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NANOG alone induces germ cells in primed epiblast in vitro by activation of enhancers
Open accessAbstract
Nanog, a core pluripotency factor in the inner cell mass of blastocysts, is also expressed in unipotent primordial germ cells (PGCs) in mice, where its precise role is yet unclear. We investigated this in an in vitro model, in which naive pluripotent embryonic stem (ES) cells cultured in basic fibroblast growth factor (bFGF) and activin A develop as epiblast-like cells (EpiLCs) and gain competence for a PGC-like fate. Consequently, bone morphogenetic protein 4 (BMP4), or ectopic expression of key germline transcription factors Prdm1, Prdm14 and Tfap2c, directly induce PGC-like cells (PGCLCs) in EpiLCs, but not in ES cells. Here we report an unexpected discovery that Nanog alone can induce PGCLCs in EpiLCs, independently of BMP4. We propose that after the dissolution of the naive ES-cell pluripotency network during establishment of EpiLCs, the epigenome is reset for cell fate determination. Indeed, we found genome-wide changes in NANOG-binding patterns between ES cells and EpiLCs, indicating epigenetic resetting of regulatory elements. Accordingly, we show that NANOG can bind and activate enhancers of Prdm1 and Prdm14 in EpiLCs in vitro; BLIMP1 (encoded by Prdm1) then directly induces Tfap2c. Furthermore, while SOX2 and NANOG promote the pluripotent state in ES cells, they show contrasting roles in EpiLCs, as Sox2 specifically represses PGCLC induction by Nanog. This study demonstrates a broadly applicable mechanistic principle for how cells acquire competence for cell fate determination, resulting in the context-dependent roles of key transcription factors during development.
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“For Dex inducible WT or APPA Mettl3 plasmid, GRBD insert from pPyCAG-cGR-IP plasmid (RIKEN, #RDB10440) was subcloned to constitutive PiggyBac plasmid to generate GRBD-FLAG-METTL3WT or GRBD-FLAG-METTL3APPA fusion protein (Murakami et al., 2016).”
From METTL3 uncouples chromatin accessibility from transcription during retinal development (2025) · cites this paper as Murakami et al., 2016
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“Recently, PGCs were defined as unipotent stem cells in Nature journal (Murakami et al., 2016) on the assumption that they only give rise to gametes in the body.”
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“Accordingly, the PGCLC induction system has been exploited for identification of transcription factors involved in PGC induction (Murakami et al., 2016; Nakaki et al., 2013), clarification of signaling mechanisms for PGC spec-ification (Aramaki et al., 2013), and, more recently, development of a PGCLC…”
Defined conditions for propagation and manipulation of mouse embryonic stem cells
Mulas, C., Kalkan, T., von Meyenn, F., Leitch, H. G., Nichols, J., & Smith, A. G. (2019) · Development · doi:10.1242/dev.173146 · Cited by 114
“The 2i platform has been widely exploited to study signalling, gene regulation and network control of naive pluripotency, and the transition path from pluripotency to lineage commitment (Blaschke et al., 2013; Carey et al., 2014; Dunn et al., 2014; Ficz et al., 2013; Habibi et al., 2013; Hackett et al., 2017; Hayashi…”
MethodDNA methylation and chromatin accessibility profiling of mouse and human fetal germ cells
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Metabolic regulation of pluripotency and germ cell fate through α‐ketoglutarate
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“…pluripotency, indicated by the sustained expression of Rex1 -GFPd2, in a dose-dependent manner ( Figs 2B While ESCs have the potential to self-renew and can generate colonies from single cells in naı¨ve pluripotency-promoting conditions, this ability is lost in 48 h EpiLCs (Murakami et al , 2016).”
Mammalian germ cells are determined after PGC colonization of the nascent gonad
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Niche stiffness sustains cancer stemness via TAZ and NANOG phase separation
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“Similar to previous reports (Murakami et al., 2016), cESCs fromEpiLC-48h ( Dox) cluster far from naive ESCs (ESCs in 2i/LIF).”
