{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,17]],"date-time":"2026-07-17T11:09:08Z","timestamp":1784286548997,"version":"3.55.0"},"reference-count":107,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,5,9]],"date-time":"2026-05-09T00:00:00Z","timestamp":1778284800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,7,17]],"date-time":"2026-07-17T00:00:00Z","timestamp":1784246400000},"content-version":"vor","delay-in-days":69,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Commun Biol"],"DOI":"10.1038\/s42003-026-10235-7","type":"journal-article","created":{"date-parts":[[2026,5,9]],"date-time":"2026-05-09T16:53:07Z","timestamp":1778345587000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Gene regulatory networks orchestrating oocyte fate bifurcation in primordial follicles revealed by single-cell transcriptomics"],"prefix":"10.1038","volume":"9","author":[{"given":"Hanwen","family":"Zhang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xingsi","family":"He","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qiuzhen","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Min","family":"Su","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huanyu","family":"Yan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiao","family":"Zeng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2548-3689","authenticated-orcid":false,"given":"Shicai","family":"Fan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7572-6354","authenticated-orcid":false,"given":"Xi","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,9]]},"reference":[{"key":"10235_CR1","doi-asserted-by":"publisher","first-page":"R229","DOI":"10.1530\/REP-13-0202","volume":"146","author":"H Zhang","year":"2013","unstructured":"Zhang, H., Adhikari, D., Zheng, W. & Liu, K. Combating ovarian aging depends on the use of existing ovarian follicles, not on putative oogonial stem cells. Reproduction 146, R229\u2013R233 (2013).","journal-title":"Reproduction"},{"key":"10235_CR2","doi-asserted-by":"publisher","first-page":"e14338","DOI":"10.1111\/acel.14338","volume":"24","author":"S Qin","year":"2025","unstructured":"Qin, S. et al. Oocytes maintain low ROS levels to support the dormancy of primordial follicles. Aging Cell 24, e14338 (2025).","journal-title":"Aging Cell"},{"key":"10235_CR3","doi-asserted-by":"publisher","first-page":"e13424","DOI":"10.1111\/acel.13424","volume":"20","author":"Y He","year":"2021","unstructured":"He, Y. et al. Single-cell RNA-Seq reveals a highly coordinated transcriptional program in mouse germ cells during primordial follicle formation. Aging Cell 20, e13424 (2021).","journal-title":"Aging Cell"},{"key":"10235_CR4","doi-asserted-by":"publisher","first-page":"920","DOI":"10.1093\/hmg\/ddt486","volume":"23","author":"W Zheng","year":"2014","unstructured":"Zheng, W. et al. Two classes of ovarian primordial follicles exhibit distinct developmental dynamics and physiological functions. Hum. Mol. Genet. 23, 920\u2013928 (2014).","journal-title":"Hum. Mol. Genet."},{"key":"10235_CR5","doi-asserted-by":"publisher","first-page":"20015","DOI":"10.1073\/pnas.2005570117","volume":"117","author":"W Niu","year":"2020","unstructured":"Niu, W. & Spradling, A. C. Two distinct pathways of pregranulosa cell differentiation support follicle formation in the mouse ovary. Proc. Natl. Acad. Sci. Usa. 117, 20015\u201320026 (2020).","journal-title":"Proc. Natl. Acad. Sci. Usa."},{"key":"10235_CR6","doi-asserted-by":"publisher","first-page":"286","DOI":"10.1093\/molehr\/gau007","volume":"20","author":"W Zheng","year":"2014","unstructured":"Zheng, W., Zhang, H. & Liu, K. The two classes of primordial follicles in the mouse ovary: their development, physiological functions and implications for future research. Mol. Hum. Reprod. 20, 286\u2013292 (2014).","journal-title":"Mol. Hum. Reprod."},{"key":"10235_CR7","doi-asserted-by":"publisher","first-page":"672890","DOI":"10.3389\/fcell.2021.672890","volume":"9","author":"SJ Chon","year":"2021","unstructured":"Chon, S. J., Umair, Z. & Yoon, M.-S. Premature ovarian insufficiency: past, present, and future. Front. Cell Develop. Biol. 9, 672890 (2021).","journal-title":"Front. Cell Develop. Biol."},{"key":"10235_CR8","doi-asserted-by":"publisher","first-page":"e3001025","DOI":"10.1371\/journal.pbio.3001025","volume":"18","author":"JJ Wang","year":"2020","unstructured":"Wang, J. J. et al. Single-cell transcriptome landscape of ovarian cells during primordial follicle assembly in mice. PLoS Biol. 18, e3001025 (2020).","journal-title":"PLoS Biol."},{"key":"10235_CR9","volume":"9","author":"M Su","year":"2022","unstructured":"Su, M. et al. Data analysis guidelines for single-cell RNA-seq in biomedical studies and clinical applications. Mil. Med. Res. 9, 68 (2022).","journal-title":"Mil. Med. Res."},{"key":"10235_CR10","doi-asserted-by":"publisher","first-page":"qzad001","DOI":"10.1093\/gpbjnl\/qzad001","volume":"22","author":"W Liu","year":"2024","unstructured":"Liu, W. et al. Transcriptome dynamics and cell dialogs between oocytes and granulosa cells in mouse follicle development. Genomics, Proteom. Bioinforma. 