{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,19]],"date-time":"2026-06-19T11:46:41Z","timestamp":1781869601454,"version":"3.54.5"},"reference-count":80,"publisher":"Oxford University Press (OUP)","issue":"4","license":[{"start":{"date-parts":[[2026,2,10]],"date-time":"2026-02-10T00:00:00Z","timestamp":1770681600000},"content-version":"vor","delay-in-days":5,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2024YFA1108100"],"award-info":[{"award-number":["2024YFA1108100"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82371625"],"award-info":[{"award-number":["82371625"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82302099"],"award-info":[{"award-number":["82302099"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002858","name":"China Postdoctoral Science Foundation","doi-asserted-by":"publisher","award":["2023M742727"],"award-info":[{"award-number":["2023M742727"]}],"id":[{"id":"10.13039\/501100002858","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100016804","name":"Shenzhen Natural Science Foundation","doi-asserted-by":"publisher","award":["JCYJ20240813153314019"],"award-info":[{"award-number":["JCYJ20240813153314019"]}],"id":[{"id":"10.13039\/100016804","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100021171","name":"Guangdong Basic and Applied Basic Research Foundation","doi-asserted-by":"publisher","award":["2024A1515012936"],"award-info":[{"award-number":["2024A1515012936"]}],"id":[{"id":"10.13039\/501100021171","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Hubei Provincial Laboratory Animal Research Special Fund","award":["2023CFA006"],"award-info":[{"award-number":["2023CFA006"]}]},{"name":"Wuhan Municipal Health Commission","award":["WX23A47"],"award-info":[{"award-number":["WX23A47"]}]},{"DOI":"10.13039\/501100003397","name":"Huazhong University of Science and Technology","doi-asserted-by":"publisher","award":["2023BR031"],"award-info":[{"award-number":["2023BR031"]}],"id":[{"id":"10.13039\/501100003397","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,2,5]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The process of male germ cell development is a central determinant of spermatogenesis. Nevertheless, the genetic regulatory mechanisms underlying male germ cell development in mammals remain largely unclear. In this study, employing a germ cell-specific Hnrnpk knockout mouse model combined with multi-omics analyses, we identified hnRNPK as a key factor necessary for maintaining normal development in differentiating spermatogonia. Phenotypically, adult mice with germ cell-specific hnRNPK deletion exhibited infertility, characterized by a near-complete absence of spermatocytes in the seminiferous tubules. Single-cell RNA sequencing (scRNA-seq) analysis revealed that hnRNPK deletion induced cell-cycle dysregulation in differentiating spermatogonia, triggering apoptotic cell death. As a consequence, the population of differentiating spermatogonia in the testes is markedly diminished, and these cells fail to undergo proper maturation or successfully enter meiosis. Mechanistically, cytoplasmic hnRNPK exerts its regulatory function at the post-transcriptional level, regulating the translation efficiency (TE) of genes involved in meiosis, the cell cycle, and transcriptional regulation. Furthermore, hnRNPK interacts with and colocalizes with DAZL at the 40S ribosome, thereby modulating the initiation of target messenger RNA translation. In the nucleus, hnRNPK interacts with splicing factors and participates in the splicing of target genes related to germ cell differentiation and meiosis. Collectively, these findings emphasize the functional role and mechanistic involvement of hnRNPK in differentiating spermatogonia, providing valuable insights into the post-transcriptional regulatory mechanisms that govern male germ cell development.