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MUC6 (mucin 6, oligomeric mucus/gel-forming)

Written2009-12Lara Cobler, Marta Garrido, Carme de Bolós
Programa de Recerca en Cancer, IMIM-Hospital del Mar, Dr Aiguader, 88, 08003, Barcelona, Spain

(Note : for Links provided by Atlas : click)

Identity

Alias_namesmucin 6
Other aliasMUC-6
HGNC (Hugo) MUC6
LocusID (NCBI) 4588
Atlas_Id 44115
Location 11p15.5  [Link to chromosome band 11p15]
Location_base_pair Starts at 1012824 and ends at 1036706 bp from pter ( according to hg19-Feb_2009)  [Mapping MUC6.png]
 
  Location of MUC6 gene.
Fusion genes
(updated 2016)
GRINA (8q24.3) / MUC6 (11p15.5)

DNA/RNA

 
  Genomic organization of MUC6 gene (not to scale).
Description MUC6 gene approximately extends 24 kb-long on the chromosome 11 in the region p15.5. The central region has sequences repeated in tandem (TR) with a consensus motif composed of 507 pb. The variable number of TR (VNTR) diverges between 15 and 26, and the difference in distinct alleles is 5 kb long. Some data suggest that short MUC6 alleles are associated with Helicobacter pylori infection and gastric cancer.
Transcription MUC6 gene is composed of 33 exons and the mRNA length is approximately 8 kb. The 5' flanking region of the MUC6 gene contains a TATA box at -35/-29 and potential transcription factor binding sites are described for NFkappaB and Sp family members.
Although MUC6 promoter contains a high percentage of CpG dinucleotides (up to 75%) and a CpG island, its regulation is not influenced by epigenetics.
At present no splice variants forms have been reported.

Protein

 
  Schematic representation of MUC6 peptide structure (not to scale).
Description At the N-terminal region D1, D2, D' and D3 domains similar to von Willebrand factor (vWF) are present.
The TR domain composes the central region. The 169 amino acid consensus sequence has a high content of Thr-Ser-Pro containing numerous potential O-glycosilation sites.
The C-terminal region consists in two distinct regions. One with a high Thr-Ser-Pro content (STP domain), very similar to the tandem repeat region in amino acid composition, and the other region that has a high content of cystein residues. This domain has approximately 25% similarity to the CK domain of the 11p15 human mucins MUC2, MUC5AC, MUC5B and the vWF.
Expression MUC6 was initially isolated from a gastric cDNA library, and it is expressed in the deep gland cells. It is the only mucin produced by acinar cells of the duodenal Brüner's glands and it is also expressed in pancreas, endocervix and gallbladder.
Under pathological conditions, MUC6 expression can be altered, as it is reported below.
Function MUC6, together with MUC5AC, is the major component of the protective layer over the gastric surface where it acts as a selective diffusion barrier for HCl. Furthermore, the O-glycans found in the MUC6 TR have antimicrobial activity against Helicobacter pylori, inhibiting the biosynthesis of cholesteryl-alpha-D-glucopyranoside, a major cell wall component.
Homology Several orthologues of MUC6 have been identified in Mus musculus, Rattus norvegicus, Pan troglodytes and Equus caballus. The chicken and mouse Muc6 have similar domain structures with human MUC6. Furthermore, the tissue-specific expression is conserved in murine organisms.

Implicated in

Note
  
Entity Gastric cancer
Disease Gastric cancer remains the second leading cause related death and the fourth most common cancer in the world, although its incidence is gradually decreasing.
Prognosis MUC6 is highly expressed in gastric mucosa and in intestinal metaplasia MUC6 levels are decreased: In incomplete metaplasia and type I complete metaplasia MUC6 is not detected, whereas it is found in type II and type III complete intestinal metaplasia. MUC6 expression is lower in intestinal-type of gastric carcinomas than in adenomas or normal mucosa suggesting that the downregulation of MUC6 may contribute to the malignant transformation of gastric epithelial cells.
MUC6 is expressed in diffuse-type gastric carcinoma and its levels are greater in early carcinomas than in advanced carcinomas, showing no relationship with the depth of the carcinoma cells invasion.
  
  
Entity Colon cancer
Disease Colorectal cancer is one of the commonest cancers and the third leading cause of cancer death. However, its incidence has decreased due to a most effective intervention and life-style changes in the western countries.
Prognosis Although in normal colon mucosa MUC6 is not expressed, it is upregulated through the adenoma-carcinoma sequence, together with the downregulation of MUC2.
  