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Goszczynski, D., Cheng, H., Demyda-Peyrás, S., Medrano, J., Wu, J., & Ross, P. (2018) · Biology of Reproduction · doi:10.1093/biolre/ioy256 · Cited by 61
“It has also been observed that the induced expression of NANOG stimulates the induction of putative PGCLCs independently of BMP signaling in EpiLCs [89].”
BackgroundBasic and Clinical Approaches for Fertility Preservation and Restoration in Cancer Patients.
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MethodEsrrb Complementation Rescues Development of Nanog-Null Germ Cells
Zhang, M., Leitch, H. G., Tang, W. W. C., Festuccia, N., Hall-Ponselé, E., Nichols, J., Surani, M., Smith, A. G., & Chambers, I. (2018) · Cell Reports · doi:10.1016/j.celrep.2017.12.060 · Cited by 46
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“In the absence of Bmp4, robust PGCLC production can also be achieved by overexpression of three transcription factors (TFs)—namely Prdm1, Prdm14, and Tfap2c (7, 14)—or by a single TF Nanog (8) from exogenous vectors.”
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Modeling human infertility with pluripotent stem cells
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“The model proposed from this work follows that NANOG functions upstream of both PRDM14 and PRDM1 by binding to their enhancers in the germline-competent in vitro epiblast-like cells to promote PGC fate (Murakami et al., 2013).”
Induced formation of primordial germ cells from zebrafish blastomeres by germplasm factors
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Gastruloid Development Competence Discriminates Different States of Pluripotency
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Mammalian Germ Cell Development: From Mechanism to In Vitro Reconstitution
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Synthetic organizer cells guide development via spatial and biochemical instructions
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“This plasmid was digested with SpeI and XhoI, and the Tesra fragment was inserted into a pPBhCMV∗1-cHA-pA vector which was cut by SpeI and XhoI [51].”
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“Additionally, NANOG plays a crucial role in the specification of mammalian PGCs via their epigenetic modification [77,78].”
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DNA methylation and the core pluripotency network
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STRA8–RB interaction is required for timely entry of meiosis in mouse female germ cells
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“Our demonstration of a functional requirement of NANOG throughout the differentiation process altogether is consistent with a critical role for NANOG in mouse PGC speci fi cation in vivo and in vitro 53,54 .”
ResultEpigenome regulation during germ cell specification and development from pluripotent stem cells.
Kurimoto, K., & Saitou, M. (2018) · Current Opinion in Genetics and Development · doi:10.1016/j.gde.2018.06.004 · Cited by 23
What Can Stem Cell Models Tell Us About Human Germ Cell Biology?
Irie, N., Sybirna, A., & Surani, M. A. (2018) · Current Topics in Developmental Biology · doi:10.1016/bs.ctdb.2018.02.010 · Cited by 22
Protein phosphatase 1 acts as a RIF1 effector to suppress DSB resection prior to Shieldin action
Isobe, S.-Y., Hiraga, S., Nagao, K., Sasanuma, H., Donaldson, A. D., & Obuse, C. (2021) · Cell Reports · doi:10.1016/j.celrep.2021.109383 · Cited by 32
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PRDM14 and BLIMP1 control the development of chicken primordial germ cells.