<\/jats:p>","DOI":"10.1093\/nar\/gkag108","type":"journal-article","created":{"date-parts":[[2026,2,10]],"date-time":"2026-02-10T09:38:18Z","timestamp":1770716298000},"source":"Crossref","is-referenced-by-count":2,"title":["Azoospermia phenotype and scRNA-seq reveal hnRNPK as a factor essential for male germ cell development in mice"],"prefix":"10.1093","volume":"54","author":[{"given":"Huihui","family":"Gao","sequence":"first","affiliation":[{"name":"Department of Obstetrics and Gynecology, The Central Hospital of Wuhan, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]},{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shiyu","family":"Yang","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ao","family":"Ning","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Sichuan Agricultural University Sichuan , Chengdu 625014 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lisha","family":"Yin","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yifei","family":"Lan","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Keren","family":"Cheng","sequence":"additional","affiliation":[{"name":"Center for Reproductive Medicine, The Fourth Affiliated Hospital, Zhejiang University School of Medicine , Yiwu 322000 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenjing","family":"Xiong","sequence":"additional","affiliation":[{"name":"Laboratory Animal Center, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinxin","family":"Xiong","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jin","family":"Zhang","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingshou","family":"Chen","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shenglei","family":"Feng","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xu","family":"Fan","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kuan","family":"Liu","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yiqian","family":"Gui","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peng","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology, The Central Hospital of Wuhan, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoli","family":"Wang","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fengli","family":"Wang","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoxu","family":"Chen","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Northwest A&F University , Yangling 712100 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qinghua","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology, The Central Hospital of Wuhan, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1460-7682","authenticated-orcid":false,"given":"Shuiqiao","family":"Yuan","sequence":"additional","affiliation":[{"name":"Institute of Reproductive Health, Tongji Medical College, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]},{"name":"Laboratory Animal Center, Huazhong University of Science and Technology , Wuhan 430030 ,","place":["China"]},{"name":"Shenzhen Huazhong University of Science and Technology Research Institute , Shenzhen 518057 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,2,10]]},"reference":[{"key":"2026021004381033000_B1","doi-asserted-by":"publisher","first-page":"339","DOI":"10.1634\/stemcells.2007-0622","article-title":"Dynamic regulation of mitotic arrest in fetal male germ cells","volume":"26","author":"Western","year":"2008","journal-title":"Stem Cells"},{"key":"2026021004381033000_B2","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1530\/REP-10-0431","article-title":"Spermatogonial morphology and kinetics during testis development in mice: a high-resolution light microscopy approach","volume":"142","author":"Drumond","year":"2011","journal-title":"Reproduction"},{"key":"2026021004381033000_B3","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1095\/biolreprod.113.114645","article-title":"Retinoic acid induces multiple hallmarks of the prospermatogonia-to-spermatogonia transition in the neonatal mouse","volume":"90","author":"Busada","year":"2014","journal-title":"Biol Reprod"},{"key":"2026021004381033000_B4","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1530\/REP-14-0653","article-title":"Marker expression reveals heterogeneity of spermatogonia in the neonatal