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AXIN2 (axin 2)

Identity

Other namesaxil (mostly in rat)
conductin
DKFZp781B0869 (single database entry only)
MGC126582 (single database entry only)
HGNC (Hugo) AXIN2
LocusID (NCBI) 8313
Location 17q24.1
Location_base_pair Starts at 63524683 and ends at 63557740 bp from pter ( according to hg19-Feb_2009)  [Mapping]

DNA/RNA

 
  The 5' end of the human AXIN2 gene. An alignment of human genomic DNA (top line) with the 5' end of different Axin2 mRNA variants. Exons are shown as boxes (non-coding: filled; coding: open) and the translational start codon is marked (ATG).
Description The AXIN2 gene spans about 35 kbp including 10 coding exons and 3 non-coding 5' exons (E0a, 0b and 0c; see above). Nearby genes: about 70 kbp upstream is CCDC46 (coiled-coil domain containing 46), about 300 kbp downstream is RGS9 (regulator of G-protein signalling 9). In addition, there is a putative gene that overlaps the AXIN2 non-coding 5' exons and coding exon 1 (E1) and is transcribed from the same strand (Gnomon model hmm119498); there is no published data on whether this is actually expressed.
Transcription Transcription occurs from three separate promoters leading to initiation at each of the three non-coding 5' exons. mRNAs are spliced so that that each non-coding exon is expressed separately, rather than in combinations. It is unclear whether transcription can initiate at the first coding exon (E1). Promoters can be activated by TCF transcription factors binding at multiple sites and by E2F1 binding at up to 4 sites, although E2F1 can also induce transcription in the absence of consensus sites. It has been reported that exon 6 can be omitted in an alternatively-spliced form.
Pseudogene None identified

Protein

 
Description Human Axin2 is an 843 amino acids protein (777 amino acids from delta exon 6 mRNAs) containing an RGS domain (regulator of G protein signalling; amino acids 81-200), a GSK-3 beta binding domain (amino acids 327-413), a beta-catenin binding domain (amino acids 413-476), and a DIX domain (domain in dishevelled and axin; amino acids 761-843).
Expression Expression appears to be ubiquitous in adult tissues (although at differing levels), but is limited to specific regions during embryonic development. Expression is regulated at multiple levels including transcription, mRNA stability, translation and protein stability.
Localisation Axin2 protein has been localised to the cytoplasm, the nucleus and the mitotic spindle.
Function Molecular functions :
1) Axin2 acts as a negative regulator of canonical Wnt/TCF signalling by enhancing formation of the beta-catenin destruction complex. Since expression of Axin2 is itself activated by canonical Wnt/TCF signalling, this results in a negative feedback-loop that restricts TCF activity.
2) Axin2 may influence TCF activity by re-localising beta-catenin to the cytoplasm.
3) Activity of the GSK-3 beta target snail1 can be regulated by Axin2's ability to influence the nucleo-cytoplasmic localisation of GSK-3 beta.
4) Axin2 binds polo-like kinase 1 (PLK1) during mitosis and influences the accuracy of chromosome segregation.
Cellular/physiological functions :
1) Axin2 expression oscillates during early embryogenesis in response to Wnt3a this is required to achieve correct the temporal TCF activity to allow somatogenesis.
2) A requirement for Axin2 for correct calvarial morphogenesis and craniosynostosis has been revealed in Axin2 -/- mice.
3) Axin2 appears to act as a tumour suppressor, and somatic mutations have been seen in many different tumour types (see below).
Homology Axin2 is 44% identical to axin in mice and knock-in experiments suggest that the proteins can be functionally equivalent.

Mutations

Note A large number of different mutations in the AXIN2 gene have been identified. In many cases (but not all) these lead to premature translational termination and protein truncation. Truncated Axin2 protein is more stable than the wild type, while there has been speculation that the mRNA may be less stable.
Germinal Heterozygous germ line mutations in exon 7 that lead to premature termination are associated with familial tooth agenesis and a predisposition to colorectal cancer. Further germ line polymorphisms associated with familial tooth agenesis have been identified in exons 2 and 7. A polymorphism within exon 1 has been identified that is associated with risk of lung cancer. Many other polymorphisms that have yet to be associated with any function have been detected.
Somatic The genomic region containing the AXIN2 gene shows loss of heterozygosity and re-arrangements in a variety of cancers. In addition somatic point mutations and deletions have been identified in colorectal cancer, hepatocellular carcinomas, ovarian endometrioid adenocarcinomas and hepatoblastomas. Many of these mutations/deletions result in translation of truncated proteins that are likely to be functionally inactive, although one report has suggested that these truncated proteins have a dominant negative activity.