Okuzaki, Y., Kaneoka, H., Suzuki, T., Hagihara, Y., Nakayama, Y., Murakami, S., Murase, Y., Kuroiwa, A., Iijima, S., & Nishijima, K. (2019) · Developmental Biology · doi:10.1016/j.ydbio.2019.06.018 · Cited by 21
Committing the primordial germ cell: An updated molecular perspective
Tan, H., & Tee, W.-W. (2018) · Wiley Interdisciplinary Reviews: Systems Biology and Medicine · doi:10.1002/wsbm.1436 · Cited by 21
“…the overexpression of Nanog in mEpiLCs also led to mPGCLC differentiation, and Nanog was found to bind and activate Prdm1 and Prdm14 expression (Murakami et al., 2016); conversely, differentiation of Nanog-null cells revealed a reduction in mPGCLC yield which could be rescued by ectopic Nanog…”
BackgroundInfluentialLincU Preserves Naïve Pluripotency by Restricting ERK Activity in Embryonic Stem Cells
Jiapaer, Z., Li, G.-P., Ye, D., Bai, M.-L., Li, J., Guo, X.-D., Du, Y.-H., Su, D., Jia, W.-W., Chen, W., et al. (2018) · Stem Cell Reports · doi:10.1016/j.stemcr.2018.06.010 · Cited by 21
“, 2003) we found that Nanog can bind the promoter region of LincU, which is located on chromosome 16 (9,133,405–9,138,147) (mm9) (Murakami et al., 2016).”
BackgroundBMP4 drives primed to naïve transition through PGC-like state
Yu, S., Zhou, C., He, J., Yao, Z., Huang, X., Rong, B., Zhu, H., Wang, S., Chen, S., Wang, X., et al. (2022) · Nature Communications · doi:10.1038/s41467-022-30325-4 · Cited by 20
Wang, X., Veerapandian, V., Yang, X., Song, K., Xu, X., Cui, M., Yuan, W.-Y., Huang, Y., Xia, X., Yao, Z., et al. (2021) · Stem Cell Reports · doi:10.1016/j.stemcr.2021.03.032 · Cited by 20
“Moreover, SOX15 peaks were detected at the proximal regulatory elements of several pluripotency-related genes, such as PRDM14, NANOG, ETV4, and ETV5 (Figures 6G and S7D) (Kalkan et al., 2019; Murakami et al., 2016; Sybirna et al., 2020).”
BackgroundFunctional characterization of NANOG in goat pre-implantation embryonic development.
Habibi, R., Hosseini, S., Zadegan, F. G., Hajian, M., Ostadhosseini, S., Vash, N. T., Naddafpour, A., & Esfahani, M. N. N. (2018) · Theriogenology · doi:10.1016/j.theriogenology.2018.07.023 · Cited by 20
Transcriptional control of human gametogenesis
Fang, F., Iaquinta, P. J., Xia, N., Liu, L., Diao, L., & Pera, R. A. R. (2022) · Human Reproduction Update · doi:10.1093/humupd/dmac002 · Cited by 23
“Pluripotency genes OCT4 and NANOG are re-expressed in both mPGCs (Murakami et al., 2016) and hPGCs (Guo et al., 2015; Tang et al., 2015), and SOX2 is absent in hPGCs (Lin et al., 2014) although .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..it is required for mouse PGC survival…”
Deb, S., Felix, D. A., Koch, P., Deb, M. K., Szafranski, K., Buder, K., Sannai, M., Groth, M., Kirkpatrick, J. M., Pietsch, S., et al. (2020) · EMBO Reports · doi:10.15252/embr.201949328 · Cited by 29
“Immunofluorescence staining was performed as previously described (Murakami et al , 2016).”
Jung, H., Hwang, Y., Park, Y. H., Ho, Y.-O.-C., Rengaraj, D., & Han, J. (2018) · Stem Cells and Development · doi:10.1089/scd.2018.0059 · Cited by 19
“The recent report showed that NANOG solely activated enhancers upstream of germ cell specifiers to generate PGC-like cells from epiblast-like cells, but not in ESCs in mouse [22], indicating that its function differs according to cell type.”