22, qzad001 (2024).","journal-title":"Genomics, Proteom. Bioinforma."},{"key":"10235_CR11","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1093\/biolre\/ioz093","volume":"101","author":"Y Wang","year":"2019","unstructured":"Wang, Y. et al. Influence of mouse defective zona pellucida in folliculogenesis on apoptosis of granulosa cells and developmental competence of oocytes. Biol. Reprod. 101, 457\u2013465 (2019).","journal-title":"Biol. Reprod."},{"key":"10235_CR12","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms10324","volume":"7","author":"E Abby","year":"2016","unstructured":"Abby, E. et al. Implementation of meiosis prophase I programme requires a conserved retinoid-independent stabilizer of meiotic transcripts. Nat. Commun. 7, 10324 (2016).","journal-title":"Nat. Commun."},{"key":"10235_CR13","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-42259-6","volume":"14","author":"R Shimada","year":"2023","unstructured":"Shimada, R. et al. STRA8\u2013RB interaction is required for timely entry of meiosis in mouse female germ cells. Nat. Commun. 14, 6443 (2023).","journal-title":"Nat. Commun."},{"key":"10235_CR14","doi-asserted-by":"publisher","first-page":"e1006128","DOI":"10.1371\/journal.pgen.1006128","volume":"12","author":"KJ Grive","year":"2016","unstructured":"Grive, K. J. et al. TAF4b regulates oocyte-specific genes essential for meiosis. PLoS Genet 12, e1006128 (2016).","journal-title":"PLoS Genet"},{"key":"10235_CR15","doi-asserted-by":"publisher","first-page":"2106","DOI":"10.1172\/JCI90281","volume":"127","author":"Y-H Shin","year":"2017","unstructured":"Shin, Y.-H. et al. Transcription factors SOHLH1 and SOHLH2 coordinate oocyte differentiation without affecting meiosis I. J. Clin. Investig. 127, 2106\u20132117 (2017).","journal-title":"J. Clin. Investig."},{"key":"10235_CR16","doi-asserted-by":"publisher","first-page":"3525","DOI":"10.1093\/nar\/gkaa101","volume":"48","author":"Z Wang","year":"2020","unstructured":"Wang, Z., Liu, C.-Y., Zhao, Y. & Dean, J. FIGLA, LHX8 and SOHLH1 transcription factor networks regulate mouse oocyte growth and differentiation. Nucleic Acids Res. 48, 3525\u20133541 (2020).","journal-title":"Nucleic Acids Res."},{"key":"10235_CR17","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1038\/383531a0","volume":"383","author":"J Dong","year":"1996","unstructured":"Dong, J. et al. Growth differentiation factor-9 is required during early ovarian folliculogenesis. Nature 383, 531\u2013535 (1996).","journal-title":"Nature"},{"key":"10235_CR18","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1016\/S0303-7207(02)00047-3","volume":"191","author":"SM Galloway","year":"2002","unstructured":"Galloway, S. M. et al. Bmp15 mutations and ovarian function. Mol. Cell. Endocrinol. 191, 15\u201318 (2002).","journal-title":"Mol. Cell. Endocrinol."},{"key":"10235_CR19","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1016\/j.bbrc.2023.05.058","volume":"667","author":"K Yamamoto","year":"2023","unstructured":"Yamamoto, K. et al. Stimulatory effects of vasopressin on progesterone production and BMP signaling by ovarian granulosa cells. Biochem. Biophys. Res. Commun. 667, 132\u2013137 (2023).","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"10235_CR20","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1038\/s41421-022-00492-1","volume":"8","author":"J Zhou","year":"2022","unstructured":"Zhou, J. et al. Dissecting the fate of Foxl2-expressing cells in fetal ovary using lineage tracing and single-cell transcriptomics. Cell Discov. 8, 139 (2022).","journal-title":"Cell Discov."},{"key":"10235_CR21","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1530\/REP-22-0053","volume":"164","author":"Y-Y Chen","year":"2022","unstructured":"Chen, Y.-Y. et al. Single-cell transcriptomics of staged oocytes and somatic cells reveal novel regulators of follicle activation. Reproduction 164, 55\u201370 (2022).","journal-title":"Reproduction"},{"key":"10235_CR22","doi-asserted-by":"publisher","first-page":"188","DOI":"10.1038\/s41594-022-00909-1","volume":"30","author":"JH Crichton","year":"2023","unstructured":"Crichton, J. H. et al. Structural maturation of SYCP1-mediated meiotic chromosome synapsis by SYCE3. Nat. Struct. Mol. Biol. 30, 188\u2013199 (2023).","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10235_CR23","doi-asserted-by":"publisher","first-page":"1107","DOI":"10.1016\/j.molcel.2022.02.025","volume":"82","author":"G Biancon","year":"2022","unstructured":"Biancon, G. et al. Precision analysis of mutant U2AF1 activity reveals deployment of stress granules in myeloid malignancies. Mol. Cell 82, 1107\u20131122 e1107 (2022).","journal-title":"Mol. Cell"},{"key":"10235_CR24","doi-asserted-by":"publisher","first-page":"4212","DOI":"10.1002\/alz.13847","volume":"20","author":"Y Hou","year":"2024","unstructured":"Hou, Y. et al. Urolithin A improves Alzheimer\u2019s disease cognition and restores mitophagy and lysosomal functions. Alzheimers Dement 20, 4212\u20134233 (2024).","journal-title":"Alzheimers Dement"},{"key":"10235_CR25","doi-asserted-by":"publisher","first-page":"42710","DOI":"10.31083\/FBL42710","volume":"30","author":"T Yamazaki","year":"2025","unstructured":"Yamazaki, T., Bari, M. W. & Kishigami, S. Requirement of lysosomal two-pore channels for normal fertilization and artificial oocyte activation in mice. Front Biosci. 