/jxb\/eraa034","article-title":"Molecular and genetic bases of heat stress responses in crop plants and breeding for increased resilience and productivity","volume":"71","author":"Janni","year":"2020","journal-title":"J. Exp. Bot."},{"key":"10.1016\/j.plaphy.2025.110394_bib20","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1111\/j.1469-8137.2011.03941.x","article-title":"S1 domain-containing STF modulates plastid transcription and chloroplast biogenesis in Nicotiana benthamiana","volume":"193","author":"Jeon","year":"2011","journal-title":"New Phytol."},{"key":"10.1016\/j.plaphy.2025.110394_bib23","doi-asserted-by":"crossref","first-page":"354","DOI":"10.1016\/j.cub.2004.02.039","article-title":"The Arabidopsis thaliana chloroplast proteome reveals pathway abundance and novel protein functions","volume":"14","author":"Kleffmann","year":"2004","journal-title":"Curr. Biol."},{"key":"10.1016\/j.plaphy.2025.110394_bib24","doi-asserted-by":"crossref","first-page":"385","DOI":"10.3389\/fpls.2014.00385","article-title":"Plastid encoded RNA polymerase activity and expression of photosynthesis genes required for embryo and seed development in arabidopsis","volume":"5","author":"Kremnev","year":"2014","journal-title":"Front. Plant Sci."},{"key":"10.1016\/j.plaphy.2025.110394_bib25","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1046\/j.1365-313X.2002.01349.x","article-title":"Comparative analysis of plastid transcription profiles of entire plastid chromosomes from tobacco attributed to wild-type and PEP-Deficient transcription machineries","volume":"31","author":"Legen","year":"2002","journal-title":"Plant J."},{"key":"10.1016\/j.plaphy.2025.110394_bib26","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1007\/s11103-010-9714-4","article-title":"Function of plastid sigma factors in higher plants: regulation of gene expression or just preservation of constitutive transcription?","volume":"76","author":"Lerbs-Mache","year":"2010","journal-title":"Plant Mol. Biol."},{"key":"10.1016\/j.plaphy.2025.110394_bib27","doi-asserted-by":"crossref","DOI":"10.15252\/embj.2020104941","article-title":"Nucleo\u2010plastidic PAP8\/pTAC6 couples chloroplast formation with photomorphogenesis","volume":"39","author":"Liebers","year":"2020","journal-title":"EMBO J."},{"key":"10.1016\/j.plaphy.2025.110394_bib28","doi-asserted-by":"crossref","first-page":"1159","DOI":"10.1093\/nar\/gkh285","article-title":"Overexpression of phage-type RNA polymerase RpoTp in tobacco demonstrates its role in chloroplast transcription by recognizing a distinct promoter type","volume":"32","author":"Liere","year":"2004","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.plaphy.2025.110394_bib29","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1089\/152308603321223568","article-title":"Redox regulation of chloroplast transcription","volume":"5","author":"Link","year":"2003","journal-title":"Antioxidants Redox Signal."},{"key":"10.1016\/j.plaphy.2025.110394_bib30","doi-asserted-by":"crossref","first-page":"3148","DOI":"10.1105\/tpc.108.061341","article-title":"A heterocomplex of iron superoxide dismutases defends chloroplast nucleoids against oxidative stress and is essential for chloroplast development inArabidopsis","volume":"20","author":"Myouga","year":"2008","journal-title":"Plant Cell"},{"key":"10.1016\/j.plaphy.2025.110394_bib31","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1074\/mcp.M500180-MCP200","article-title":"The oligomeric stromal proteome of Arabidopsis thaliana chloroplasts","volume":"5","author":"Peltier","year":"2006","journal-title":"Mol. Cell. Proteomics"},{"key":"10.1016\/j.plaphy.2025.110394_bib32","doi-asserted-by":"crossref","first-page":"2042","DOI":"10.1038\/emboj.2009.121","article-title":"Site-specific binding of a PPR protein defines and stabilizes 5\u2032 and 3\u2032 mRNA termini in chloroplasts","volume":"28","author":"Pfalz","year":"2009","journal-title":"EMBO J."},{"key":"10.1016\/j.plaphy.2025.110394_bib34","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1016\/j.tplants.2012.11.003","article-title":"Essential nucleoid proteins in early chloroplast development","volume":"18","author":"Pfalz","year":"2013","journal-title":"Trends Plant Sci."},{"key":"10.1016\/j.plaphy.2025.110394_bib35","doi-asserted-by":"crossref","first-page":"6957","DOI":"10.1093\/jxb\/erv415","article-title":"Plastid RNA polymerases: orchestration of enzymes with different evolutionary origins controls chloroplast biogenesis during the plant life cycle","volume":"66","author":"Pfannschmidt","year":"2015","journal-title":"J. Exp. Bot."},{"key":"10.1016\/j.plaphy.2025.110394_bib36","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1089\/ars.2005.7.607","article-title":"Redox regulation and modification of proteins controlling chloroplast gene expression","volume":"7","author":"Pfannschmidt","year":"2005","journal-title":"Antioxidants Redox Signal."},{"key":"10.1016\/j.plaphy.2025.110394_bib37","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1016\/j.fcr.2016.09.024","article-title":"Field crops and the fear of heat stress\u2014Opportunities, challenges and future directions","volume":"200","author":"Prasad","year":"2017","journal-title":"Field Crops Res."},{"key":"10.1016\/j.plaphy.2025.110394_bib38","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1038\/nclimate3352","article-title":"Less than 2 \u00b0C warming by 2100 unlikely","volume":"7","author":"Raftery","year":"2017","journal-title":"Nat. Clim. Change"},{"key":"10.1016\/j.plaphy.2025.110394_bib40","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1046\/j.1365-313X.1999.00473.x","article-title":"Targeted disruption of the plastid RNA polymerase genes rpoA, B and C1: molecular biology, biochemistry and ultrastructure","volume":"18","author":"Santis-Maciossek","year":"1999","journal-title":"Plant J."