mouse testis","volume":"149","author":"Niedenberger","year":"2015","journal-title":"Reproduction"},{"key":"2026021004381033000_B5","doi-asserted-by":"publisher","first-page":"783","DOI":"10.1095\/biolreprod.110.085811","article-title":"Retinoic acid availability drives the asynchronous initiation of spermatogonial differentiation in the mouse","volume":"83","author":"Snyder","year":"2010","journal-title":"Biol Reprod"},{"key":"2026021004381033000_B6","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1095\/biolreprod.107.066795","article-title":"Expression of stimulated by retinoic acid gene 8 (Stra8) in spermatogenic cells induced by retinoic acid: an in vivo study in vitamin A-sufficient postnatal murine testes","volume":"79","author":"Zhou","year":"2008","journal-title":"Biol Reprod"},{"key":"2026021004381033000_B7","doi-asserted-by":"publisher","first-page":"5894","DOI":"10.1210\/endo.140.12.7172","article-title":"Differential expression of c-kit in mouse undifferentiated and differentiating type A spermatogonia","volume":"140","author":"Schrans-Stassen","year":"1999","journal-title":"Endocrinology"},{"key":"2026021004381033000_B8","doi-asserted-by":"publisher","first-page":"e979623","DOI":"10.4161\/21565562.2014.979623","article-title":"Mechanisms of spermiogenesis and spermiation and how they are disturbed","volume":"4","author":"O\u2019Donnell","year":"2014","journal-title":"Spermatogenesis"},{"key":"2026021004381033000_B9","doi-asserted-by":"publisher","first-page":"743","DOI":"10.1530\/REP-08-0537","article-title":"Control of KIT signalling in male germ cells: what can we learn from other systems?","volume":"138","author":"Mithraprabhu","year":"2009","journal-title":"Reproduction"},{"key":"2026021004381033000_B10","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1111\/j.1365-2605.2005.00614.x","article-title":"Mouse models for genes involved in impaired spermatogenesis","volume":"29","author":"O\u2019Bryan","year":"2006","journal-title":"Int J of Andrology"},{"key":"2026021004381033000_B11","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-031-31574-9","volume-title":"Andrology: Male Reproductive Health and Dysfunction","author":"Nieschlag","year":"2023"},{"key":"2026021004381033000_B12","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1146\/annurev.bi.62.070193.001445","article-title":"hnRNP proteins and the biogenesis of mRNA","volume":"62","author":"Dreyfuss","year":"1993","journal-title":"Annu Rev Biochem"},{"key":"2026021004381033000_B13","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1042\/BJ20100396","article-title":"Functional diversity of the hnRNPs: past, present and perspectives","volume":"430","author":"Han","year":"2010","journal-title":"Biochem J"},{"key":"2026021004381033000_B14","doi-asserted-by":"publisher","first-page":"314","DOI":"10.4103\/aja202273","article-title":"Characterization of the protein expression and localization of hnRNP family members during murine spermatogenesis","volume":"25","author":"Wang","year":"2023","journal-title":"Asian J Androl"},{"key":"2026021004381033000_B15","doi-asserted-by":"publisher","first-page":"2803","DOI":"10.1093\/hmg\/ddn179","article-title":"Haploinsufficiency of the germ cell-specific nuclear RNA binding protein hnRNP G-T prevents functional spermatogenesis in the mouse","volume":"17","author":"Ehrmann","year":"2008","journal-title":"Hum Mol Genet"},{"key":"2026021004381033000_B16","doi-asserted-by":"publisher","first-page":"2879","DOI":"10.1016\/j.jprot.2011.12.040","article-title":"HnRNPL as a key factor in spermatogenesis: lesson from functional proteomic studies of azoospermia patients with sertoli cell only syndrome","volume":"75","author":"Li","year":"2012","journal-title":"J Proteomics"},{"key":"2026021004381033000_B17","doi-asserted-by":"publisher","first-page":"3588","DOI":"10.1038\/s41467-022-31364-7","article-title":"hnRNPH1 recruits PTBP2 and SRSF3 to modulate alternative splicing in germ cells","volume":"13","author":"Feng","year":"2022","journal-title":"Nat