Dynamics of male canine germ cell development
de Souza, A. F., Pieri, N. C. G., Roballo, K., Bressan, F., Casals, J. B., Ambrósio, C., Perecin, F., & Martins, D. S. (2018) · PLoS ONE · doi:10.1371/journal.pone.0193026 · Cited by 19
Shirazi, R., Zarnani, A., Soleimani, M., Nayernia, K., & Kashani, I. R. (2017) · Microscopy research and technique (Print) · doi:10.1002/jemt.22812 · Cited by 19
Molecular versatility during pluripotency progression
Furlan, G., Huyghe, A., Combémorel, N., & Lavial, F. (2023) · Nature Communications · doi:10.1038/s41467-022-35775-4 · Cited by 18
Zhou, J.-L., He, H., Zhang, J.-J., Liu, X., Yao, W., Li, C., Xu, T., Yin, S.-Y., Wu, D., Dou, C.-L., et al. (2022) · Autophagy · doi:10.1080/15548627.2022.2055285 · Cited by 18
“Indeed, overexpression of NANOG in EpiLCs cannot maintain primed pluripotency, but promotes germline specification through binding active enhancers proximate to the Prdm14 and Prdm1 loci [57], suggesting the epigenetic resetting of NANOG binding profiles in EpiLCs was associated with germline specification.”
BackgroundEnhanced cultivation of chicken primordial germ cells
Dehdilani, N., Taemeh, S. Y., Rival-Gervier, S., Montillet, G., Kress, C., Jean, C., Goshayeshi, L., Dehghani, H., & Pain, B. (2023) · Scientific Reports · doi:10.1038/s41598-023-39536-1 · Cited by 17
Jung, K. M., Seo, M., Kim, Y. M., Kim, J. L., & Han, J. Y. (2021) · Frontiers in Cell and Developmental Biology · doi:10.3389/fcell.2021.791335 · Cited by 16
“Avian PGCs follow a preformation mode that is formed autonomously by maternally-inherited germ plasm, unlike mouse PGCs that are induced by signals received from surrounding tissues during development (Murakami et al., 2016; Jung et al., 2018).”
BackgroundZhang, M., Ji, J., Wang, X.-X., Zhang, X.-B., Zhang, Y., Li, Y.-T., Wang, X., Li, X.-F., Ban, Q., & Ye, S.-D. (2021) · Journal of Biological Chemistry · doi:10.1016/j.jbc.2021.101217 · Cited by 17
“However, induced expression of estrogen related receptor beta (Esrrb) can restore PGCLC numbers in Nanog-null mouse ESCs (34).”
BackgroundInfluentialZhao, R.-F., Zuo, Q.-S., Yuan, X.-Y., Jin, K., Jin, J., Ding, Y., Zhang, C., Li, T.-T., Jiang, J.-Y., Li, J., et al. (2021) · Nature Communications · doi:10.1038/s41467-021-23242-5 · Cited by 17
RetractedAgrawal, P., Blinka, S. M., Pulakanti, K., Reimer, M. H., Stelloh, C. T., Meyer, A. E., & Rao, S. (2021) · Journal of Biological Chemistry · doi:10.1074/jbc.ra120.015152 · Cited by 28
“Of importance, this DNA element may function as an alternative promoter in other contexts, such as alternative pluripotent states or in primordial germ cells, where Nanog is also expressed (37).”
BackgroundChoi, H.-J., Jin, S., Rengaraj, D., Kim, J. H., Pain, B., & Han, J. (2020) · Journal of Animal Science and Biotechnology · doi:10.1186/s40104-021-00563-5 · Cited by 17
Evolving Role of RING1 and YY1 Binding Protein in the Regulation of Germ-Cell-Specific Transcription
Bajusz, I., Henry, S., Sutus, E., Kovács, G. E., & Pirity, M. K. (2019) · Genes · doi:10.3390/genes10110941 · Cited by 20
“Ectopic induction of Nanog expression alone can trigger the differentiation of PGCLCs from ES cells and the activation of Prdm1 and Prdm14 transcription independently of the activation of BMP4 and RA signaling [61].”
BackgroundHu, B., Wang, R.-J., Wu, D., Long, R., Ruan, J., Jin, L., Ma, D., Sun, C.-Y., & Liao, S.-J. (2023) · Frontiers in Physiology · doi:10.3389/fphys.2023.1177443 · Cited by 16
“NANOG is upstream of PRDM14 and BLIMP1, and NANOG binds to their enhancers (Murakami et al., 2016).”