  
Entity Salivary glands tumors
Disease Salivary gland tumors are relatively uncommon. However, mucoepidermoid carcinoma is the most frequent malignant tumor of salivary glands.
Prognosis Normal salivary glands do not express MUC6 whereas in mucoepidermoid carcinomas MUC6 is detected preferentially in mucous cells.
  
  
Entity Esophagus adenocarcinoma
Disease The frequency of adenocarcinoma of the esophagus is increasing in the western world from a result of a higher prevalence in Barrett's mucosa.
Prognosis MUC6 showed a decrease in expression with progression from Barrett's esophagus to dysplasia and to adenocarcinoma.
  
  
Entity Pancreatic cancer
Disease Pancreas cancer is a very aggressive tumor with a 5-year survival of less than 5%, and approximately 85% of them correspond to ductal adenocarcinomas.
Prognosis In normal pancreatic epithelium MUC6 is only expressed in ductal and in a minority of centroacinar cells. However, it is upregulated at early stages of pancreatic carcinogenesis and in pancreatic cancer. MUC6 expression has been related to clinicopathological factors and patient prognosis and survival in invasive ductal carcinoma.
  
  
Entity Biliary tract cancer
Disease Biliary tract carcinomas are uncommon tumors that include cholangiocarcinomas and gallbladder carcinomas. These tumors have a poor prognosis: more than 80% of the patients are unresectable with a 6-9 month survival, and this rate is increased to 5 year after surgery.
Prognosis MUC6 is expressed in normal gallbladder mucosa. However in adenomas, dysplasias and carcinomas, MUC6 tends to decrease and its expression is related to non-invasive growth. MUC6 is expressed frequently in pseudopyloric gland metaplasia as well as dysplasia and carcinoma.
  
  
Entity Lung cancer
Disease Lung cancer is the leading cause of cancer-related mortality worldwide. Despite of this its incidence is decreasing.
Prognosis MUC6 is not expressed in normal respiratory epithelium, but it has been focally detected in normal and distal epithelium from cancer patients. MUC6 levels increase significantly in the progression from atypical adenomatous hyperplasia, through bronchio alveolar carcinoma, to adenocarcinoma with mixed subtypes, suggesting that its expression might be associated with the progression of the lung adenocarcinoma. No expression of MUC6 is found in squamous lesions.
  
  
Entity Breast cancer
Disease Breast cancer is the most common cancer worldwide and the leading cause of cancer mortality in women.
Prognosis MUC6 is generally not detected in normal breast epithelium but it is overexpressed in benign breast disease (fibrocystic disease without atypia and atypical fibrocystic disease) and in breast carcinoma.
  
  
Entity Endometrial adenocarcinoma
Disease Endometrial carcinoma is the most common malignant neoplasm of the female genital tract in developed countries, and it occurs predominantly after menopause.
Prognosis MUC6 is not expressed in normal endometrial epithelium. However during endometrial neoplasia transformation, increased levels of MUC6 are detected: from simple hyperplasia, to complex hyperplasia, and in endometrial adenocarcinomas a MUC6 upregulation is found.
  
  
Entity Uterine cervix adenocarcinoma
Disease Adenocarcinoma is the second most common malignancy of the uterine cervix. Its incidence has been increasing and constitutes 10-20% of invasive cervical cancers.
Prognosis Although MUC6 is not expressed in normal cervical epithelium, in adenocarcinomas from uterine cervix; MUC6 can be detected at different levels associated to the histological type.
  

Bibliography

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Mucinous bronchioloalveolar carcinomas display a specific pattern of mucin gene expression among primary lung adenocarcinomas.
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MUC6 apomucin shows a distinct normal tissue distribution that correlates with Lewis antigen expression in the human stomach.
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The mouse secreted gel-forming mucin gene cluster.
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MUC6 gene polymorphism in healthy individuals and in gastric cancer patients from northern Portugal.
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DX2 expression is increased in gastric cancers with less invasiveness and intestinal mucin phenotype.
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Natural antibiotic function of a human gastric mucin against Helicobacter pylori infection.
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PMID 15310903
 
An inventory of mucin genes in the chicken genome shows that the mucin domain of Muc13 is encoded by multiple exons and that ovomucin is part of a locus of related gel-forming mucins.
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PMID 16887038
 