Commun"},{"key":"2026021004381033000_B18","doi-asserted-by":"publisher","first-page":"114113","DOI":"10.1016\/j.celrep.2024.114113","article-title":"hnRNPU is required for spermatogonial stem cell pool establishment in mice","volume":"43","author":"Wen","year":"2024","journal-title":"Cell Rep"},{"key":"2026021004381033000_B19","doi-asserted-by":"publisher","first-page":"113769","DOI":"10.1016\/j.celrep.2024.113769","article-title":"hnRNPA2B1 represses the disassembly of arsenite-induced stress granules and is essential for male fertility","volume":"43","author":"Wang","year":"2024","journal-title":"Cell Rep"},{"key":"2026021004381033000_B20","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1186\/s12958-024-01340-5","article-title":"HnRNPM modulates alternative splicing in germ cells by recruiting PTBP1","volume":"23","author":"Lv","year":"2025","journal-title":"Reprod Biol Endocrinol"},{"key":"2026021004381033000_B21","doi-asserted-by":"publisher","first-page":"e2412196","DOI":"10.1002\/advs.202412196","article-title":"hnRNPC functions with HuR to regulate alternative splicing in an m6A-dependent manner and is essential for meiosis","volume":"12","author":"Xiong","year":"2025","journal-title":"Adv Sci"},{"key":"2026021004381033000_B22","doi-asserted-by":"publisher","first-page":"dev201040","DOI":"10.1242\/dev.201040","article-title":"hnRNPH1 establishes Sertoli\u2013germ cell crosstalk through cooperation with PTBP1 and AR, and is essential for male fertility in mice","volume":"150","author":"Feng","year":"2023","journal-title":"Development"},{"key":"2026021004381033000_B23","doi-asserted-by":"publisher","first-page":"2712","DOI":"10.1111\/j.1742-4658.2008.06411.x","article-title":"Structure and function of KH domains","volume":"275","author":"Valverde","year":"2008","journal-title":"FEBS J"},{"key":"2026021004381033000_B24","doi-asserted-by":"publisher","first-page":"1995","DOI":"10.1002\/jcp.29186","article-title":"The emerging roles of hnRNPK","volume":"235","author":"Wang","year":"2020","journal-title":"J Cell Physioyl"},{"key":"2026021004381033000_B25","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1016\/j.ccell.2015.09.001","article-title":"hnRNP K is a haploinsufficient tumor suppressor that regulates proliferation and differentiation programs in hematologic malignancies","volume":"28","author":"Gallardo","year":"2015","journal-title":"Cancer Cell"},{"key":"2026021004381033000_B26","doi-asserted-by":"publisher","first-page":"2350","DOI":"10.1128\/MCB.16.5.2350","article-title":"Heterogeneous nuclear ribonucleoprotein K is a transcription factor","volume":"16","author":"Michelotti","year":"1996","journal-title":"Mol Cell Biol"},{"key":"2026021004381033000_B27","doi-asserted-by":"publisher","first-page":"18249","DOI":"10.1016\/S0021-9258(17)46837-2","article-title":"Specific binding of heterogeneous ribonucleoprotein particle protein K to the human c-myc promoter, in vitro","volume":"268","author":"Takimoto","year":"1993","journal-title":"J Biol Chem"},{"key":"2026021004381033000_B28","doi-asserted-by":"publisher","first-page":"1502","DOI":"10.1093\/nar\/gkg246","article-title":"Identification of the SRC pyrimidine-binding protein (SPy) as hnRNP K: implications in the regulation of SRC1A transcription","volume":"31","author":"Ritchie","year":"2003","journal-title":"Nucleic Acids Res"},{"key":"2026021004381033000_B29","doi-asserted-by":"publisher","first-page":"8012","DOI":"10.1038\/sj.onc.1206645","article-title":"Members of the poly (rC) binding protein family stimulate the activity of the c-myc internal ribosome entry segment in vitro and in vivo","volume":"22","author":"Evans","year":"2003","journal-title":"Oncogene"},{"key":"2026021004381033000_B30","doi-asserted-by":"publisher","first-page":"180239","DOI":"10.1098\/rsob.180239","article-title":"Post-translational modification control of RNA-binding protein hnRNPK function","volume":"9","author":"Xu","year":"2019","journal-title":"Open Biol"},{"key":"2026021004381033000_B31","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1186\/s12935-019-1020-x","article-title":"hnRNPK