{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T19:59:21Z","timestamp":1787774361992,"version":"build-2784847793"},"reference-count":76,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2025,8,5]],"date-time":"2025-08-05T00:00:00Z","timestamp":1754352000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2025,8,5]],"date-time":"2025-08-05T00:00:00Z","timestamp":1754352000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["22K06081"],"award-info":[{"award-number":["22K06081"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["21H02401"],"award-info":[{"award-number":["21H02401"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["21H05275"],"award-info":[{"award-number":["21H05275"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100016925","name":"Takeda Foundation","doi-asserted-by":"publisher","award":["J241503007"],"award-info":[{"award-number":["J241503007"]}],"id":[{"id":"10.13039\/100016925","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012630","name":"Life Science Foundation of Japan","doi-asserted-by":"publisher","award":["J231503018"],"award-info":[{"award-number":["J231503018"]}],"id":[{"id":"10.13039\/501100012630","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007428","name":"Naito Foundation","doi-asserted-by":"publisher","award":["J241503011"],"award-info":[{"award-number":["J241503011"]}],"id":[{"id":"10.13039\/100007428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100005927","name":"Daiichi Sankyo Foundation of Life Science","doi-asserted-by":"publisher","award":["J241503010"],"award-info":[{"award-number":["J241503010"]}],"id":[{"id":"10.13039\/501100005927","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Genes to Cells"],"published-print":{"date-parts":[[2025,9]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Testis\u2010specific gene regulatory mechanisms govern the differentiation of germ cells into mature sperm. However, the molecular underpinnings are not fully elucidated. Here, we show the subunits forming Paf1C, a transcription regulator complex conserved across eukaryotes, have their individual paralogs predominantly expressed in\n                    <jats:italic>Drosophila<\/jats:italic>\n                    testes. One of these, namely, Ctr9 paralog enriched in testes (Ctr9t) was found to play a critical and nonredundant role in postmeiotic spermatid differentiation and male fertility in\n                    <jats:styled-content style=\"fixed-case\">\n                      <jats:italic>D. melanogaster<\/jats:italic>\n                    <\/jats:styled-content>\n                    . A proximity proteome analysis provides evidence that Ctr9t prefers the interaction between paralog members. We show endogenous Ctr9t is expressed and functional in germ cells at spermatocyte stages, accumulating in a distinct compartment within the nucleolus. There, Ctr9t co\u2010localizes with Spermatocyte arrest (Sa), a testis\u2010specific paralog of TATA\u2010binding protein (TBP)\u2010associated factor 8 (TAF8). We further demonstrate that\n                    <jats:italic>ctr9t<\/jats:italic>\n                    function is crucial for maintaining Sa in the nucleolus, but not vice versa. Transcriptome profiling reveals that Ctr9t acts as an activator for the set of male fertility genes on the Y chromosome, but it also acts as a global repressor of X chromosome genes. Collectively, our results shed light on the nucleolus\u2010associated, paralog\u2010dependent regulation of gene expression from sex chromosomes, which ensures the terminal differentiation of male germ cells.\n                  <\/jats:p>","DOI":"10.1111\/gtc.70040","type":"journal-article","created":{"date-parts":[[2025,8,6]],"date-time":"2025-08-06T01:00:03Z","timestamp":1754442003000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Paralog\u2010Dependent Specialization of\n                    <scp>Paf1C<\/scp>\n                    Subunit, Ctr9, for Sex Chromosome Gene Regulation and Male Germline Differentiation in Drosophila"],"prefix":"10.1111","volume":"30","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8675-8469","authenticated-orcid":false,"given":"Toshie","family":"Kai","sequence":"first","affiliation":[{"name":"Laboratory of Germline Biology, Graduate School of Frontier Biosciences Osaka University  Osaka Japan"},{"name":"Graduate School of Biostudies Kyoto University  Kyoto Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinglan","family":"Zheng","sequence":"additional","affiliation":[{"name":"Laboratory of Germline Biology, Graduate School of Frontier Biosciences Osaka University  Osaka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3215-4552","authenticated-orcid":false,"given":"Taichiro","family":"Iki","sequence":"additional","affiliation":[{"name":"Laboratory of Germline Biology, Graduate School of Frontier Biosciences Osaka University  Osaka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2025,8,5]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-34295-5"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1128\/MCB.26.1.250-260.2006"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.091553"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.89373"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1534\/g3.116.034231"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.1516607"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-29146-2"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrm2184"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1093\/genetics\/120.4.1015"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1038\/nbt.4201"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/evo.13229"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1534\/genetics.118.301765"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1534\/g3.116.035196"},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1118101"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.056572"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2017.07.052"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M115.646315"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ydbio.2011.04.030"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1003842"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.4161\/spmg.21798"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.349969.122"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1008028"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jmb.2021.166979"},{"key":"e_1_2_9_25_1","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2148-7-87"},{"key":"e_1_2_9_26_1","first-page":"E9224","article-title":"PAF1 