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NANOG (Nanog homeobox)

Written2012-05Hubert Schorle, Daniel Nettersheim
University of Bonn Medical School, Institute of Pathology, Department of Developmental Pathology, Sigmund-Freud-Str. 25, 53127 Bonn, Germany

(Note : for Links provided by Atlas : click)

Identity

Alias_symbol (synonym)FLJ12581
FLJ40451
Other alias
HGNC (Hugo) NANOG
LocusID (NCBI) 79923
Atlas_Id 46540
Location 12p13.31  [Link to chromosome band 12p13]
Location_base_pair Starts at 7941992 and ends at 7948657 bp from pter ( according to hg19-Feb_2009)  [Mapping NANOG.png]

DNA/RNA

 
  Figure 1. (A) Genomic sequence of the human NANOG promoter region. Important regulatory elements, binding sites and CpG-dinucleotides are indicated in different colors. (B) Graphical illustration of the four different human NANOG transcripts.
Description The NANOG gene encompasses 8265 bp of DNA in humans and is located on the short arm of chromosome 12 (12p13.31) from 7940390 - 7948655. A 299 bp large NANOG promoter region (-264 to +35) upstream of exon 1 (Fig. 1A) contains five CpG-dinucleotides, which are subjected to DNA-methylation (Nettersheim et al., 2011a). Furthermore, this NANOG promoter region contains an OCT3/4-SOX2 binding motif, a TATA-box and binding sites for the transcription factors AP-2,SP1 and TFIID (Rodda et al., 2005; Nettersheim et al., 2011a) (PromoterScan 1.7). Functionality of this NANOG promoter element has been demonstrated by Luciferase-reporter experiments (Nettersheim et al., 2011a). A similar region (-229 to +193) has also been analysed by Wu et al. in mice (Wu da and Yao, 2005). There, binding sites for AP-1, SP1 and Oct3/4 could be found and functionality of this region in regulation of Nanog expression was demonstrated in F9 EC cells by Luciferase-assays.
Transcription The mRNA encoded by NANOG has four transcript variants (Fig. 1B). Transcript variants NANOG-001 (2101 bp mRNA) and -002 (870 bp mRNA) are known protein coding, while variant -004 (561 bp mRNA) is putatively protein coding and variant -003 is known as nonsense mediated decay. NANOG transcription can be regulated by binding of OCT3/4 and SOX2 to their binding motifs in the NANOG promoter (see Fig. 1A) (Nettersheim et al., 2011a; Kuroda et al., 2005; Rodda et al., 2005).
Pseudogene Ten NANOG-related nucleotide sequences (NANOGP2 - NANOGP11) plus one non-functional duplication (NANOGP1) have been discovered in the human genome (Booth and Holland, 2004). NANOGP2 - NANOGP11 are processed but are lacking introns, except NANOGP8 which is a retrogene.