promotes gastric tumorigenesis through regulating CD44E alternative splicing","volume":"19","author":"Peng","year":"2019","journal-title":"Cancer Cell Int"},{"key":"2026021004381033000_B32","doi-asserted-by":"publisher","first-page":"1006","DOI":"10.1111\/gtc.12400","article-title":"Control of the heat stress-induced alternative splicing of a subset of genes by hnRNP K","volume":"21","author":"Yamamoto","year":"2016","journal-title":"Genes Cells"},{"key":"2026021004381033000_B33","doi-asserted-by":"publisher","first-page":"1479","DOI":"10.1177\/15353702221110649","article-title":"HnRNP K regulates inflammatory gene expression by mediating splicing pattern of transcriptional factors","volume":"248","author":"Liu","year":"2023","journal-title":"Exp Biol Med (Maywood)"},{"key":"2026021004381033000_B34","doi-asserted-by":"crossref","first-page":"1277","DOI":"10.3390\/cells11081277","article-title":"Deletion of Hnrnpk gene causes infertility in male mice by disrupting spermatogenesis","volume":"11","author":"Xu","year":"2022","journal-title":"Cells"},{"key":"2026021004381033000_B35","doi-asserted-by":"publisher","first-page":"6438","DOI":"10.1038\/s41598-025-91081-1","article-title":"Insights into the role of hnRNPK in spermatogenesis via the piRNA pathway","volume":"15","author":"Xu","year":"2025","journal-title":"Sci Rep"},{"key":"2026021004381033000_B36","doi-asserted-by":"publisher","first-page":"1216","DOI":"10.1038\/cr.2017.117","article-title":"Mettl3-\/Mettl14-mediated mRNA N6-methyladenosine modulates murine spermatogenesis","volume":"27","author":"Lin","year":"2017","journal-title":"Cell Res"},{"key":"2026021004381033000_B37","first-page":"1331","article-title":"CHD4 acts as a critical regulator in the survival of spermatogonial stem cells in micedagger","volume":"107","author":"Li","year":"2022","journal-title":"Biol Reprod"},{"key":"2026021004381033000_B38","doi-asserted-by":"publisher","first-page":"pdb.prot094185","DOI":"10.1101\/pdb.prot094185","article-title":"Isolation of the spermatogonial stem cell-containing fraction from testes","volume":"2017","author":"Goodyear","year":"2017","journal-title":"Cold Spring Harb Protoc"},{"key":"2026021004381033000_B39","doi-asserted-by":"publisher","first-page":"1523","DOI":"10.1038\/s41467-019-09234-6","article-title":"Metascape provides a biologist-oriented resource for the analysis of systems-level datasets","volume":"10","author":"Zhou","year":"2019","journal-title":"Nat Commun"},{"key":"2026021004381033000_B40","doi-asserted-by":"publisher","first-page":"E5593","DOI":"10.1073\/pnas.1419161111","article-title":"rMATS: robust and flexible detection of differential alternative splicing from replicate RNA-seq data","volume":"111","author":"Shen","year":"2014","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026021004381033000_B41","doi-asserted-by":"publisher","first-page":"e61","DOI":"10.1093\/nar\/gkr1291","article-title":"MATS: a Bayesian framework for flexible detection of differential alternative splicing from RNA-seq data","volume":"40","author":"Shen","year":"2012","journal-title":"Nucleic Acids Res"},{"key":"2026021004381033000_B42","doi-asserted-by":"publisher","first-page":"879","DOI":"10.1038\/s41422-018-0074-y","article-title":"Single-cell RNA-seq uncovers dynamic processes and critical regulators in mouse spermatogenesis","volume":"28","author":"Chen","year":"2018","journal-title":"Cell Res"},{"key":"2026021004381033000_B43","doi-asserted-by":"publisher","first-page":"101282","DOI":"10.1016\/j.mam.2024.101282","article-title":"Mechanisms of meiosis initiation and meiotic prophase progression during spermatogenesis","volume":"97","author":"Ishiguro","year":"2024","journal-title":"Mol Aspects Med"},{"key":"2026021004381033000_B44","doi-asserted-by":"publisher","first-page":"11590","DOI":"10.1074\/jbc.M209233200","article-title":"Synthesis and functional analyses of nuclear clusterin, a cell death protein","volume":"278","author":"Leskov","year":"2003","journal-title":"J