30, 42710 (2025).","journal-title":"Front Biosci."},{"key":"10235_CR26","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1530\/REP-16-0268","volume":"152","author":"JT Busada","year":"2016","unstructured":"Busada, J. T. et al. Rhox13 is required for a quantitatively normal first wave of spermatogenesis in mice. Reproduction 152, 379\u2013388 (2016).","journal-title":"Reproduction"},{"key":"10235_CR27","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1038\/s42255-022-00576-y","volume":"4","author":"E Danileviciute","year":"2022","unstructured":"Danileviciute, E. et al. PARK7\/DJ-1 promotes pyruvate dehydrogenase activity and maintains T(reg) homeostasis during ageing. Nat. Metab. 4, 589\u2013607 (2022).","journal-title":"Nat. Metab."},{"key":"10235_CR28","doi-asserted-by":"publisher","unstructured":"Cunatova, K. et al. Loss of COX4I1 leads to combined respiratory chain deficiency and impaired mitochondrial protein synthesis. Cells 10, https:\/\/doi.org\/10.3390\/cells10020369 (2021).","DOI":"10.3390\/cells10020369"},{"key":"10235_CR29","doi-asserted-by":"publisher","first-page":"E384","DOI":"10.1152\/ajpendo.00226.2010","volume":"299","author":"A Falcon","year":"2010","unstructured":"Falcon, A. et al. FATP2 is a hepatic fatty acid transporter and peroxisomal very long-chain acyl-CoA synthetase. Am. J. Physiol. Endocrinol. Metab. 299, E384\u2013E393 (2010).","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"10235_CR30","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-47222-7","volume":"15","author":"D Muoio","year":"2024","unstructured":"Muoio, D. et al. PARP2 promotes break induced replication-mediated telomere fragility in response to replication stress. Nat. Commun. 15, 2857 (2024).","journal-title":"Nat. Commun."},{"key":"10235_CR31","doi-asserted-by":"publisher","first-page":"2113","DOI":"10.1016\/j.molcel.2022.04.027","volume":"82","author":"ME Pesenti","year":"2022","unstructured":"Pesenti, M. E. et al. Structure of the human inner kinetochore CCAN complex and its significance for human centromere organization. Mol. Cell 82, 2113\u20132131.e2118 (2022).","journal-title":"Mol. Cell"},{"key":"10235_CR32","doi-asserted-by":"publisher","first-page":"937","DOI":"10.1038\/ncb2802","volume":"15","author":"Z Holubcova","year":"2013","unstructured":"Holubcova, Z., Howard, G. & Schuh, M. Vesicles modulate an actin network for asymmetric spindle positioning. Nat. Cell Biol. 15, 937\u2013947 (2013).","journal-title":"Nat. Cell Biol."},{"key":"10235_CR33","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1016\/j.tcb.2014.02.004","volume":"24","author":"T Welz","year":"2014","unstructured":"Welz, T., Wellbourne-Wood, J. & Kerkhoff, E. Orchestration of cell surface proteins by Rab11. Trends Cell Biol. 24, 407\u2013415 (2014).","journal-title":"Trends Cell Biol."},{"key":"10235_CR34","doi-asserted-by":"publisher","first-page":"3487","DOI":"10.4161\/cc.10.20.17742","volume":"10","author":"BB Chen","year":"2011","unstructured":"Chen, B. B., Glasser, J. R., Coon, T. A. & Mallampalli, R. K. FBXL2 is a ubiquitin E3 ligase subunit that triggers mitotic arrest. Cell Cycle 10, 3487\u20133494 (2011).","journal-title":"Cell Cycle"},{"key":"10235_CR35","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-024-55376-z","volume":"14","author":"C Simerly","year":"2024","unstructured":"Simerly, C. et al. Male meiotic spindle poles are stabilized by TACC3 and cKAP5\/chTOG differently from female meiotic or somatic mitotic spindles in mice. Sci. Rep. 14, 4808 (2024).","journal-title":"Sci. Rep."},{"key":"10235_CR36","doi-asserted-by":"publisher","first-page":"8111","DOI":"10.1073\/pnas.1222166110","volume":"110","author":"Y Fujihara","year":"2013","unstructured":"Fujihara, Y. et al. Expression of TEX101, regulated by ACE, is essential for the production of fertile mouse spermatozoa. Proc. Natl. Acad. Sci. USA 110, 8111\u20138116 (2013).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10235_CR37","doi-asserted-by":"publisher","first-page":"eabj6647","DOI":"10.1126\/science.abj6647","volume":"377","author":"JY Kang","year":"2022","unstructured":"Kang, J. Y. et al. LLPS of FXR1 drives spermiogenesis by activating translation of stored mRNAs. Science 377, eabj6647 (2022).","journal-title":"Science"},{"key":"10235_CR38","doi-asserted-by":"publisher","unstructured":"Han, S. et al. Myo-Inositol Oxygenase (MIOX): a pivotal regulator and therapeutic target in multiple diseases. Curr. Issues Mol. Biol. 47 https:\/\/doi.org\/10.3390\/cimb47090745 (2025).","DOI":"10.3390\/cimb47090745"},{"key":"10235_CR39","doi-asserted-by":"publisher","unstructured":"Zhao, Y. et al. Current understandings of core pathways