Protein

 
  Figure 2: Graphical illustration of the NANOG protein sequence (A) and secondary structure motifs of the NANOG protein (B).
Description NANOG mRNA variants NANOG-001 and -002 encode for a protein with a length of 305 amino acid (aa) and 289 aa, respectively. NANOG-004 is putatively protein coding; translation would result in a 186 aa long protein.
NANOG-001, the 305 aa long protein with a molecular weight of 34.6 kDa is usually analyzed to study the role of NANOG (Fig. 2A). It consists of a Serine-, Threonine- and Proline-rich N-terminal region as well as eight W-repeats at its C-terminus (aa 104-151). The DNA-binding facilitating homeodomain spans from aa 95-155. Formation of secondary structures (helix, strand and turn) occurs mainly within the homeobox-coding region (Fig. 2B).
The NANOG protein is unstable with a half-life of 120 min in human ESCs (Ramakrishna et al., 2011).
Expression NANOG is expressed in undifferentiated embryonic stem cells (ESCs) (Ezeh et al., 2005), glioma stem cells (Zbinden et al., 2010), fetal gonocytes (Hart et al., 2005; Hoei-Hansen et al., 2005), carcinoma-in-situ (CIS) of the testis (Hart et al., 2005; Hoei-Hansen et al., 2005), seminoma (Ezeh et al., 2005; Hoei-Hansen et al., 2005), embryonal carcinoma (Hoei-Hansen et al., 2005) and mammary carcinoma / breast cancer (Ezeh et al., 2005). Ezeh et al. found NANOG expression in adult germ cells throughout germ cell differentiation (spermatogonia to spermatids) (Ezeh et al., 2005). In contrast to the data of Ezeh et al., Hoei-Hansen et al. were unable to detect NANOG protein in adult testis tissue (Hoei-Hansen et al., 2005). NANOG is also expressed in the seminoma cell line TCam-2 and the embryonal carcinoma cell lines 2102EP, NCCIT, Tera-1, Tera-2, NT2/D1 and 833Ke (Nettersheim et al., 2011a; Nettersheim et al., 2011b; Eckert et al., 2008; You et al., 2009; Deb-Rinker et al., 2005; Freberg et al., 2007).
In germ cell tumors (GCTs), NANOG expression is regulated by binding of OCT3/4 and SOX2 (SOX17 putatively in seminomas) to the NANOG promoter (Nettersheim et al., 2011a). Furthermore, in GCTs NANOG expression can be epigenetically suppressed by DNA-methylation of CpG-dinucleotides in the promoter (Nettersheim et al., 2011a). NANOG expression correlates to the differentiation status of GCTs, i.e. the more differentiated the GCT, the lower the NANOG expression (Nettersheim et al., 2011a).
Expression of the NANOG pseudogene NANOGP8 has been detected in uterine cervix tumor tissue, breast tumor tissue, urinary bladder tissue and smooth muscle cells as well as in the cell lines OS732, HepG3 and MCF-7 (Zhang et al., 2006; Ambady et al., 2010). Expression of pseudogenes NANOGP4 and NANOGP5 has been demonstrated in the cell lines OS732, HepG3 and MCF-7, too (Zhang et al., 2006).
Localisation Nuclear.
Function NANOG promotes self-renewal and maintains pluripotency in undifferentiated cells, like ESCs (Boyer et al., 2005; Wang et al., 2006). Thereby, NANOG acts in concert with the transcription factors SOX2, OCT3/4 (POU5F1) and FOXD3 (Pan and Thomson, 2007) (Wang et al., 2006). NANOG activates expression of SOX2 and OCT3/4 (Pan and Thomson, 2007).
A NANOG knock-down leads to differentiation of ESCs into the extraembryonic endoderm and trophectoderm lineages (Hyslop et al., 2005), demonstrating NANOG's important role in repression of differentiation. However, siRNA/shRNA-mediated reduction of the NANOG protein in the seminoma cell line TCam-2 is tolerated without induction of differentiation (Nettersheim et al., 2011b). It is assumed that the observed upregulation of OCT3/4 and SOX17 compensates for reduced NANOG levels in TCam-2 cells (Nettersheim et al., 2011b). NANOG-shRNA transduced cancer cells exhibit decreased long-term clonal and clonogenic growth, reduced proliferation and altered differentiation (Jeter et al., 2009). Overexpression of NANOG in ESC enables their feeder-cell-free propagation for multiple passages during which the cells remain pluripotent (Darr et al., 2006).
NANOG is able to modulate gliomasphere clonogenicity and CD133+ stem cell behaviour (Zbinden et al., 2010). There, NANOG is regulated by the Hedgehog-GLI signalling pathway. GLI1 requires NANOG activity to form a positive loop, which is negatively controlled by p53 and vice versa (Zbinden et al., 2010).
Besides SOX2 and OCT3/4, protein interaction databases predict interaction of NANOG with SALL4, KLF4, ZFP42, ISYNA1, LIN28, FAM48A, SMAD1, GATA6, RIF1, TRIM28, SMARCA4 and SMARCA2 (by similarity) (String 9.0, UniProt, I2D).
Homology NANOG and its pseudogenes (except NANOGP8) are present at their expected orthologous chromosomal positions in the chimpanzee genome when compared to the human genome (Fairbanks and Maughan, 2006). NANOGP8 is not found in the genome of the chimpanzee (Fairbanks and Maughan, 2006). The homeodomain of the human NANOG protein shares 87% identity to the murine NANOG homeodomain (Hart et al., 2004). A sequence comparison of murine, cow, chimp and human Nanog/NANOG promoters revealed that the CpG-dinucleotides are completely conserved in chimp and human (Nettersheim et al., 2011a).