Biol Chem"},{"key":"2026021004381033000_B45","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1007\/s10735-006-9052-7","article-title":"Clusterin and DNA repair: a new function in cancer for a key player in apoptosis and cell cycle control","volume":"37","author":"Shannan","year":"2006","journal-title":"J Mol Hist"},{"key":"2026021004381033000_B46","doi-asserted-by":"publisher","first-page":"110307","DOI":"10.1016\/j.cellsig.2022.110307","article-title":"GPx8 regulates apoptosis and autophagy in esophageal squamous cell carcinoma through the IRE1\/JNK pathway","volume":"93","author":"Yin","year":"2022","journal-title":"Cell Signalling"},{"key":"2026021004381033000_B47","doi-asserted-by":"publisher","first-page":"1695","DOI":"10.1093\/emboj\/16.7.1695","article-title":"Induction of apoptosis by the transcription factor c-Jun","volume":"16","author":"Bossy-Wetzel","year":"1997","journal-title":"EMBO J"},{"key":"2026021004381033000_B48","doi-asserted-by":"publisher","first-page":"6537","DOI":"10.1128\/.20.17.6537-6549.2000","article-title":"A bromodomain protein, MCAP, associates with mitotic chromosomes and affects G(2)-to-M transition","volume":"20","author":"Dey","year":"2000","journal-title":"Mol Cell Biol"},{"key":"2026021004381033000_B49","doi-asserted-by":"publisher","first-page":"609","DOI":"10.1007\/BF03343784","article-title":"Role of c-kit in mammalian spermatogenesis","volume":"23","author":"Rossi","year":"2000","journal-title":"J Endocrinol Invest"},{"key":"2026021004381033000_B50","doi-asserted-by":"publisher","first-page":"1133","DOI":"10.1242\/dev.073478","article-title":"An essential role for a mammalian SWI\/SNF chromatin-remodeling complex during male meiosis","volume":"139","author":"Kim","year":"2012","journal-title":"Development"},{"key":"2026021004381033000_B51","doi-asserted-by":"publisher","first-page":"1598","DOI":"10.1002\/2211-5463.12304","article-title":"The transcription factor ATF7 mediates in vitro fertilization-induced gene expression changes in mouse liver","volume":"7","author":"Liu","year":"2017","journal-title":"FEBS Open Bio"},{"key":"2026021004381033000_B52","doi-asserted-by":"publisher","first-page":"104293","DOI":"10.1016\/j.yexmp.2019.104293","article-title":"Wnt suppressor and stem cell regulator TCF7L1 is a sensitive immunohistochemical marker to differentiate testicular seminoma from non-seminomatous germ cell tumor","volume":"110","author":"Bu","year":"2019","journal-title":"Exp Mol Pathol"},{"key":"2026021004381033000_B53","doi-asserted-by":"publisher","first-page":"1225","DOI":"10.1016\/j.celrep.2018.10.012","article-title":"DAZL regulates germ cell survival through a network of PolyA-proximal mRNA interactions","volume":"25","author":"Zagore","year":"2018","journal-title":"Cell Rep"},{"key":"2026021004381033000_B54","doi-asserted-by":"publisher","first-page":"e56523","DOI":"10.7554\/eLife.56523","article-title":"DAZL mediates a broad translational program regulating expansion and differentiation of spermatogonial progenitors","volume":"9","author":"Mikedis","year":"2020","journal-title":"eLife"},{"key":"2026021004381033000_B55","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1093\/nsr\/nwy163","article-title":"DAZL is a master translational regulator of murine spermatogenesis","volume":"6","author":"Li","year":"2019","journal-title":"Natl Sci Rev"},{"key":"2026021004381033000_B56","doi-asserted-by":"publisher","first-page":"1276","DOI":"10.1038\/s41418-017-0038-7","article-title":"BRAP-2 promotes DNA damage induced germline apoptosis in C. elegans through the regulation of SKN-1 and AKT-1","volume":"25","author":"D\u2019Amora","year":"2018","journal-title":"Cell Death Differ"},{"key":"2026021004381033000_B57","doi-asserted-by":"publisher","first-page":"25572","DOI":"10.1074\/jbc.M002218200","article-title":"Stem cell factor\/c-kit up-regulates cyclin D3 and promotes cell cycle progression via the phosphoinositide 3-kinase\/p70 S6 kinase pathway in spermatogonia","volume":"275","author":"Feng","year":"2000","journal-title":"J