BackgroundGermline specification from pluripotent stem cells
Yao, C., Yao, R., Luo, H., & Shuai, L. (2022) · Stem cell research & therapeutics · doi:10.1186/s13287-022-02750-1 · Cited by 16
“T cells directly activate Blimp1 and Prdm14.”
BackgroundInfluentialWang, X., Liao, T.-T., Wan, C., Yang, X.-Y., Zhao, J., Fu, R., Yao, Z., Huang, Y., Shi, Y., Chang, G., et al. (2019) · PeerJ · doi:10.7717/peerj.6143 · Cited by 16
Tischler, J., Swank, Z., Hsiung, H.-A., Vianello, S., Lutolf, M., & Maerkl, S. J. (2022) · Cell Reports Methods · doi:10.1016/j.crmeth.2022.100244 · Cited by 15
Shono, M., Kishimoto, K., Hikabe, O., Hayashi, M., Semi, K., Takashima, Y., Sasaki, E., Kato, K., & Hayashi, K. (2023) · Scientific Reports · doi:10.1038/s41598-023-29850-z · Cited by 19
Nanog regulates Pou3f1 expression at the exit from pluripotency during gastrulation
Barral, A., Rollan, I., Sánchez-Iranzo, H., Jawaid, W., Badía-Careaga, C., Menchero, S., Gómez, M., Torroja, C., Sánchez-Cabo, F., Göttgens, B., et al. (2019) · Biology Open · doi:10.1242/bio.046367 · Cited by 14
“Sequence (right) of the genomic regions (A, -9kb; B, -11.5kb; C, +9kb) from the Pou3f1 locus showing NANOG binding as determined by ChIP-seq by from Murakami et al. (2016), and PCR genotyping of single embryos (left) showing the wild type and the corresponding deleted bands by gel electrophoresis.”
BackgroundMethodInfluentialOf mice and men: in vivo and in vitro studies of primordial germ cell specification
Kumar, D. L., & DeFalco, T. (2017) · Seminars in Reproductive Medicine · doi:10.1055/s-0037-1599085 · Cited by 14
Capturing Pluripotency and Beyond
Yeh, C.-Y., Huang, W., Chen, H., & Meir, Y. J. (2021) · Cells · doi:10.3390/cells10123558 · Cited by 16
Chimeric NANOG repressors inhibit glioblastoma growth in vivo in a context-dependent manner
Kuciak, M., Mas, C., Borges, I., Sánchez‐Gómez, P., & Altaba, A. R. I. (2019) · Scientific Reports · doi:10.1038/s41598-019-39473-y · Cited by 14
Yu, S. E., Kim, M. S., Park, S. H., Yoo, B., Kim, K.-H., & Jang, Y. (2017) · Cell Biochemistry and Function · doi:10.1002/cbf.3269 · Cited by 13
Yuan, F., Yang, J., Ma, F., Hu, Z., Malik, V., Zang, R., Li, D., Shi, X., Huang, X., Zhou, H., et al. (2025) · Nature Communications · doi:10.1038/s41467-025-57165-2 · Cited by 14
“Given that Tex10 interacts with and regulates Nanog in mouse ESCs 20 and that Nanog acts as a potent inducer of PGC speci fi cation from ESC-derived EpiLCs 7 , we next asked if Tex10 overexpression (OE) would promote PGC development.”
Tissue and cell interactions in mammalian PGC development
Cooke, C. B., & Moris, N. (2021) · Development · doi:10.1242/dev.200093 · Cited by 14
Shi, G. W., Bai, Y., Zhang, X., Su, J., Pang, J., He, Q., Zeng, P., Ding, J., Xiong, Y., Zhang, J., et al. (2021) · Protein & Cell · doi:10.1007/s13238-021-00884-1 · Cited by 15
“, Stella, BlIMP1, NANOS2) and is essential for germ cell lineage determination (Kehler et al., 2004; van den Berg et al., 2008; Murakami et al., 2016; Fang et al., 2018).”