Mucins as differentiation markers in bronchial epithelium. Squamous cell carcinoma and adenocarcinoma display similar expression patterns.
Lopez-Ferrer A, Curull V, Barranco C, Garrido M, Lloreta J, Real FX, de Bolos C.
Am J Respir Cell Mol Biol. 2001 Jan;24(1):22-29.
PMID 11152646
 
Gastrointestinal immunophenotype in adenocarcinomas of the uterine cervix and related glandular lesions: a possible link between lobular endocervical glandular hyperplasia/pyloric gland metaplasia and 'adenoma malignum'.
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PMID 14623937
 
Short mucin 6 alleles are associated with H pylori infection.
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Immunohistochemical study of the expression of MUC6 mucin and co-expression of other secreted mucins (MUC5AC and MUC2) in human gastric carcinomas.
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Citation

This paper should be referenced as such :
Cobler, L ; Garrido, M ; de, Bolòs C
MUC6 (mucin 6, oligomeric mucus/gel-forming)
Atlas Genet Cytogenet Oncol Haematol. 2010;14(10):918-922.
Free journal version : [ pdf ]   [ DOI ]
On line version : http://AtlasGeneticsOncology.org/Genes/MUC6ID44115ch11p15.html


External links

Nomenclature
HGNC (Hugo)MUC6   7517
Cards
AtlasMUC6ID44115ch11p15
Entrez_Gene (NCBI)MUC6  4588  mucin 6, oligomeric mucus/gel-forming
AliasesMUC-6
GeneCards (Weizmann)MUC6
Ensembl hg19 (Hinxton)ENSG00000184956 [Gene_View]
Ensembl hg38 (Hinxton)ENSG00000184956 [Gene_View]  chr11:1012824-1036706 [Contig_View]  MUC6 [Vega]
ICGC DataPortalENSG00000184956
TCGA cBioPortalMUC6
AceView (NCBI)MUC6
Genatlas (Paris)MUC6
WikiGenes4588
SOURCE (Princeton)MUC6
Genetics Home Reference (NIH)MUC6
Genomic and cartography
GoldenPath hg38 (UCSC)MUC6  -     chr11:1012824-1036706 -  11p15.5   [Description]    (hg38-Dec_2013)
GoldenPath hg19 (UCSC)MUC6  -     11p15.5   [Description]    (hg19-Feb_2009)
EnsemblMUC6 - 11p15.5 [CytoView hg19]  MUC6 - 11p15.5 [CytoView hg38]
Mapping of homologs : NCBIMUC6 [Mapview hg19]  MUC6 [Mapview hg38]
OMIM158374   
Gene and transcription
Genbank (Entrez)AK092533 AK096772 AK098503 AY312160 AY458429
RefSeq transcript (Entrez)NM_005961
RefSeq genomic (Entrez)NC_000011 NC_018922 NG_052845 NT_187656 NT_187681 NW_015148966
Consensus coding sequences : CCDS (NCBI)MUC6
Cluster EST : UnigeneHs.528432 [ NCBI ]
CGAP (NCI)Hs.528432
Alternative Splicing GalleryENSG00000184956
Gene ExpressionMUC6 [ NCBI-GEO ]   MUC6 [ EBI - ARRAY_EXPRESS ]   MUC6 [ SEEK ]   MUC6 [ MEM ]
Gene Expression Viewer (FireBrowse)MUC6 [ Firebrowse - Broad ]
SOURCE (Princeton)Expression in : [Datasets]   [Normal Tissue Atlas]  [carcinoma Classsification]  [NCI60]
GenevisibleExpression in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
BioGPS (Tissue expression)4588
GTEX Portal (Tissue expression)MUC6
Protein : pattern, domain, 3D structure
UniProt/SwissProtQ6W4X9   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
NextProtQ6W4X9  [Sequence]  [Exons]  [Medical]  [Publications]
With graphics : InterProQ6W4X9
Splice isoforms : SwissVarQ6W4X9
PhosPhoSitePlusQ6W4X9
Domaine pattern : Prosite (Expaxy)CTCK_2 (PS01225)    VWFD (PS51233)   
Domains : Interpro (EBI)Cys_knot_C    MUC6    TIL_dom    Unchr_dom_Cys-rich    VWF_dom    VWF_type-D   
Domain families : Pfam (Sanger)C8 (PF08742)    TIL (PF01826)    VWD (PF00094)   