Mutations

Somatic NANOG is localized on the short arm of chromosome 12, which is often gained in type II germ cell tumors, mostly due to isochromosome formation (i(12p)) (Looijenga et al., 2003).

Implicated in

Note
  
Entity Testicular germ cell tumors
Oncogenesis A consistent structural chromosomal abnormality in invasive testicular germ cell tumors are gains of the short arm of chromosome 12 (12p), due to isochromosome formation (Looijenga et al., 2003). This gain results in an increase in copy numbers of genes on 12p, which is speculated to lead to suppression of apoptosis and Sertoli-cell independency of CIS cells (Looijenga et al., 2003). NANOG is located in 12p and therefore amplified in invasive germ cell tumors. Increased NANOG copy numbers, resulting in overexpression of NANOG in seminomas and non-seminomas might lead to maintenance of self-renewal and mediates a pluripotency-like status. During differentiation of embryonal carcinomas into teratoma, choriocarcinoma and yolk-sac tumors the 12p gain is maintained, but NANOG expression is downregulated.
  

To be noted

In the adult testis, global DNA-methylation levels decrease from spermatogonia (hypermethylated) to mature sperm (hypomethylated). Interestingly the NANOG promoter is hypomethylated in spermatogonia and hypermethylated in sperm (Nettersheim et al., 2011a). In sperm, it is hypothesized that NANOG promoter methylation represents a way to epigenetically repress NANOG expression in order to control the pluripotency program and to prevent germ cell malignancies (Nettersheim et al., 2011a).
NANOG promoter hypomethylation could be found in seminoma, embryonal carcinoma, fetal gonocytes as well as fetal and adult spermatogonia. NANOG promoter hypermethylation was demonstrated in adult testis tissue, sperm, teratoma, choriocarcinoma, yolk-sac tumor and mixed non-seminoma (Nettersheim et al., 2011a). The cell lines TCam-2, 2102EP, Tera-1, Tera-2 and NT2/D1 harbor a hypomethylated NANOG promoter, while cell lines 833Ke, NCCIT and JKT-1 are hypermethylated at the NANOG promoter (Nettersheim et al., 2011a).
Upon differentiation of the seminoma cell line TCam-2 into a cell type resembling a mixed non-seminoma NANOG expression is downregulated (Nettersheim et al., 2011c).
NANOG has been shown to regulate self-renewal of cancer stem cells through the IGF pathway in hepatocellular carcinoma (Shan et al., 2012). Furthermore, increased expression of NANOG in cancer cells correlated to a worse clinical outcome. In addition, NANOG positive cancer stem cells were resistant to the therapeutic agents, sorafenib and cisplatin and have a high capacity for tumor invasion and metastasis.

Bibliography

Expression of NANOG and NANOGP8 in a variety of undifferentiated and differentiated human cells.
Ambady S, Malcuit C, Kashpur O, Kole D, Holmes WF, Hedblom E, Page RL, Dominko T.
Int J Dev Biol. 2010;54(11-12):1743-54.
PMID 21136380
 
Eleven daughters of NANOG.
Booth HA, Holland PW.
Genomics. 2004 Aug;84(2):229-38.
PMID 15233988
 
Core transcriptional regulatory circuitry in human embryonic stem cells.
Boyer LA, Lee TI, Cole MF, Johnstone SE, Levine SS, Zucker JP, Guenther MG, Kumar RM, Murray HL, Jenner RG, Gifford DK, Melton DA, Jaenisch R, Young RA.
Cell. 2005 Sep 23;122(6):947-56.
PMID 16153702
 