for the activation of mammalian primordial follicles. Cells 10 https:\/\/doi.org\/10.3390\/cells10061491 (2021).","DOI":"10.3390\/cells10061491"},{"key":"10235_CR40","doi-asserted-by":"publisher","first-page":"695","DOI":"10.1146\/annurev-biochem-011520-105310","volume":"89","author":"ES Litscher","year":"2020","unstructured":"Litscher, E. S. & Wassarman, P. M. Zona pellucida proteins, fibrils, and matrix. Annu. Rev. Biochem 89, 695\u2013715 (2020).","journal-title":"Annu. Rev. Biochem"},{"key":"10235_CR41","doi-asserted-by":"publisher","first-page":"R161","DOI":"10.1530\/REP-17-0465","volume":"154","author":"D Monk","year":"2017","unstructured":"Monk, D., Sanchez-Delgado, M. & Fisher, R. NLRPs, the subcortical maternal complex and genomic imprinting. Reproduction 154, R161\u2013R170 (2017).","journal-title":"Reproduction"},{"key":"10235_CR42","doi-asserted-by":"publisher","first-page":"1014","DOI":"10.1016\/j.molmed.2017.09.004","volume":"23","author":"X Lu","year":"2017","unstructured":"Lu, X., Gao, Z., Qin, D. & Li, L. A maternal functional module in the mammalian oocyte-to-embryo transition. Trends Mol. Med. 23, 1014\u20131023 (2017).","journal-title":"Trends Mol. Med."},{"key":"10235_CR43","doi-asserted-by":"publisher","first-page":"1707","DOI":"10.1007\/s00018-018-2750-y","volume":"75","author":"C Liu","year":"2018","unstructured":"Liu, C., Ma, Y., Shang, Y., Huo, R. & Li, W. Post-translational regulation of the maternal-to-zygotic transition. Cell Mol. Life Sci. 75, 1707\u20131722 (2018).","journal-title":"Cell Mol. Life Sci."},{"key":"10235_CR44","doi-asserted-by":"publisher","first-page":"2300043","DOI":"10.1002\/advs.202300043","volume":"10","author":"H Li","year":"2023","unstructured":"Li, H. et al. LSM14B is an oocyte-specific RNA-binding protein indispensable for maternal mRNA metabolism and oocyte development in mice. Adv. Sci. 10, 2300043 (2023).","journal-title":"Adv. Sci."},{"key":"10235_CR45","doi-asserted-by":"publisher","first-page":"578","DOI":"10.1017\/S0967199415000556","volume":"24","author":"AY Cavalcante","year":"2016","unstructured":"Cavalcante, A. Y. et al. Kit ligand promotes the transition from primordial to primary follicles after in vitro culture of ovine ovarian tissue. Zygote 24, 578\u2013582 (2016).","journal-title":"Zygote"},{"key":"10235_CR46","doi-asserted-by":"publisher","first-page":"111460","DOI":"10.1016\/j.mce.2021.111460","volume":"539","author":"J Chen","year":"2022","unstructured":"Chen, J. et al. Overexpression of Lin28a induces a primary ovarian insufficiency phenotype via facilitation of primordial follicle activation in mice. Mol. Cell. Endocrinol. 539, 111460 (2022).","journal-title":"Mol. Cell. Endocrinol."},{"key":"10235_CR47","doi-asserted-by":"publisher","unstructured":"Sutherland, J. M. et al. Janus kinase JAK1 maintains the ovarian reserve of primordial follicles in the mouse ovary. MHR: Basic science of reproductive medicine https:\/\/doi.org\/10.1093\/molehr\/gay041 (2018).","DOI":"10.1093\/molehr\/gay041"},{"key":"10235_CR48","doi-asserted-by":"publisher","first-page":"576","DOI":"10.1086\/519496","volume":"81","author":"Y Qin","year":"2007","unstructured":"Qin, Y. et al. NOBOX Homeobox Mutation Causes Premature Ovarian Failure.  Am. J. Hum. Genet. 81, 576\u2013581 (2007).","journal-title":"Am. J. Hum. Genet."},{"key":"10235_CR49","doi-asserted-by":"publisher","first-page":"1157","DOI":"10.1126\/science.1099755","volume":"305","author":"A Rajkovic","year":"2004","unstructured":"Rajkovic, A., Pangas, S. A., Ballow, D., Suzumori, N. & Matzuk, M. M. NOBOX Deficiency Disrupts Early Folliculogenesis and Oocyte-Specific Gene Expression. Science 305, 1157\u20131159 (2004).","journal-title":"Science"},{"key":"10235_CR50","doi-asserted-by":"publisher","first-page":"e53810","DOI":"10.1371\/journal.pone.0053810","volume":"8","author":"D Adhikari","year":"2013","unstructured":"Adhikari, D., Risal, S., Liu, K. & Shen, Y. Pharmacological inhibition of mTORC1 prevents over-activation of the primordial follicle pool in response to elevated PI3K signaling. PLoS ONE 8, e53810 (2013).","journal-title":"PLoS ONE"},{"key":"10235_CR51","doi-asserted-by":"publisher","first-page":"397","DOI":"10.1093\/hmg\/ddp483","volume":"19","author":"D Adhikari","year":"2010","unstructured":"Adhikari, D. et al. Tsc\/mTORC1 signaling in oocytes governs the quiescence and activation of primordial follicles. Hum. Mol. Genet. 19, 397\u2013410 (2010).","journal-title":"Hum. Mol. Genet."},{"key":"10235_CR52","doi-asserted-by":"publisher","first-page":"e110491","DOI":"10.1371\/journal.pone.0110491","volume":"9","author":"N Gorre","year":"2014","unstructured":"Gorre, N. et al. mTORC1 signaling in oocytes is dispensable for the survival of primordial follicles and for female fertility. PLoS ONE 9, e110491 (2014).","journal-title":"PLoS ONE"},{"key":"10235_CR53","doi-asserted-by":"publisher","first-page":"8299","DOI":"10.1074\/jbc.M113.532952","volume":"289","author":"Z-P Wang","year":"2014","unstructured":"Wang, Z.