Biol Chem"},{"key":"2026021004381033000_B58","doi-asserted-by":"publisher","first-page":"102905","DOI":"10.1016\/j.jbc.2023.102905","article-title":"DNA repair protein FANCD2 has both ubiquitination-dependent and ubiquitination-independent functions during germ cell development","volume":"299","author":"Zhao","year":"2023","journal-title":"J Biol Chem"},{"key":"2026021004381033000_B59","doi-asserted-by":"publisher","first-page":"633","DOI":"10.1136\/jmedgenet-2017-104514","article-title":"Mutation in TDRD9 causes non-obstructive azoospermia in infertile men","volume":"54","author":"Arafat","year":"2017","journal-title":"J Med Genet"},{"key":"2026021004381033000_B60","doi-asserted-by":"publisher","first-page":"775","DOI":"10.1016\/j.devcel.2009.10.012","article-title":"The TDRD9\u2013MIWI2 complex is essential for piRNA-mediated retrotransposon silencing in the mouse male germline","volume":"17","author":"Shoji","year":"2009","journal-title":"Dev Cell"},{"key":"2026021004381033000_B61","doi-asserted-by":"publisher","first-page":"e1004954","DOI":"10.1371\/journal.pgen.1004954","article-title":"Polycomb protein SCML2 associates with USP7 and counteracts histone H2A ubiquitination in the XY chromatin during male meiosis","volume":"11","author":"Luo","year":"2015","journal-title":"PLoS Genet"},{"key":"2026021004381033000_B62","doi-asserted-by":"publisher","first-page":"3233","DOI":"10.1242\/jcs.035071","article-title":"STRA8-deficient spermatocytes initiate, but fail to complete, meiosis and undergo premature chromosome condensation","volume":"121","author":"Mark","year":"2008","journal-title":"J Cell Sci"},{"key":"2026021004381033000_B63","doi-asserted-by":"publisher","first-page":"4982","DOI":"10.1073\/pnas.0505123103","article-title":"TEX14 is essential for intercellular bridges and fertility in male mice","volume":"103","author":"Greenbaum","year":"2006","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026021004381033000_B64","doi-asserted-by":"publisher","first-page":"744","DOI":"10.1126\/science.1142612","article-title":"Developmentally regulated piRNA clusters implicate MILI in transposon control","volume":"316","author":"Aravin","year":"2007","journal-title":"Science"},{"key":"2026021004381033000_B65","doi-asserted-by":"publisher","first-page":"3003","DOI":"10.1091\/mbc.E18-07-0459","article-title":"Depletion of SMC5\/6 sensitizes male germ cells to DNA damage","volume":"29","author":"Hwang","year":"2018","journal-title":"MBoC"},{"key":"2026021004381033000_B66","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1002\/cyto.a.23898","article-title":"Apoptotic cell exclusion and bias-free single-cell selection are important quality control requirements for successful single-cell sequencing applications","volume":"97","author":"Ordonez-Rueda","year":"2020","journal-title":"Cytometry Pt A"},{"key":"2026021004381033000_B67","doi-asserted-by":"publisher","first-page":"1251","DOI":"10.1038\/s41467-019-09182-1","article-title":"Staged developmental mapping and X chromosome transcriptional dynamics during mouse spermatogenesis","volume":"10","author":"Ernst","year":"2019","journal-title":"Nat Commun"},{"key":"2026021004381033000_B68","first-page":"473","article-title":"ERK1\/2-mediated cytoplasmic accumulation of hnRNPK antagonizes TRAIL-induced apoptosis through upregulation of XIAP in H1299 cells","volume":"30","author":"Huang","year":"2017","journal-title":"Biomed Environ Sci"},{"key":"2026021004381033000_B69","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1038\/35060131","article-title":"ERK phosphorylation drives cytoplasmic accumulation of hnRNP-K and inhibition of mRNA translation","volume":"3","author":"Habelhah","year":"2001","journal-title":"Nat Cell Biol"},{"key":"2026021004381033000_B70","doi-asserted-by":"publisher","first-page":"338","DOI":"10.1002\/1878-0261.12406","article-title":"O-GlcNAc-induced nuclear translocation of hnRNP-K is associated with progression and metastasis of cholangiocarcinoma","volume":"13","author":"Phoomak","year":"2019","journal-title":"Mol