},{"key":"10.1016\/j.plaphy.2025.110394_bib42","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1016\/j.ejcb.2010.06.016","article-title":"Role and regulation of plastid sigma factors and their functional interactors during chloroplast transcription \u2013 recent lessons from Arabidopsis thaliana","volume":"89","author":"Schweer","year":"2010","journal-title":"Eur. J. Cell Biol."},{"key":"10.1016\/j.plaphy.2025.110394_bib43","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0074-7696(05)44001-2","article-title":"Plastid RNA polymerases, promoters, and transcription regulators in higher plants","volume":"244","author":"Shiina","year":"2005","journal-title":"Int. Rev. Cytol."},{"key":"10.1016\/j.plaphy.2025.110394_bib44","doi-asserted-by":"crossref","first-page":"1043","DOI":"10.1104\/pp.111.184515","article-title":"Identification of essential subunits in the plastid-encoded RNA polymerase complex reveals building blocks for proper plastid development","volume":"157","author":"Steiner","year":"2011","journal-title":"Plant Physiology"},{"key":"10.1016\/j.plaphy.2025.110394_bib45","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1016\/j.cell.2024.01.036","article-title":"Structure of the plant plastid-encoded RNA polymerase","volume":"187","author":"Vergara-Cruces","year":"2024","journal-title":"Cell"},{"key":"10.1016\/j.plaphy.2025.110394_bib46","doi-asserted-by":"crossref","first-page":"1433","DOI":"10.1111\/nph.19243","article-title":"Interaction of PALE CRESS with PAP2\/pTAC2 and PAP3\/pTAC10 affects the accumulation of plastid-encoded RNA polymerase complexes in arabidopsis","volume":"240","author":"Wang","year":"2023","journal-title":"New Phytol."},{"key":"10.1016\/j.plaphy.2025.110394_bib47","doi-asserted-by":"crossref","first-page":"2182","DOI":"10.3390\/plants12112182","article-title":"During water stress, fertility modulated by ROS scavengers abundant in arabidopsis pistils","volume":"12","author":"Wang","year":"2023","journal-title":"Plants"},{"key":"10.1016\/j.plaphy.2025.110394_bib48","doi-asserted-by":"crossref","first-page":"8593","DOI":"10.1128\/JVI.00537-06","article-title":"The spread of rice dwarf virus among cells of its insect vector exploits virus-induced tubular structures","volume":"80","author":"Wei","year":"2006","journal-title":"J. Virol."},{"key":"10.1016\/j.plaphy.2025.110394_bib51","doi-asserted-by":"crossref","first-page":"1127","DOI":"10.1016\/j.cell.2024.01.026","article-title":"Cryo-EM structures of the plant plastid-encoded RNA polymerase","volume":"187","author":"Wu","year":"2024","journal-title":"Cell"},{"key":"10.1016\/j.plaphy.2025.110394_bib53","doi-asserted-by":"crossref","first-page":"61.","DOI":"10.3389\/fpls.2014.00061","article-title":"Recent advances in the study of chloroplast gene expression and its evolution","volume":"5","author":"Yagi","year":"2014","journal-title":"Front. Plant Sci."},{"key":"10.1016\/j.plaphy.2025.110394_bib54","article-title":"NCP activates chloroplast transcription by controlling phytochrome-dependent dual nuclear and plastidial switches","volume":"10, 2630","author":"Yang","year":"2019","journal-title":"Nat. Commun."},{"key":"10.1016\/j.plaphy.2025.110394_bib55","doi-asserted-by":"crossref","first-page":"316","DOI":"10.3389\/fpls.2014.00316","article-title":"Nuclear-encoded factors associated with the chloroplast transcription machinery of higher plants","volume":"5","author":"Yu","year":"2014","journal-title":"Front. Plant Sci."},{"key":"10.1016\/j.plaphy.2025.110394_bib56","doi-asserted-by":"crossref","first-page":"641","DOI":"10.1038\/nprot.2006.97","article-title":"Agrobacterium-mediated transformation of Arabidopsis thaliana using the floral dip method","volume":"1","author":"Zhang","year":"2006","journal-title":"Nat. Protoc."},{"key":"10.1016\/j.plaphy.2025.110394_bib57","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1111\/j.1365-313X.2008.03766.x","article-title":"The arabidopsis gene YS1 encoding a DYW protein is required for editing of rpoB transcripts and the rapid development of chloroplasts during early growth","volume":"58","author":"Zhou","year":"2008","journal-title":"Plant J."}],"container-title":["Plant Physiology and Biochemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0981942825009222?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0981942825009222?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,3,7]],"date-time":"2026-03-07T10:34:14Z","timestamp":1772879654000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0981942825009222"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12]]},"references-count":47,"alternative-id":["S0981942825009222"],"URL":"https:\/\/doi.org\/10.1016\/j.plaphy.2025.110394","relation":{},"ISSN":["0981-9428"],"issn-type":[{"value":"0981-9428","type":"print"}],"subject":[],"published":{"date-parts":[[2025,12]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Thermal protection of the plastid-encoded RNA polymerase complex by PAP9 ensures chloroplast function in Arabidopsis","name":"articletitle","label":"Article Title"},{"value":"Plant Physiology and Biochemistry","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.plaphy.2025.110394","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2025 Elsevier Masson SAS. All rights are reserved, including those for text and data mining, AI training, and similar technologies.","name":"copyright","label":"Copyright"}],"article-number":"110394"}}