-P. et al. Transforming growth factor-\u03b2 signaling participates in the maintenance of the primordial follicle pool in the mouse ovary. J. Biol. Chem. 289, 8299\u20138311 (2014).","journal-title":"J. Biol. Chem."},{"key":"10235_CR54","doi-asserted-by":"publisher","first-page":"3863","DOI":"10.7150\/ijbs.95020","volume":"20","author":"J Zhang","year":"2024","unstructured":"Zhang, J. et al. Participation of preovulatory follicles in the activation of primordial follicles in mouse ovaries. Int. J. Biol. Sci. 20, 3863\u20133880 (2024).","journal-title":"Int. J. Biol. Sci."},{"key":"10235_CR55","doi-asserted-by":"publisher","first-page":"eaav9960","DOI":"10.1126\/sciadv.aav9960","volume":"5","author":"G Nagamatsu","year":"2019","unstructured":"Nagamatsu, G., Shimamoto, S., Hamazaki, N., Nishimura, Y. & Hayashi, K. Mechanical stress accompanied with nuclear rotation is involved in the dormant state of mouse oocytes. Sci. Adv. 5, eaav9960 (2019).","journal-title":"Sci. Adv."},{"key":"10235_CR56","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1038\/nbt.2859","volume":"32","author":"C Trapnell","year":"2014","unstructured":"Trapnell, C. et al. The dynamics and regulators of cell fate decisions are revealed by pseudotemporal ordering of single cells. Nat. Biotechnol. 32, 381\u2013386 (2014).","journal-title":"Nat. Biotechnol."},{"key":"10235_CR57","doi-asserted-by":"publisher","first-page":"12321","DOI":"10.1073\/pnas.1817223116","volume":"116","author":"S Shimamoto","year":"2019","unstructured":"Shimamoto, S. et al. Hypoxia induces the dormant state in oocytes through expression of Foxo3. Proc. Natl. Acad. Sci. USA 116, 12321\u201312326 (2019).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10235_CR58","doi-asserted-by":"publisher","first-page":"1247","DOI":"10.1038\/s42003-024-06900-4","volume":"7","author":"L Lei","year":"2024","unstructured":"Lei, L. et al. The mouse Balbiani body regulates primary oocyte quiescence via RNA storage. Commun. Biol. 7, 1247 (2024).","journal-title":"Commun. Biol."},{"key":"10235_CR59","doi-asserted-by":"publisher","first-page":"1158510","DOI":"10.3389\/fragi.2023.1158510","volume":"4","author":"G Nagamatsu","year":"2023","unstructured":"Nagamatsu, G. Oocyte aging in comparison to stem cells in mice. Front. Aging 4, 1158510 (2023).","journal-title":"Front. Aging"},{"key":"10235_CR60","doi-asserted-by":"publisher","first-page":"264","DOI":"10.1038\/s41586-020-3027-9","volume":"589","author":"N Hamazaki","year":"2021","unstructured":"Hamazaki, N. et al. Reconstitution of the oocyte transcriptional network with transcription factors. Nature 589, 264\u2013269 (2021).","journal-title":"Nature"},{"key":"10235_CR61","doi-asserted-by":"publisher","unstructured":"Abbasi, F. et al. CRISPR\/Cas9-mediated genome editing reveals oosp family genes are dispensable for female fertility in mice. Cells 9 https:\/\/doi.org\/10.3390\/cells9040821 (2020).","DOI":"10.3390\/cells9040821"},{"key":"10235_CR62","doi-asserted-by":"publisher","first-page":"962","DOI":"10.1038\/s41556-024-01422-x","volume":"26","author":"B Liu","year":"2024","unstructured":"Liu, B. et al. Mapping putative enhancers in mouse oocytes and early embryos reveals TCF3\/12 as key folliculogenesis regulators. Nat. Cell Biol. 26, 962\u2013974 (2024).","journal-title":"Nat. Cell Biol."},{"key":"10235_CR63","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1210\/er.2008-0048","volume":"30","author":"D Adhikari","year":"2009","unstructured":"Adhikari, D. & Liu, K. Molecular mechanisms underlying the activation of mammalian primordial follicles. Endocr. Rev. 30, 438\u2013464 (2009).","journal-title":"Endocr. Rev."},{"key":"10235_CR64","doi-asserted-by":"publisher","first-page":"4939","DOI":"10.1242\/dev.124.24.4939","volume":"124","author":"L-f Liang","year":"1997","unstructured":"Liang, L. -f, Soyal, S. M. & Dean, J. FIG\u03b1, a germ cell specific transcription factor involved in the coordinate expression of the zona pellucida genes. Development 124, 4939\u20134947 (1997).","journal-title":"Development"},{"key":"10235_CR65","doi-asserted-by":"publisher","first-page":"8090","DOI":"10.1073\/pnas.0601083103","volume":"103","author":"SA Pangas","year":"2006","unstructured":"Pangas, S. A. et al. Oogenesis requires germ cell-specific transcriptional regulators Sohlh1 and Lhx8. Proc. Natl. Acad. Sci. USA 103, 8090\u20138095 (2006).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10235_CR66","doi-asserted-by":"publisher","first-page":"1021","DOI":"10.1016\/j.molcel.2018.10.029","volume":"72","author":"Y Zhang","year":"2018","unstructured":"Zhang, Y. et al. Transcriptome landscape of human folliculogenesis reveals oocyte and granulosa cell interactions. Mol. Cell 72, 1021\u20131034.e1024 (2018).","journal-title":"Mol. Cell"},{"key":"10235_CR67","doi-asserted-by":"publisher","first-page":"2051","DOI":"10.1042\/BST20201125","volume":"49","author":"L Tora","year":"2021","unstructured":"Tora, L. & Vincent, S. D. What defines the maternal transcriptome. Biochem Soc. Trans. 49, 2051\u20132062 (2021).","journal-title":"Biochem Soc. Trans."