BackgroundInfluentialRore, H., Owen, N., Piña-Aguilar, R., Docherty, K., & Sekido, R. (2021) · Communications Biology · doi:10.1038/s42003-021-02322-8 · Cited by 12
The transcription factor GABPA is a master regulator of naive pluripotency
Zhou, C.-J., Wang, M., Zhang, C., & Zhang, Y. (2025) · Nature Cell Biology · doi:10.1038/s41556-024-01554-0 · Cited by 11
Giovannantonio, L. D. D., Acampora, D., Omodei, D., Nigro, V., Barba, P., Barbieri, E., Chambers, I., & Simeone, A. (2021) · Development · doi:10.1242/dev.199166 · Cited by 11
YAP establishes epiblast responsiveness to inductive signals for germ cell fate
Kagiwada, S., Aramaki, S., Wu, G.-M., Shin, B., Kutejova, E., Obridge, D., Adachi, K., Wrana, J., Hübner, K., & Schöler, H. (2021) · Development · doi:10.1242/dev.199732 · Cited by 10
“We would like the reviewer to note that the previous study clearly indicates that integrinb3/ SSEA1 double-positive cells are equivalent to the cell fraction of functional PGCLCs with further differentiation competence, and the methodology is a reliable method for identifying specified PGCLCs in the relevant field (Ha…”
MethodTranscriptional and epigenetic control of germline competence and specification.
Bleckwehl, T., & Rada-Iglesias, Á. (2019) · Current Opinion in Cell Biology · doi:10.1016/j.ceb.2019.05.006 · Cited by 10
LIN28A activates the transcription of NANOG in dairy goat male germline stem cells
Ma, F.-L., Du, X., Wei, Y., Zhou, Z., Clotaire, D. Z. J., Li, N., Peng, S., Li, G.-P., & Hua, J. (2018) · Journal of Cellular Physiology · doi:10.1002/jcp.27593 · Cited by 10
“Furthermore, NANOG promoted the proliferation of the embryonic stem cells (ESCs) (Camp et al., 2009; Schuff et al., 2012) and alone induced germ cells to primed epiblast (Murakami et al., 2016).”
BackgroundDong, G., Shang, Z., Liu, L., Liu, C., Ge, Y., Wang, Q., Wu, L. L., Chen, F., Li, B., Liu, X., et al. (2017) · Bioscience Reports · doi:10.1042/bsr20170637 · Cited by 11
Li, Z., Fang, F., Long, Y., Zhao, Q., Wang, X., Ye, Z., Meng, T., Gu, X., Xiang, W., Xiong, C., et al. (2022) · Cell & Bioscience · doi:10.1186/s13578-022-00917-0 · Cited by 11
“Conversely, SOX17 is not critical for mPGC differentiation, and overexpression of Nanog along can induce PGCLCs in mouse EpiLCs (epiblast-like cells) [49], which illustrates that NANOG plays different roles in human and mouse PGC specification (Fig.”
BackgroundAdvances in Female Germ Cell Induction from Pluripotent Stem Cells
Gulimiheranmu, M., Wang, X., & Zhou, J.-M. (2021) · Stem Cells International · doi:10.1155/2021/8849230 · Cited by 9
“Migrating human PGCs mainly express early GCs markers BLIMP1, TFAP2C, and SSEA1 as well as pluripotency markers OCT4 and NANOG.”
BackgroundMethodInfluentialEnhancer priming by H3K4 methylation safeguards germline competence
Bleckwehl, T., Schaaf, K., Crispatzu, G., Respuela, P., Bartusel, M., Benson, L., Clark, S. J., Dorighi, K. M., Barral, A., Laugsch, M., et al. (2020) · bioRxiv · doi:10.1101/2020.07.07.192427 · Cited by 9
Segregation of the mouse germline and soma
Zhang, M., & Chambers, I. (2019) · Cell Cycle · doi:10.1080/15384101.2019.1672466 · Cited by 9
“In addition, regulatory interactions by pluripotency TFs on genes encoding the PGC TFs have been reported [35].”