Overexpression of NANOG in human ES cells enables feeder-free growth while inducing primitive ectoderm features.
Darr H, Mayshar Y, Benvenisty N.
Development. 2006 Mar;133(6):1193-201.
PMID 16501172
 
Sequential DNA methylation of the Nanog and Oct-4 upstream regions in human NT2 cells during neuronal differentiation.
Deb-Rinker P, Ly D, Jezierski A, Sikorska M, Walker PR.
J Biol Chem. 2005 Feb 25;280(8):6257-60. Epub 2004 Dec 21.
PMID 15615706
 
TCam-2 but not JKT-1 cells resemble seminoma in cell culture.
Eckert D, Nettersheim D, Heukamp LC, Kitazawa S, Biermann K, Schorle H.
Cell Tissue Res. 2008 Feb;331(2):529-38. Epub 2007 Nov 15.
PMID 18008088
 
Human embryonic stem cell genes OCT4, NANOG, STELLAR, and GDF3 are expressed in both seminoma and breast carcinoma.
Ezeh UI, Turek PJ, Reijo RA, Clark AT.
Cancer. 2005 Nov 15;104(10):2255-65.
PMID 16228988
 
Evolution of the NANOG pseudogene family in the human and chimpanzee genomes.
Fairbanks DJ, Maughan PJ.
BMC Evol Biol. 2006 Feb 9;6:12.
PMID 16469101
 
Epigenetic reprogramming of OCT4 and NANOG regulatory regions by embryonal carcinoma cell extract.
Freberg CT, Dahl JA, Timoskainen S, Collas P.
Mol Biol Cell. 2007 May;18(5):1543-53. Epub 2007 Feb 21.
PMID 17314394
 
The pluripotency homeobox gene NANOG is expressed in human germ cell tumors.
Hart AH, Hartley L, Parker K, Ibrahim M, Looijenga LH, Pauchnik M, Chow CW, Robb L.
Cancer. 2005 Nov 15;104(10):2092-8.
PMID 16206293
 
Stem cell pluripotency factor NANOG is expressed in human fetal gonocytes, testicular carcinoma in situ and germ cell tumours.
Hoei-Hansen CE, Almstrup K, Nielsen JE, Brask Sonne S, Graem N, Skakkebaek NE, Leffers H, Rajpert-De Meyts E.
Histopathology. 2005 Jul;47(1):48-56.
PMID 15982323
 
Downregulation of NANOG induces differentiation of human embryonic stem cells to extraembryonic lineages.
Hyslop L, Stojkovic M, Armstrong L, Walter T, Stojkovic P, Przyborski S, Herbert M, Murdoch A, Strachan T, Lako M.
Stem Cells. 2005 Sep;23(8):1035-43. Epub 2005 Jun 27.
PMID 15983365
 
Functional evidence that the self-renewal gene NANOG regulates human tumor development.
Jeter CR, Badeaux M, Choy G, Chandra D, Patrawala L, Liu C, Calhoun-Davis T, Zaehres H, Daley GQ, Tang DG.
Stem Cells. 2009 May;27(5):993-1005.
PMID 19415763
 
Octamer and Sox elements are required for transcriptional cis regulation of Nanog gene expression.
Kuroda T, Tada M, Kubota H, Kimura H, Hatano SY, Suemori H, Nakatsuji N, Tada T.
Mol Cell Biol. 2005 Mar;25(6):2475-85.
PMID 15743839
 
Role of gain of 12p in germ cell tumour development.
Looijenga LH, Zafarana G, Grygalewicz B, Summersgill B, Debiec-Rychter M, Veltman J, Schoenmakers EF, Rodriguez S, Jafer O, Clark J, van Kessel AG, Shipley J, van Gurp RJ, Gillis AJ, Oosterhuis JW.
APMIS. 2003 Jan;111(1):161-71; discussion 172-3.
PMID 12752258
 