Domain families : Pfam (NCBI)pfam08742    pfam01826    pfam00094   
Domain families : Smart (EMBL)C8 (SM00832)  CT (SM00041)  VWC_out (SM00215)  VWD (SM00216)  
Conserved Domain (NCBI)MUC6
DMDM Disease mutations4588
Blocks (Seattle)MUC6
SuperfamilyQ6W4X9
Human Protein AtlasENSG00000184956
Peptide AtlasQ6W4X9
IPIIPI00941125   IPI00980896   IPI00979688   IPI00980291   
Protein Interaction databases
DIP (DOE-UCLA)Q6W4X9
IntAct (EBI)Q6W4X9
FunCoupENSG00000184956
BioGRIDMUC6
STRING (EMBL)MUC6
ZODIACMUC6
Ontologies - Pathways
QuickGOQ6W4X9
Ontology : AmiGOstimulatory C-type lectin receptor signaling pathway  extracellular matrix structural constituent  extracellular region  Golgi lumen  plasma membrane  O-glycan processing  maintenance of gastrointestinal epithelium  
Ontology : EGO-EBIstimulatory C-type lectin receptor signaling pathway  extracellular matrix structural constituent  extracellular region  Golgi lumen  plasma membrane  O-glycan processing  maintenance of gastrointestinal epithelium  
REACTOMEQ6W4X9 [protein]
REACTOME PathwaysR-HSA-977068 [pathway]   
NDEx NetworkMUC6
Atlas of Cancer Signalling NetworkMUC6
Wikipedia pathwaysMUC6
Orthology - Evolution
OrthoDB4588
GeneTree (enSembl)ENSG00000184956
Phylogenetic Trees/Animal Genes : TreeFamMUC6
HOVERGENQ6W4X9
HOGENOMQ6W4X9
Homologs : HomoloGeneMUC6
Homology/Alignments : Family Browser (UCSC)MUC6
Gene fusions - Rearrangements
Fusion : MitelmanGRINA/MUC6 [8q24.3/11p15.5]  [t(8;11)(q24;p15)]  
Fusion: TCGAGRINA 8q24.3 MUC6 11p15.5 PRAD
Polymorphisms : SNP and Copy number variants
NCBI Variation ViewerMUC6 [hg38]
dbSNP Single Nucleotide Polymorphism (NCBI)MUC6
dbVarMUC6
ClinVarMUC6
1000_GenomesMUC6 
Exome Variant ServerMUC6
ExAC (Exome Aggregation Consortium)MUC6 (select the gene name)
Genetic variants : HAPMAP4588
Genomic Variants (DGV)MUC6 [DGVbeta]
DECIPHERMUC6 [patients]   [syndromes]   [variants]   [genes]  
CONAN: Copy Number AnalysisMUC6 
Mutations
ICGC Data PortalMUC6 
TCGA Data PortalMUC6 
Broad Tumor PortalMUC6
OASIS PortalMUC6 [ Somatic mutations - Copy number]
Somatic Mutations in Cancer : COSMICMUC6  [overview]  [genome browser]  [tissue]  [distribution]  
Mutations and Diseases : HGMDMUC6
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 MUC6
DgiDB (Drug Gene Interaction Database)MUC6
DoCM (Curated mutations)MUC6 (select the gene name)
CIViC (Clinical Interpretations of Variants in Cancer)MUC6 (select a term)
intoGenMUC6
NCG5 (London)MUC6
Cancer3DMUC6(select the gene name)
Impact of mutations[PolyPhen2] [SIFT Human Coding SNP] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
Diseases
OMIM158374   
Orphanet
MedgenMUC6
Genetic Testing Registry MUC6
NextProtQ6W4X9 [Medical]
TSGene4588
GENETestsMUC6
Target ValidationMUC6
Huge Navigator MUC6 [HugePedia]
snp3D : Map Gene to Disease4588
BioCentury BCIQMUC6
ClinGenMUC6
Clinical trials, drugs, therapy
Chemical/Protein Interactions : CTD4588
Chemical/Pharm GKB GenePA31322
Clinical trialMUC6
Miscellaneous
canSAR (ICR)MUC6 (select the gene name)
Probes
Litterature
PubMed55 Pubmed reference(s) in Entrez
GeneRIFsGene References Into Functions (Entrez)
CoreMineMUC6
EVEXMUC6
GoPubMedMUC6
iHOPMUC6
REVIEW articlesautomatic search in PubMed
Last year publicationsautomatic search in PubMed

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