},{"key":"10235_CR68","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1095\/biolreprod.110.087122","volume":"84","author":"S Matoba","year":"2011","unstructured":"Matoba, S. & Ogura, A. Generation of functional oocytes and spermatids from fetal primordial germ cells after ectopic transplantation in adult mice. Biol. Reprod. 84, 631\u2013638 (2011).","journal-title":"Biol. Reprod."},{"key":"10235_CR69","doi-asserted-by":"publisher","first-page":"971","DOI":"10.1126\/science.1226889","volume":"338","author":"K Hayashi","year":"2012","unstructured":"Hayashi, K. et al. Offspring from oocytes derived from in vitro primordial germ cell-like cells in mice. Science 338, 971\u2013975 (2012).","journal-title":"Science"},{"key":"10235_CR70","doi-asserted-by":"publisher","DOI":"10.1038\/s41419-018-1208-3","volume":"10","author":"H Yan","year":"2019","unstructured":"Yan, H. et al. Oocyte-derived E-cadherin acts as a multiple functional factor maintaining the primordial follicle pool in mice. Cell Death Dis. 10, 160 (2019).","journal-title":"Cell Death Dis."},{"key":"10235_CR71","doi-asserted-by":"publisher","first-page":"946","DOI":"10.1016\/j.ccell.2024.04.009","volume":"42","author":"Y Liu","year":"2024","unstructured":"Liu, Y., Su, Z., Tavana, O. & Gu, W. Understanding the complexity of p53 in a new era of tumor suppression. Cancer Cell 42, 946\u2013967 (2024).","journal-title":"Cancer Cell"},{"key":"10235_CR72","doi-asserted-by":"publisher","first-page":"150","DOI":"10.1111\/dgd.12653","volume":"62","author":"K Hayashi","year":"2020","unstructured":"Hayashi, K., Shimamoto, S. & Nagamatsu, G. Environmental factors for establishment of the dormant state in oocytes. Dev. Growth Differ. 62, 150\u2013157 (2020).","journal-title":"Dev. Growth Differ."},{"key":"10235_CR73","doi-asserted-by":"publisher","first-page":"2189","DOI":"10.1210\/me.2007-0172","volume":"21","author":"S Rajareddy","year":"2007","unstructured":"Rajareddy, S. et al. p27kip1 (Cyclin-Dependent Kinase Inhibitor 1B) controls ovarian development by suppressing follicle endowment and activation and promoting follicle atresia in mice. Mol. Endocrinol. 21, 2189\u20132202 (2007).","journal-title":"Mol. Endocrinol."},{"key":"10235_CR74","doi-asserted-by":"publisher","DOI":"10.1186\/s12915-018-0541-4","volume":"16","author":"H Yan","year":"2018","unstructured":"Yan, H. et al. CDC42 controls the activation of primordial follicles by regulating PI3K signaling in mouse oocytes. BMC Biol. 16, 73 (2018).","journal-title":"BMC Biol."},{"key":"10235_CR75","doi-asserted-by":"publisher","first-page":"727614","DOI":"10.3389\/fmolb.2021.727614","volume":"8","author":"Y He","year":"2021","unstructured":"He, Y. et al. Pervasive 3\u2019-UTR isoform switches during mouse oocyte maturation. Front Mol. Biosci. 8, 727614 (2021).","journal-title":"Front Mol. Biosci."},{"key":"10235_CR76","doi-asserted-by":"publisher","DOI":"10.1038\/s41419-021-03842-1","volume":"12","author":"T Zhang","year":"2021","unstructured":"Zhang, T. et al. HDAC6 regulates primordial follicle activation through mTOR signaling pathway. Cell Death Dis. 12, 559 (2021).","journal-title":"Cell Death Dis."},{"key":"10235_CR77","doi-asserted-by":"publisher","DOI":"10.1038\/s41419-023-05984-w","volume":"14","author":"W Chen","year":"2023","unstructured":"Chen, W. et al. PARP1-catalyzed PARylation of YY1 mediates endoplasmic reticulum stress in granulosa cells to determine primordial follicle activation. Cell Death Dis. 14, 524 (2023).","journal-title":"Cell Death Dis."},{"key":"10235_CR78","doi-asserted-by":"publisher","first-page":"747","DOI":"10.1038\/s42003-024-06418-9","volume":"7","author":"JF Madsen","year":"2024","unstructured":"Madsen, J. F. et al. Dorsomorphin inhibits AMPK, upregulates Wnt and Foxo genes and promotes the activation of dormant follicles. Commun. Biol. 7, 747 (2024).","journal-title":"Commun. Biol."},{"key":"10235_CR79","doi-asserted-by":"publisher","first-page":"756","DOI":"10.1038\/s41586-022-04979-5","volume":"607","author":"A Rodriguez-Nuevo","year":"2022","unstructured":"Rodriguez-Nuevo, A. et al. Oocytes maintain ROS-free mitochondrial metabolism by suppressing complex I. Nature 607, 756\u2013761 (2022).","journal-title":"Nature"},{"key":"10235_CR80","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1016\/j.ydbio.2005.08.023","volume":"286","author":"H Pan","year":"2005","unstructured":"Pan, H., O\u2019Brien, M. J., Wigglesworth, K., Eppig, J. J. & Schultz, R. M. Transcript profiling during mouse oocyte development and the effect of gonadotropin priming and development in vitro. Develop. Biol. 286, 493\u2013506 (2005).","journal-title":"Develop. Biol."},{"key":"10235_CR81","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1071\/RD09216","volume":"22","author":"T Fair","year":"2010","unstructured":"Fair, T. Mammalian oocyte development: checkpoints for competence. Reprod. Fertil. Dev. 22, 13\u201320 (2010).","journal-title":"Reprod. Fertil. Dev."