TGF-b1, EGF and FGF4 synergistically induce differentiation of the seminoma cell line TCam-2 into a cell type resembling mixed non-seminoma.
Nettersheim D, Gillis AJ, Looijenga LH, Schorle H.
Int J Androl. 2011c Aug;34(4 Pt 2):e189-203. doi: 10.1111/j.1365-2605.2011.01172.x. Epub 2011 Jun 8.
PMID 21649665
 
Nanog and transcriptional networks in embryonic stem cell pluripotency.
Pan G, Thomson JA.
Cell Res. 2007 Jan;17(1):42-9.
PMID 17211451
 
PEST motif sequence regulating human NANOG for proteasomal degradation.
Ramakrishna S, Suresh B, Lim KH, Cha BH, Lee SH, Kim KS, Baek KH.
Stem Cells Dev. 2011 Sep;20(9):1511-9. Epub 2011 Mar 12.
PMID 21299413
 
Transcriptional regulation of nanog by OCT4 and SOX2.
Rodda DJ, Chew JL, Lim LH, Loh YH, Wang B, Ng HH, Robson P.
J Biol Chem. 2005 Jul 1;280(26):24731-7. Epub 2005 Apr 27.
PMID 15860457
 
Nanog regulates self-renewal of cancer stem cell through IGF pathway in human hepatocellular carcinoma.
Shan J, Shen J, Liu L, Xia F, Xu C, Duan G, Xu Y, Ma Q, Yang Z, Zhang Q, Ma L, Liu J, Xu S, Yan X, Bie P, Cui Y, Bian XW, Qian C.
Hepatology. 2012 Apr 2. doi: 10.1002/hep.25745. [Epub ahead of print]
PMID 22473773
 
A protein interaction network for pluripotency of embryonic stem cells.
Wang J, Rao S, Chu J, Shen X, Levasseur DN, Theunissen TW, Orkin SH.
Nature. 2006 Nov 16;444(7117):364-8. Epub 2006 Nov 8.
PMID 17093407
 
Isolation and characterization of the murine Nanog gene promoter.
Wu da Y, Yao Z.
Cell Res. 2005 May;15(5):317-24.
PMID 15916719
 
Depletion of embryonic stem cell signature by histone deacetylase inhibitor in NCCIT cells: involvement of Nanog suppression.
You JS, Kang JK, Seo DW, Park JH, Park JW, Lee JC, Jeon YJ, Cho EJ, Han JW.
Cancer Res. 2009 Jul 15;69(14):5716-25. Epub 2009 Jun 30.
PMID 19567677
 
NANOG regulates glioma stem cells and is essential in vivo acting in a cross-functional network with GLI1 and p53.
Zbinden M, Duquet A, Lorente-Trigos A, Ngwabyt SN, Borges I, Ruiz i Altaba A.
EMBO J. 2010 Aug 4;29(15):2659-74. Epub 2010 Jun 25.
PMID 20581802
 
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FEBS J. 2006 Apr;273(8):1723-30.
PMID 16623708
 

Citation

This paper should be referenced as such :
Schorle, H ; Nettersheim, D
NANOG (Nanog homeobox)
Atlas Genet Cytogenet Oncol Haematol. 2012;16(10):727-731.
Free journal version : [ pdf ]   [ DOI ]
On line version : http://AtlasGeneticsOncology.org/Genes/NANOGID46540ch12p13.html


Other Leukemias implicated (Data extracted from papers in the Atlas) [ 1 ]
  t(10;11)(q22;q23) KMT2A/TET1


Other Solid tumors implicated (Data extracted from papers in the Atlas) [ 1 ]
  Testis: Spermatocytic seminoma