Implicated in

Entity Colorectal cancer (CRC)
Oncogenesis Axin2 is often over-expressed in CRC as a result of the deregulation of canonical Wnt/beta-catenin signalling that is an early event in CRC development (usually caused by mutations/deletions in APC or beta-catenin). Somatic inactivating mutations within Axin2 have been reported in CRC and theoretically these could contribute to further deregulation of Wnt/beta-catenin suggesting that Axin2 is a tumour suppressor. However mutations have only been seen in microsatellite unstable tumours and often within regions of mono-nucleotide repeats (exon 7), hence whether Axin2 mutations are cause or effect in these tumours remains undetermined. In support of Axin2's role as a tumour suppressor are observations that Axin2 is silenced by promoter methylation in many microsatellite unstable tumours. As discussed above, heterozygotes for some germ line mutations in AXIN2 are predisposed to CRC although this seems to be involved with only a very small proportion of familial colorectal cancer.
  
Entity Other cancers (hepatocellular carcinomas, hepatoblastomas, ovarian endometrioid adenocarcinomas)
Oncogenesis Somatic mutations in Axin2 have been detected in a range of cancer types. It is usually assumed that these lead to partial inactivation of Axin2 function thereby deregulation of canonical Wnt/beta-catenin signalling. In most cases this has not formally been demonstrated, and the contribution of Axin2 mutations to any putative change in Wnt/beta-catenin activity and to the development of these cancers remains mostly unclear.
  
Entity Familial Tooth Agenesis (see above)
  