},{"key":"10235_CR82","doi-asserted-by":"publisher","first-page":"eabq4835","DOI":"10.1126\/science.abq4835","volume":"378","author":"S Cheng","year":"2022","unstructured":"Cheng, S. et al. Mammalian oocytes store mRNAs in a mitochondria-associated membraneless compartment. Science 378, eabq4835 (2022).","journal-title":"Science"},{"key":"10235_CR83","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1016\/j.tcb.2024.07.006","volume":"35","author":"JW Pek","year":"2025","unstructured":"Pek, J. W. The idiosyncrasies of oocytes. Trends Cell Biol. 35, 305\u2013315 (2025).","journal-title":"Trends Cell Biol."},{"key":"10235_CR84","doi-asserted-by":"publisher","first-page":"6053","DOI":"10.1093\/nar\/gkab293","volume":"49","author":"M Mancini","year":"2021","unstructured":"Mancini, M., Magnani, E., Macchi, F. & Bonapace, I. M. The multi-functionality of UHRF1: epigenome maintenance and preservation of genome integrity. Nucleic Acids Res. 49, 6053\u20136068 (2021).","journal-title":"Nucleic Acids Res."},{"key":"10235_CR85","doi-asserted-by":"publisher","first-page":"4645","DOI":"10.1242\/dev.127.21.4645","volume":"127","author":"SM Soyal","year":"2000","unstructured":"Soyal, S. M., Amleh, A. & Dean, J. FIGalpha, a germ cell-specific transcription factor required for ovarian follicle formation. Development 127, 4645\u20134654 (2000).","journal-title":"Development"},{"key":"10235_CR86","doi-asserted-by":"publisher","first-page":"1176","DOI":"10.1095\/biolreprod.108.071217","volume":"79","author":"Y Choi","year":"2008","unstructured":"Choi, Y., Yuan, D. & Rajkovic, A. Germ cell-specific transcriptional regulator Sohlh2 is essential for early mouse folliculogenesis and oocyte-specific gene expression1. Biol. Reprod. 79, 1176\u20131182 (2008).","journal-title":"Biol. Reprod."},{"key":"10235_CR87","doi-asserted-by":"publisher","first-page":"6145","DOI":"10.7150\/ijbs.75063","volume":"18","author":"P Li","year":"2022","unstructured":"Li, P. et al. ELK1-mediated YTHDF1 drives prostate cancer progression by facilitating the translation of Polo-like kinase 1 in an m6A dependent manner. Int. J. Biol. Sci. 18, 6145\u20136162 (2022).","journal-title":"Int. J. Biol. Sci."},{"key":"10235_CR88","doi-asserted-by":"publisher","first-page":"1623135","DOI":"10.3389\/fcell.2025.1623135","volume":"13","author":"A Abdurahman","year":"2025","unstructured":"Abdurahman, A., Ga, Y. & Ma, X. ELK1 regulates BMPR1B transcriptional activity in ovine granulosa cells. Front. Cell Develop. Biol. 13, 1623135 (2025).","journal-title":"Front. Cell Develop. Biol."},{"key":"10235_CR89","doi-asserted-by":"publisher","first-page":"7994","DOI":"10.1073\/pnas.141002798","volume":"98","author":"SE Yi","year":"2001","unstructured":"Yi, S. E. et al. The type I BMP receptor BmprIB is essential for female reproductive function. Proc. Natl. Acad. Sci. USA 98, 7994\u20137999 (2001).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10235_CR90","doi-asserted-by":"publisher","first-page":"6998","DOI":"10.1074\/jbc.M807670200","volume":"284","author":"GA Baltus","year":"2009","unstructured":"Baltus, G. A., Kowalski, M. P., Tutter, A. V. & Kadam, S. A positive regulatory role for the mSin3A-HDAC complex in pluripotency through Nanog and Sox2. J. Biol. Chem. 284, 6998\u20137006 (2009).","journal-title":"J. Biol. Chem."},{"key":"10235_CR91","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1007\/s13402-024-00973-3","volume":"48","author":"C Bret","year":"2025","unstructured":"Bret, C., Desmots-Loyer, F., Moreaux, J. & Fest, T. BHLHE41, a transcriptional repressor involved in physiological processes and tumor development. Cell Oncol. 48, 43\u201366 (2025).","journal-title":"Cell Oncol."},{"key":"10235_CR92","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-42578-z","volume":"9","author":"Y Makino","year":"2019","unstructured":"Makino, Y. et al. Single cell RNA-sequencing identified Dec2 as a suppressive factor for spermatogonial differentiation by inhibiting Sohlh1 expression. Sci. Rep. 9, 6063 (2019).","journal-title":"Sci. Rep."},{"key":"10235_CR93","doi-asserted-by":"publisher","first-page":"3661","DOI":"10.1128\/MCB.00201-10","volume":"30","author":"W Hu","year":"2010","unstructured":"Hu, W., Gauthier, L., Baibakov, B., Jimenez-Movilla, M. & Dean, J. FIGLA, a basic helix-loop-helix transcription factor, balances sexually dimorphic gene expression in postnatal oocytes. Mol. Cell Biol. 30, 3661\u20133671 (2010).","journal-title":"Mol. Cell Biol."},{"key":"10235_CR94","doi-asserted-by":"publisher","first-page":"eadi5516","DOI":"10.1126\/science.adi5516","volume":"382","author":"L Li","year":"2023","unstructured":"Li, L. et al. Multifaceted SOX2-chromatin interaction underpins pluripotency progression in early embryos. Science 382, eadi5516 (2023).","journal-title":"Science"},{"key":"10235_CR95","doi-asserted-by":"publisher","first-page":"506","DOI":"10.1007\/s11427-025-3041-4","volume":"69","author":"X He","year":"2026","unstructured":"He, X. et al. Plat safeguards maternally aged oocytes against programmed cell death through activating the Erk1\/2 pathway. Sci. China Life Sci. 69, 506\u2013522 (2026).","journal-title":"Sci. China Life Sci."