External links

Nomenclature
HGNC (Hugo)NANOG   20857
Cards
AtlasNANOGID46540ch12p13
Entrez_Gene (NCBI)NANOG  79923  Nanog homeobox
Aliases
GeneCards (Weizmann)NANOG
Ensembl hg19 (Hinxton)ENSG00000111704 [Gene_View]  chr12:7941992-7948657 [Contig_View]  NANOG [Vega]
Ensembl hg38 (Hinxton)ENSG00000111704 [Gene_View]  chr12:7941992-7948657 [Contig_View]  NANOG [Vega]
ICGC DataPortalENSG00000111704
TCGA cBioPortalNANOG
AceView (NCBI)NANOG
Genatlas (Paris)NANOG
WikiGenes79923
SOURCE (Princeton)NANOG
Genetics Home Reference (NIH)NANOG
Genomic and cartography
GoldenPath hg19 (UCSC)NANOG  -     chr12:7941992-7948657 +  12p13.31   [Description]    (hg19-Feb_2009)
GoldenPath hg38 (UCSC)NANOG  -     12p13.31   [Description]    (hg38-Dec_2013)
EnsemblNANOG - 12p13.31 [CytoView hg19]  NANOG - 12p13.31 [CytoView hg38]
Mapping of homologs : NCBINANOG [Mapview hg19]  NANOG [Mapview hg38]
OMIM607937   
Gene and transcription
Genbank (Entrez)AB093576 AI656990 AK022643 AK290896 AY230262
RefSeq transcript (Entrez)NM_001297698 NM_024865
RefSeq genomic (Entrez)NC_000012 NC_018923 NT_009714 NW_004929383
Consensus coding sequences : CCDS (NCBI)NANOG
Cluster EST : UnigeneHs.635882 [ NCBI ]
CGAP (NCI)Hs.635882
Alternative Splicing GalleryENSG00000111704
Gene ExpressionNANOG [ NCBI-GEO ]   NANOG [ EBI - ARRAY_EXPRESS ]   NANOG [ SEEK ]   NANOG [ MEM ]
Gene Expression Viewer (FireBrowse)NANOG [ Firebrowse - Broad ]
SOURCE (Princeton)Expression in : [Datasets]   [Normal Tissue Atlas]  [carcinoma Classsification]  [NCI60]
GenevisibleExpression in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
BioGPS (Tissue expression)79923
GTEX Portal (Tissue expression)NANOG
Protein : pattern, domain, 3D structure
UniProt/SwissProtQ9H9S0   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
NextProtQ9H9S0  [Sequence]  [Exons]  [Medical]  [Publications]
With graphics : InterProQ9H9S0
Splice isoforms : SwissVarQ9H9S0
PhosPhoSitePlusQ9H9S0
Domaine pattern : Prosite (Expaxy)HOMEOBOX_1 (PS00027)    HOMEOBOX_2 (PS50071)   
Domains : Interpro (EBI)Homeobox_CS    Homeobox_dom    Homeodomain-like   
Domain families : Pfam (Sanger)Homeobox (PF00046)   
Domain families : Pfam (NCBI)pfam00046   
Domain families : Smart (EMBL)HOX (SM00389)  
Conserved Domain (NCBI)NANOG
DMDM Disease mutations79923
Blocks (Seattle)NANOG
PDB (SRS)2KT0   
PDB (PDBSum)2KT0   
PDB (IMB)2KT0   
PDB (RSDB)2KT0   
Structural Biology KnowledgeBase2KT0   
SCOP (Structural Classification of Proteins)2KT0   
CATH (Classification of proteins structures)2KT0   
SuperfamilyQ9H9S0
Human Protein AtlasENSG00000111704
Peptide AtlasQ9H9S0
HPRD06397
IPIIPI00552676   
Protein Interaction databases
DIP (DOE-UCLA)Q9H9S0
IntAct (EBI)Q9H9S0
FunCoupENSG00000111704
BioGRIDNANOG
STRING (EMBL)NANOG
ZODIACNANOG
Ontologies - Pathways
QuickGOQ9H9S0
Ontology : AmiGOendodermal cell fate specification  DNA binding  transcription factor activity, sequence-specific DNA binding  transcription corepressor activity  protein binding  nucleus  nucleus  nucleoplasm  nucleolus  transcription, DNA-templated  regulation of transcription, DNA-templated  cell proliferation  regulation of gene expression  stem cell population maintenance  cell differentiation  somatic stem cell population maintenance  somatic stem cell population maintenance  sequence-specific DNA binding  regulation of cell differentiation  positive regulation of transcription from RNA polymerase II promoter  negative regulation of nucleic acid-templated transcription  