External links

Nomenclature
HGNC (Hugo)AXIN2   904
Cards
AtlasAXIN2ID456ch17q24
Entrez_Gene (NCBI)AXIN2  8313  axin 2
GeneCards (Weizmann)AXIN2
Ensembl hg19 (Hinxton)ENSG00000168646 [Gene_View]  chr17:63524683-63557740 [Contig_View]  AXIN2 [Vega]
Ensembl hg38 (Hinxton)ENSG00000168646 [Gene_View]  chr17:63524683-63557740 [Contig_View]  AXIN2 [Vega]
ICGC DataPortalENSG00000168646
cBioPortalAXIN2
AceView (NCBI)AXIN2
Genatlas (Paris)AXIN2
WikiGenes8313
SOURCE (Princeton)AXIN2
Genomic and cartography
GoldenPath hg19 (UCSC)AXIN2  -     chr17:63524683-63557740 -  17q23-q24   [Description]    (hg19-Feb_2009)
GoldenPath hg38 (UCSC)AXIN2  -     17q23-q24   [Description]    (hg38-Dec_2013)
EnsemblAXIN2 - 17q23-q24 [CytoView hg19]  AXIN2 - 17q23-q24 [CytoView hg38]
Mapping of homologs : NCBIAXIN2 [Mapview hg19]  AXIN2 [Mapview hg38]
OMIM604025   608615   
Gene and transcription
Genbank (Entrez)AB052751 AF078165 AF205888 AK025718 AK315303
RefSeq transcript (Entrez)NM_004655
RefSeq genomic (Entrez)AC_000149 NC_000017 NC_018928 NG_012142 NT_010783 NW_001838454 NW_004929407
Consensus coding sequences : CCDS (NCBI)AXIN2
Cluster EST : UnigeneHs.156527 [ NCBI ]
CGAP (NCI)Hs.156527
Alternative Splicing : Fast-db (Paris)GSHG0013671
Alternative Splicing GalleryENSG00000168646
Gene ExpressionAXIN2 [ NCBI-GEO ]     AXIN2 [ SEEK ]   AXIN2 [ MEM ]
SOURCE (Princeton)Expression in : [Normal Tissue Atlas]  [carcinoma Classsification]  [NCI60]
Protein : pattern, domain, 3D structure
UniProt/SwissProtQ9Y2T1 (Uniprot)
NextProtQ9Y2T1  [Medical]
With graphics : InterProQ9Y2T1
Splice isoforms : SwissVarQ9Y2T1 (Swissvar)
Domaine pattern : Prosite (Expaxy)DIX (PS50841)    RGS (PS50132)   
Domains : Interpro (EBI)Axin_b-cat-bd    DIX    Regulat_G_prot_signal_dom1    Regulat_G_prot_signal_superfam    RGS_dom    Ubiquitin-rel_dom   
Related proteins : CluSTrQ9Y2T1
Domain families : Pfam (Sanger)Axin_b-cat_bind (PF08833)    DIX (PF00778)    RGS (PF00615)   
Domain families : Pfam (NCBI)pfam08833    pfam00778    pfam00615   
Domain families : Smart (EMBL)DAX (SM00021)  RGS (SM00315)  
DMDM Disease mutations8313
Blocks (Seattle)Q9Y2T1
Human Protein AtlasENSG00000168646
Peptide AtlasQ9Y2T1
HPRD04935
IPIIPI00294274   IPI00788621   IPI01009779   
Protein Interaction databases
DIP (DOE-UCLA)Q9Y2T1
IntAct (EBI)Q9Y2T1
FunCoupENSG00000168646
BioGRIDAXIN2
IntegromeDBAXIN2
STRING (EMBL)AXIN2
Ontologies - Pathways
QuickGOQ9Y2T1
Ontology : AmiGOpositive regulation of protein phosphorylation  intramembranous ossification  chondrocyte differentiation involved in endochondral bone morphogenesis  signal transducer activity  GTPase activator activity  protein binding  nucleus  cytoplasm  centrosome  cytoplasmic microtubule  plasma membrane  cell cortex  beta-catenin binding  beta-catenin binding  cell proliferation  negative regulation of cell proliferation  dorsal/ventral axis specification  positive regulation of epithelial to mesenchymal transition  positive regulation of cell death  postsynaptic density  cytoplasmic membrane-bounded vesicle  enzyme binding  protein kinase binding  bone mineralization  beta-catenin destruction complex  ubiquitin protein ligase binding  regulation of mismatch repair  cellular protein localization  negative regulation of catenin import into nucleus  termination of G-protein coupled receptor signaling pathway  odontogenesis  positive regulation of GTPase activity  maintenance of DNA repeat elements  negative regulation of osteoblast differentiation  mRNA stabilization  regulation of chondrocyte development  armadillo repeat domain binding  I-SMAD binding  regulation of centromeric sister chromatid cohesion  cellular response to organic cyclic compound  negative regulation of canonical Wnt signaling pathway  negative regulation of canonical Wnt signaling pathway  Wnt signaling pathway involved in somitogenesis  
Ontology : EGO-EBIpositive regulation of protein phosphorylation  intramembranous ossification  chondrocyte differentiation involved in endochondral bone morphogenesis  signal transducer activity  GTPase activator activity  protein binding  nucleus  cytoplasm  centrosome  cytoplasmic microtubule  plasma membrane  cell cortex  beta-catenin binding  beta-catenin binding  cell proliferation  negative regulation of cell proliferation  dorsal/ventral axis specification  positive regulation of epithelial to mesenchymal transition  positive regulation of cell death  postsynaptic density  cytoplasmic membrane-bounded vesicle  enzyme binding  protein kinase binding  bone mineralization  beta-catenin destruction complex  ubiquitin protein ligase binding  regulation of mismatch repair  cellular protein localization  negative regulation of catenin import into nucleus  termination of G-protein coupled receptor signaling pathway  odontogenesis  positive regulation of GTPase activity  maintenance of DNA repeat elements  negative regulation of osteoblast differentiation  mRNA stabilization  regulation of chondrocyte development  armadillo repeat domain binding  I-SMAD binding  regulation of centromeric sister chromatid cohesion  cellular response to organic cyclic compound  negative regulation of canonical Wnt signaling pathway  negative regulation of canonical Wnt signaling pathway  Wnt signaling pathway involved in somitogenesis  
Pathways : KEGGWnt signaling pathway    Hippo signaling pathway    Pathways in cancer    Colorectal cancer    Endometrial cancer    Basal cell carcinoma   
Protein Interaction DatabaseAXIN2
DoCM (Curated mutations)AXIN2
Wikipedia pathwaysAXIN2
Gene fusion - rearrangements
Polymorphisms : SNP, variants
NCBI Variation ViewerAXIN2 [hg38]
dbSNP Single Nucleotide Polymorphism (NCBI)AXIN2
dbVarAXIN2
ClinVarAXIN2
1000_GenomesAXIN2 
Exome Variant ServerAXIN2
SNP (GeneSNP Utah)AXIN2
SNP : HGBaseAXIN2
Genetic variants : HAPMAPAXIN2
Genomic VariantsAXIN2  AXIN2 [DGVbeta]
Mutations
ICGC Data PortalENSG00000168646 
Somatic Mutations in Cancer : COSMICAXIN2 
CONAN: Copy Number AnalysisAXIN2 
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
LOVD (Leiden Open Variation Database)Zhejiang University Center for Genetic and Genomic Medicine (ZJU-CGGM)
Impact of mutations[PolyPhen2] [SIFT Human Coding SNP] [Buck Institute : MutDB] [Mutation Assessor] 
Diseases
DECIPHER (Syndromes)17:63524683-63557740
Mutations and Diseases : HGMDAXIN2
OMIM604025    608615   
MedgenAXIN2
NextProtQ9Y2T1 [Medical]
GENETestsAXIN2
Disease Genetic AssociationAXIN2
Huge Navigator AXIN2 [HugePedia]  AXIN2 [HugeCancerGEM]
snp3D : Map Gene to Disease8313
DGIdb (Drug Gene Interaction db)AXIN2
General knowledge
Homologs : HomoloGeneAXIN2
Homology/Alignments : Family Browser (UCSC)AXIN2
Phylogenetic Trees/Animal Genes : TreeFamAXIN2
Chemical/Protein Interactions : CTD8313
Chemical/Pharm GKB GenePA25196
Clinical trialAXIN2
Cancer Resource (Charite)ENSG00000168646
Other databases
Other databasehttp://cancergenome.broadinstitute.org/index.php?tgene=AXIN2
Probes
Litterature
PubMed83 Pubmed reference(s) in Entrez
CoreMineAXIN2
GoPubMedAXIN2
iHOPAXIN2