},{"key":"10235_CR96","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","volume":"29","author":"A Dobin","year":"2013","unstructured":"Dobin, A. et al. STAR: ultrafast universal RNA-seq aligner. Bioinformatics 29, 15\u201321 (2013).","journal-title":"Bioinformatics"},{"key":"10235_CR97","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1093\/bioinformatics\/btu638","volume":"31","author":"S Anders","year":"2015","unstructured":"Anders, S., Pyl, P. T. & Huber, W. HTSeq-a Python framework to work with high-throughput sequencing data. Bioinformatics 31, 166\u2013169 (2015).","journal-title":"Bioinformatics"},{"key":"10235_CR98","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2105-12-323","volume":"12","author":"B Li","year":"2011","unstructured":"Li, B. & Dewey, C. N. RSEM: accurate transcript quantification from RNA-Seq data with or without a reference genome. BMC Bioinforma. 12, 323 (2011).","journal-title":"BMC Bioinforma."},{"key":"10235_CR99","doi-asserted-by":"publisher","first-page":"495","DOI":"10.1038\/nbt.3192","volume":"33","author":"R Satija","year":"2015","unstructured":"Satija, R., Farrell, J. A., Gennert, D., Schier, A. F. & Regev, A. Spatial reconstruction of single-cell gene expression data. Nat. Biotechnol. 33, 495\u2013502 (2015).","journal-title":"Nat. Biotechnol."},{"key":"10235_CR100","doi-asserted-by":"publisher","first-page":"778","DOI":"10.1038\/s41587-023-01887-5","volume":"42","author":"K Wang","year":"2024","unstructured":"Wang, K. et al. PhyloVelo enhances transcriptomic velocity field mapping using monotonically expressed genes. Nat. Biotechnol. 42, 778\u2013789 (2024).","journal-title":"Nat. Biotechnol."},{"key":"10235_CR101","doi-asserted-by":"publisher","DOI":"10.1186\/s13059-014-0550-8","volume":"15","author":"MI Love","year":"2014","unstructured":"Love, M. I., Huber, W. & Anders, S. Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2. Genome Biol. 15, 550 (2014).","journal-title":"Genome Biol."},{"key":"10235_CR102","doi-asserted-by":"publisher","first-page":"W216","DOI":"10.1093\/nar\/gkac194","volume":"50","author":"BT Sherman","year":"2022","unstructured":"Sherman, B. T. et al. DAVID: a web server for functional enrichment analysis and functional annotation of gene lists (2021 update). Nucleic Acids Res 50, W216\u2013W221 (2022).","journal-title":"Nucleic Acids Res"},{"key":"10235_CR103","doi-asserted-by":"publisher","first-page":"2247","DOI":"10.1038\/s41596-020-0336-2","volume":"15","author":"B Van de Sande","year":"2020","unstructured":"Van de Sande, B. et al. A scalable SCENIC workflow for single-cell gene regulatory network analysis. Nat. Protoc. 15, 2247\u20132276 (2020).","journal-title":"Nat. Protoc."},{"key":"10235_CR104","doi-asserted-by":"publisher","first-page":"2498","DOI":"10.1101\/gr.1239303","volume":"13","author":"P Shannon","year":"2003","unstructured":"Shannon, P. et al. Cytoscape: a software environment for integrated models of biomolecular interaction networks. Genome Res. 13, 2498\u20132504 (2003).","journal-title":"Genome Res."},{"key":"10235_CR105","doi-asserted-by":"publisher","first-page":"1754","DOI":"10.1093\/bioinformatics\/btp324","volume":"25","author":"H Li","year":"2009","unstructured":"Li, H. & Durbin, R. Fast and accurate short read alignment with Burrows-Wheeler transform. Bioinformatics 25, 1754\u20131760 (2009).","journal-title":"Bioinformatics"},{"key":"10235_CR106","doi-asserted-by":"publisher","DOI":"10.1186\/gb-2008-9-9-r137","volume":"9","author":"Y Zhang","year":"2008","unstructured":"Zhang, Y. et al. Model-based analysis of ChIP-Seq MACS. Genome Biol. 9, R137 (2008).","journal-title":"Genome Biol."},{"key":"10235_CR107","doi-asserted-by":"publisher","first-page":"576","DOI":"10.1016\/j.molcel.2010.05.004","volume":"38","author":"S Heinz","year":"2010","unstructured":"Heinz, S. et al. Simple combinations of lineage-determining transcription factors prime cis-regulatory elements required for macrophage and B cell identities. Mol. Cell 38, 576\u2013589 (2010).","journal-title":"Mol. Cell"}],"container-title":["Communications Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s42003-026-10235-7","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-026-10235-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-026-10235-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,17]],"date-time":"2026-07-17T11:02:41Z","timestamp":1784286161000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s42003-026-10235-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,9]]},"references-count":107,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["10235"],"URL":"https:\/\/doi.org\/10.1038\/s42003-026-10235-7","relation":{},"ISSN":["2399-3642"],"issn-type":[{"value":"2399-3642","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,9]]},"assertion":[{"value":"29 March 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 April 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 May 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Ethics"}}],"article-number":"978"}}