Ontology : EGO-EBIendodermal cell fate specification  DNA binding  transcription factor activity, sequence-specific DNA binding  transcription corepressor activity  protein binding  nucleus  nucleus  nucleoplasm  nucleolus  transcription, DNA-templated  regulation of transcription, DNA-templated  cell proliferation  regulation of gene expression  stem cell population maintenance  cell differentiation  somatic stem cell population maintenance  somatic stem cell population maintenance  sequence-specific DNA binding  regulation of cell differentiation  positive regulation of transcription from RNA polymerase II promoter  negative regulation of nucleic acid-templated transcription  
Pathways : KEGGProteoglycans in cancer   
REACTOMEQ9H9S0 [protein]
REACTOME Pathways2892245 [pathway]   2892247 [pathway]   452723 [pathway]   6785807 [pathway]   
NDEx NetworkNANOG
Atlas of Cancer Signalling NetworkNANOG
Wikipedia pathwaysNANOG
Orthology - Evolution
OrthoDB79923
GeneTree (enSembl)ENSG00000111704
Phylogenetic Trees/Animal Genes : TreeFamNANOG
HOVERGENQ9H9S0
HOGENOMQ9H9S0
Homologs : HomoloGeneNANOG
Homology/Alignments : Family Browser (UCSC)NANOG
Gene fusions - Rearrangements
Polymorphisms : SNP and Copy number variants
NCBI Variation ViewerNANOG [hg38]
dbSNP Single Nucleotide Polymorphism (NCBI)NANOG
dbVarNANOG
ClinVarNANOG
1000_GenomesNANOG 
Exome Variant ServerNANOG
ExAC (Exome Aggregation Consortium)NANOG (select the gene name)
Genetic variants : HAPMAP79923
Genomic Variants (DGV)NANOG [DGVbeta]
DECIPHER (Syndromes)12:7941992-7948657  ENSG00000111704
CONAN: Copy Number AnalysisNANOG 
Mutations
ICGC Data PortalNANOG 
TCGA Data PortalNANOG 
Broad Tumor PortalNANOG
OASIS PortalNANOG [ Somatic mutations - Copy number]
Somatic Mutations in Cancer : COSMICNANOG  [overview]  [genome browser]  [tissue]  [distribution]  
Mutations and Diseases : HGMDNANOG
LOVD (Leiden Open Variation Database)Whole genome datasets
LOVD (Leiden Open Variation Database)LOVD - Leiden Open Variation Database
LOVD (Leiden Open Variation Database)LOVD 3.0 shared installation
BioMutasearch NANOG
DgiDB (Drug Gene Interaction Database)NANOG
DoCM (Curated mutations)NANOG (select the gene name)
CIViC (Clinical Interpretations of Variants in Cancer)NANOG (select a term)
intoGenNANOG
NCG5 (London)NANOG
Cancer3DNANOG(select the gene name)
Impact of mutations[PolyPhen2] [SIFT Human Coding SNP] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
Diseases
OMIM607937   
Orphanet
MedgenNANOG
Genetic Testing Registry NANOG
NextProtQ9H9S0 [Medical]
TSGene79923
GENETestsNANOG
Huge Navigator NANOG [HugePedia]
snp3D : Map Gene to Disease79923
BioCentury BCIQNANOG
ClinGenNANOG
Clinical trials, drugs, therapy
Chemical/Protein Interactions : CTD79923
Chemical/Pharm GKB GenePA134864904
Clinical trialNANOG
Miscellaneous
canSAR (ICR)NANOG (select the gene name)
Other databaseString 9.0
Other databaseI2D
Other databasePromoterScan 1.7
Probes
Litterature
PubMed202 Pubmed reference(s) in Entrez
GeneRIFsGene References Into Functions (Entrez)
CoreMineNANOG
EVEXNANOG
GoPubMedNANOG
iHOPNANOG
REVIEW articlesautomatic search in PubMed
Last year publicationsautomatic search in PubMed

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