Oncol"},{"key":"2026021004381033000_B71","doi-asserted-by":"publisher","first-page":"865","DOI":"10.1016\/j.bbrc.2010.12.076","article-title":"Identification of the methylation preference region in heterogeneous nuclear ribonucleoprotein K by protein arginine methyltransferase 1 and its implication in regulating nuclear\/cytoplasmic distribution","volume":"404","author":"Chang","year":"2011","journal-title":"Biochem Biophys Res Commun"},{"key":"2026021004381033000_B72","doi-asserted-by":"publisher","first-page":"8551","DOI":"10.1073\/pnas.0402889101","article-title":"Sumoylation of heterogeneous nuclear ribonucleoproteins, zinc finger proteins, and nuclear pore complex proteins: a proteomic analysis","volume":"101","author":"Li","year":"2004","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026021004381033000_B73","doi-asserted-by":"publisher","first-page":"851","DOI":"10.1007\/s00439-016-1683-5","article-title":"The hnRNP family: insights into their role in health and disease","volume":"135","author":"Geuens","year":"2016","journal-title":"Hum Genet"},{"key":"2026021004381033000_B74","doi-asserted-by":"publisher","first-page":"626","DOI":"10.1038\/nrg1125","article-title":"The power of the 3\u2032 UTR: translational control and development","volume":"4","author":"Kuersten","year":"2003","journal-title":"Nat Rev Genet"},{"key":"2026021004381033000_B75","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/j.semcdb.2004.11.007","article-title":"Mechanisms of translational control by the 3' UTR in development and differentiation","volume":"16","author":"de\u00a0Moor","year":"2005","journal-title":"Semin Cell Dev Biol"},{"key":"2026021004381033000_B76","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1093\/biolre\/ioz070","article-title":"Functions of cyclins and CDKs in mammalian gametogenesisdagger","volume":"101","author":"Chotiner","year":"2019","journal-title":"Biol Reprod"},{"key":"2026021004381033000_B77","doi-asserted-by":"publisher","first-page":"679","DOI":"10.3390\/cells10030679","article-title":"SRSF9 regulates cassette exon splicing of caspase-2 by interacting with its downstream exon","volume":"10","author":"Ha","year":"2021","journal-title":"Cells"},{"key":"2026021004381033000_B78","doi-asserted-by":"publisher","first-page":"12074","DOI":"10.3390\/ijms252212074","article-title":"The intricate functional networks of pre-mRNA alternative splicing in mammalian spermatogenesis","volume":"25","author":"Jiang","year":"2024","journal-title":"Int J Mol Sci"},{"key":"2026021004381033000_B79","doi-asserted-by":"publisher","first-page":"e78211","DOI":"10.7554\/eLife.78211","article-title":"SRSF10 is essential for progenitor spermatogonia expansion by regulating alternative splicing","volume":"11","author":"Liu","year":"2022","journal-title":"eLife"},{"key":"2026021004381033000_B80","doi-asserted-by":"publisher","first-page":"601","DOI":"10.1016\/j.molcel.2013.04.026","article-title":"Multiple epigenetic mechanisms and the piRNA pathway enforce LINE1 silencing during adult spermatogenesis","volume":"50","author":"Di\u00a0Giacomo","year":"2013","journal-title":"Mol Cell"}],"container-title":["Nucleic Acids Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/nar\/article-pdf\/54\/4\/gkag108\/66838561\/gkag108.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/nar\/article-pdf\/54\/4\/gkag108\/66838561\/gkag108.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,10]],"date-time":"2026-02-10T09:38:20Z","timestamp":1770716300000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/nar\/article\/doi\/10.1093\/nar\/gkag108\/8471004"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,5]]},"references-count":80,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2026,2,5]]}},"URL":"https:\/\/doi.org\/10.1093\/nar\/gkag108","relation":{},"ISSN":["0305-1048","1362-4962"],"issn-type":[{"value":"0305-1048","type":"print"},{"value":"1362-4962","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,2,27]]},"published":{"date-parts":[[2026,2,5]]},"article-number":"gkag108"}}