BackgroundDNMT1 can induce primary germ layer differentiation through de novo DNA methylation
Ito, T., Kubiura-Ichimaru, M., Miura, F., Tajima, S., Surani, M., Ito, T., Yamaguchi, S., & Tada, M. (2024) · Genes to Cells · doi:10.1111/gtc.13130 · Cited by 8
“Furthermore, NANOG can induce the expression of PGC-related genes without BMP4 signaling (Murakami et al., 2016).”
BackgroundResultLi, Q., Huang, Y., Xu, J., Mao, F., Zhou, B. O., Sun, L., Basinski, B. W., Aksu, M., Liu, J., Dou, Y., et al. (2021) · Stem Cell Reports · doi:10.1016/j.stemcr.2021.10.002 · Cited by 8
“Growth advantage of mESCs by p53 deletion and upregulation of p53 target gene Nanog may account for enriched PGCLC differentiation (Lin et al., 2005; Murakami et al., 2016).”
Background
Metadata
- Authors
- Murakami, K., Günesdogan, U., Zylicz, J. J., Tang, W. W. C., Sengupta, R., Kobayashi, T., Kim, S., Butler, R., Dietmann, S., & Azim Surani, M.
- Affiliations
- Hokkaido University (JP)
- Wellcome/MRC Cambridge Stem Cell Institute (GB)
- University of Cambridge (GB)
- ORCID
- Venue
- Nature
- Published
- 2016-01
- Volume
- 529
- Issue
- 7586
- Pages
- 403-407
- ISSN
- 0028-0836 (ISSN-L)
- 1476-4687
- PMC
- PMC4724940
- OpenAlex
- W2238949050
- Cited by
- 180
- References
- 40
- Citations by year
- 2026: 4 · 2025: 11 · 2024: 12 · 2023: 15 · 2022: 19 · 2021: 28 · 2020: 15 · 2019: 26 · 2018: 22 · 2017: 17 · 2016: 11
- FWCI
- 13.17
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Licence
- www.springer.com/tdm (tdm)
- Locations
- Nature (publishedVersion)
- PubMed (publishedVersion)
- Europe PMC (PubMed Central) (acceptedVersion)
- PubMed Central (acceptedVersion)
- Apollo (University of Cambridge) (acceptedVersion)
- RePEc: Research Papers in Economics (submittedVersion)
- Research at the University of Copenhagen (University of Copenhagen) (submittedVersion)
- Funders
- Wellcome Trust (RG44593, 096738, 092096)
- Cancer Research UK (C6946 /A14492)
- Medical Research Council (MC_PC_12009)
- Japan Society for the Promotion of Science (15K18539)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Pluripotent Stem Cells Research
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Developmental Biology and Gene Regulation
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Epigenetics and DNA Methylation
- Keywords
- Nanog, epiblast-like cells, enhancer activation, chromatin accessibility, Blimp-1, PRDM14, TFAP2C, SOX2, pluripotency, naïve-to-primed transition, transcription factor binding sites
- MeSH terms
- Mouse Embryonic Stem Cells, Nanog Homeobox Protein, Positive Regulatory Domain I-Binding Factor 1, Animals, Cell Differentiation, Chromatin, DNA-Binding Proteins, Enhancer Elements, Genetic, Female, Germ Cells, Germ Layers, Male, Protein Binding, Transcription Factors, Fibroblast Growth Factor 2, RNA-Binding Proteins, Genome, Homeodomain Proteins, Gene Expression Regulation, Developmental, Activins, Pluripotent Stem Cells, Epigenesis, Genetic, Transcription Factor AP-2, Mice, Bone Morphogenetic Protein 4, SOXB1 Transcription Factors
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