Complex Component Leo1 Helps Recruit Drosophila Myc to Promoters","volume":"114","author":"Gerlach J. M.","year":"2017","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1534\/g3.114.014126"},{"key":"e_1_2_9_28_1","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-023-05717-x"},{"key":"e_1_2_9_29_1","doi-asserted-by":"publisher","DOI":"10.1534\/genetics.120.302672"},{"key":"e_1_2_9_30_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF02524646"},{"key":"e_1_2_9_31_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.0050273"},{"key":"e_1_2_9_32_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.01314"},{"key":"e_1_2_9_33_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.869101"},{"key":"e_1_2_9_34_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0213177"},{"key":"e_1_2_9_35_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tcb.2019.04.003"},{"key":"e_1_2_9_36_1","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.adh0397"},{"key":"e_1_2_9_37_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2020.108380"},{"key":"e_1_2_9_38_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbagrm.2010.01.001"},{"key":"e_1_2_9_39_1","doi-asserted-by":"publisher","DOI":"10.1093\/bib\/bbx108"},{"key":"e_1_2_9_40_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msq355"},{"key":"e_1_2_9_41_1","doi-asserted-by":"publisher","DOI":"10.1098\/rsob.140145"},{"key":"e_1_2_9_42_1","doi-asserted-by":"publisher","DOI":"10.3390\/ijms23084184"},{"key":"e_1_2_9_43_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature14337"},{"key":"e_1_2_9_44_1","doi-asserted-by":"publisher","DOI":"10.1186\/s13072-018-0183-3"},{"key":"e_1_2_9_45_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msu002"},{"key":"e_1_2_9_46_1","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkx976"},{"key":"e_1_2_9_47_1","doi-asserted-by":"publisher","DOI":"10.1242\/bio.20121255"},{"key":"e_1_2_9_48_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msp030"},{"key":"e_1_2_9_49_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.69.1.182"},{"key":"e_1_2_9_50_1","doi-asserted-by":"publisher","DOI":"10.4161\/spmg.21775"},{"key":"e_1_2_9_51_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.122.4.1331"},{"key":"e_1_2_9_52_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1005701"},{"key":"e_1_2_9_53_1","doi-asserted-by":"publisher","DOI":"10.1080\/15384101.2014.998070"},{"key":"e_1_2_9_54_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-20897-y"},{"key":"e_1_2_9_55_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1001126"},{"key":"e_1_2_9_56_1","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.21294"},{"key":"e_1_2_9_57_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.056945"},{"key":"e_1_2_9_58_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2006.01.053"},{"key":"e_1_2_9_59_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.mod.2009.02.002"},{"key":"e_1_2_9_60_1","doi-asserted-by":"publisher","DOI":"10.1089\/cmb.2022.0121"},{"key":"e_1_2_9_61_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2010.02.046"},{"key":"e_1_2_9_62_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-36021-1"},{"key":"e_1_2_9_63_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.01032"},{"key":"e_1_2_9_64_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbagrm.2013.08.004"},{"key":"e_1_2_9_65_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00412-011-0321-1"},{"key":"e_1_2_9_66_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0925-4773(98)00157-9"},{"key":"e_1_2_9_67_1","doi-asserted-by":"publisher","DOI":"10.1006\/dbio.1995.8049"},{"key":"e_1_2_9_68_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0603620103"},{"key":"e_1_2_9_69_1","first-page":"439","article-title":"TBRD\u20101 and tBRD\u20102 Regulate Expression of Genes Necessary for Spermatid Differentiation","volume":"6","author":"Theofel I.","year":"2017","journal-title":"Biology Open"},{"key":"e_1_2_9_70_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0108267"},{"key":"e_1_2_9_71_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1000731"},{"key":"e_1_2_9_72_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.351930.124"},{"key":"e_1_2_9_73_1","doi-asserted-by":"publisher","DOI":"10.1530\/REP-09-0083"},{"key":"e_1_2_9_74_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1009728"},{"key":"e_1_2_9_75_1","doi-asserted-by":"publisher","DOI":"10.1534\/genetics.120.303324"},{"key":"e_1_2_9_76_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2007.03.036"},{"key":"e_1_2_9_77_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1009655"}],"container-title":["Genes to Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/gtc.70040","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/gtc.70040","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/gtc.70040","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,9]],"date-time":"2026-03-09T02:54:33Z","timestamp":1773024873000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/gtc.70040"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,5]]},"references-count":76,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2025,9]]}},"alternative-id":["10.1111\/gtc.70040"],"URL":"https:\/\/doi.org\/10.1111\/gtc.70040","archive":["Portico"],"relation":{},"ISSN":["1356-9597","1365-2443"],"issn-type":[{"value":"1356-9597","type":"print"},{"value":"1365-2443","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,5]]},"assertion":[{"value":"2025-06-27","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2025-07-23","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2025-08-05","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70040"}}