Bibliography

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Subcellular distribution of Wnt pathway proteins in normal and neoplastic colon.
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PMID 12072559
 
Wnt/beta-catenin/Tcf signaling induces the transcription of Axin2, a negative regulator of the signaling pathway.
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PMID 11809808
 
Activation of AXIN2 expression by beta-catenin-T cell factor. A feedback repressor pathway regulating Wnt signaling.
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PMID 11940574
 
Negative feedback loop of Wnt signaling through upregulation of conductin/axin2 in colorectal and liver tumors.
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PMID 11809809
 
Mutational spectrum of beta-catenin, AXIN1, and AXIN2 in hepatocellular carcinomas and hepatoblastomas.
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Oncogene. 2002 ; 21 (31) : 4863-4871.
PMID 12101426
 
Wnt3a plays a major role in the segmentation clock controlling somitogenesis.
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Immunohistochemical analysis and mutational analyses of beta-catenin, Axin family and APC genes in hepatocellular carcinomas.
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PMID 15067328
 
Mutations and elevated transcriptional activity of conductin (AXIN2) in hepatoblastomas.
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The Journal of pathology. 2004 ; 204 (5) : 546-554.
PMID 15538750
 
Mutations in AXIN2 cause familial tooth agenesis and predispose to colorectal cancer.
Lammi L, Arte S, Somer M, Jarvinen H, Lahermo P, Thesleff I, Pirinen S, Nieminen P
American journal of human genetics. 2004 ; 74 (5) : 1043-1050.
PMID 15042511
 
Mouse axin and axin2/conductin proteins are functionally equivalent in vivo.
Chia IV, Costantini F
Molecular and cellular biology. 2005 ; 25 (11) : 4371-4376.
PMID 15899843
 
Cross-talk between pRb/E2F and Wnt/beta-catenin pathways: E2F1 induces axin2 leading to repression of Wnt signalling and to increased cell death.
Hughes TA, Brady HJ
Experimental cell research. 2005 ; 303 (1) : 32-46.
PMID 15572025
 
Mouse axin and axin2/conductin proteins are functionally equivalent in vivo.
Chia IV, Costantini F
Molecular and cellular biology. 2005 ; 25 (11) : 4371-4376.
PMID 15899843
 
Expression of axin2 is regulated by the alternative 5'-untranslated regions of its mRNA.
Hughes TA, Brady HJ
The Journal of biological chemistry. 2005 ; 280 (9) : 8581-8588.
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Germline mutations of AXIN2 are not associated with nonsyndromic colorectal cancer.
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Human mutation. 2005 ; 25 (5) : 498-500.
PMID 15841489
 
The links between axin and carcinogenesis.
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Genetic and epigenetic changes of components affecting the WNT pathway in colorectal carcinomas stratified by microsatellite instability.
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Neoplasia (New York, N.Y.). 2005 ; 7 (2) : 99-108.
PMID 15802015
 
The role of Axin2 in calvarial morphogenesis and craniosynostosis.
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PMID 15790973
 
Aberrant Wnt/beta-catenin signaling can induce chromosomal instability in colon cancer.
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PMID 16815967
 
Regulation of axin2 expression at the levels of transcription, translation and protein stability in lung and colon cancer.
Hughes TA, Brady HJ
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PMID 15885887
 
Single nucleotide polymorphism of the AXIN2 gene is preferentially associated with human lung cancer risk in a Japanese population.
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Nucleo-cytoplasmic distribution of beta-catenin is regulated by retention.
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Contributor(s)

Written01-2007Thomas A Hughes

Citation

This paper should be referenced as such :
Hughes, TA
AXIN2 (axin 2)
Atlas Genet Cytogenet Oncol Haematol. 2007;11(2):113-116.
Free online version   Free pdf version   [Bibliographic record ]
URL : http://AtlasGeneticsOncology.org/Genes/AXIN2ID456ch17q24.html

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