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CST4 Gene

protein-coding   GIFtS: 56
GCID: GC20M023666

Cystatin S

  See CST4-related disease
at  

(According to 1HGNC, 2Entrez Gene,
3UniProtKB/Swiss-Prot, 4UniProtKB/TrEMBL, 5OMIM, 6GeneLoc, 7Ensembl, 8DME, 9miRBase, 10fRNAdb, 12H-InvDB, 13NCBI, 14NONCODE, and/or 15RNAdb)
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Aliases
Cystatin S1 2     cystatin-S2
Salivary Acidic Protein 12 3     cystatin-SA-III2
Cystatin 42     Cystatin-43
cystatin-42     Cystatin-SA-III3

External Ids:    HGNC: 24761   Entrez Gene: 14722   Ensembl: ENSG000001014417   OMIM: 1238575   UniProtKB: P010363   

Export aliases for CST4 gene to outside databases

Previous GC identifers: GC20M023654 GC20M023661 GC20M023614 GC20M023626


(According to Entrez Gene, GeneCards, Tocris Bioscience, Wikipedia's Gene Wiki, PharmGKB,
UniProtKB/Swiss-Prot, and/or UniProtKB/TrEMBL)
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Entrez Gene summary for CST4 Gene:
The cystatin superfamily encompasses proteins that contain multiple cystatin-like sequences. Some of the members
are active cysteine protease inhibitors, while others have lost or perhaps never acquired this inhibitory
activity. There are three inhibitory families in the superfamily, including the type 1 cystatins (stefins), type
2 cystatins and the kininogens. The type 2 cystatin proteins are a class of cysteine proteinase inhibitors found
in a variety of human fluids and secretions. The cystatin locus on chromosome 20 contains the majority of the
type 2 cystatin genes and pseudogenes. This gene is located in the cystatin locus and encodes a type 2 salivary
cysteine peptidase inhibitor. The protein is an S-type cystatin, based on its high level of expression in saliva,
tears and seminal plasma. The specific role in these fluids is unclear but antibacterial and antiviral activity
is present, consistent with a protective function. (provided by RefSeq, Jul 2008)

GeneCards Summary for CST4 Gene:
CST4 (cystatin S) is a protein-coding gene. Diseases associated with CST4 include blepharitis. GO annotations related to this gene include cysteine-type endopeptidase inhibitor activity. An important paralog of this gene is CST2.

UniProtKB/Swiss-Prot: CYTS_HUMAN, P01036
Function: This protein strongly inhibits papain and ficin, partially inhibits stem bromelain and bovine cathepsin
C, but does not inhibit porcine cathepsin B or clostripain. Papain is inhibited non-competitively

Gene Wiki entry for CST4 Gene


(According to GeneLoc and/or HGNC, and/or
Entrez Gene (NCBI build 37),
and/or miRBase,
Genomic Views according to UCSC (hg19) and Ensembl (release 75), Regulatory elements and Epigenetics data according to QIAGEN, and/or SwitchGear Genomics)
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RefSeq DNA sequence at NCBI GenBank:
NC_000020.10  NC_018931.2  NT_011387.9  
Regulatory elements:
   Regulatory transcription factor binding sites in the CST4 gene promoter:
         ER-alpha   SREBP-1c   Nkx5-1   SREBP-1b   POU2F1   SREBP-1a   POU2F1a   Zic1   c-Myb   NF-kappaB1   
         Other transcription factors

SwitchGear Promoter luciferase reporter plasmidCST4 promoter sequence
   Search Chromatin IP Primers for CST4

Epigenetics:
DNA Methylation CpG Assay Predesigned for Pyrosequencing in human, mouse, rat CST4


Genomic Location:
Genomic View: UCSC Golden Path with GeneCards custom track

Entrez Gene cytogenetic band: 20p11.21   Ensembl cytogenetic band:  20p11.21   HGNC cytogenetic band: 20p11.2

CST4 Gene in genomic location: bands according to Ensembl, locations according to (and/or Entrez Gene and/or Ensembl if different)
CST4 gene location

GeneLoc information about chromosome 20         GeneLoc Exon Structure

GeneLoc location for GC20M023666:  view genomic region     (about GC identifiers)

Start:
23,666,277 bp from pter      End:
23,669,677 bp from pter
Size:
3,401 bases      Orientation:
minus strand

(According to 1UniProtKB, HORDE, 2neXtProt, Ensembl, and/or Reactome, Modification sites according to PhosphoSitePlus, Specific Peptides from DME, RefSeq according to NCBI, PDB rendering according to OCA and/or Proteopedia, Recombinant Proteins from EMD Millipore, R&D Systems, GenScript, Enzo Life Sciences, OriGene, Novus Biologicals, Sino Biological, ProSpec, Cloud-Clone Corp., and/or eBioscience,
Biochemical Assays by EMD Millipore, R&D Systems, OriGene, GenScript, Cell Signaling Technology, Enzo Life Sciences, Cloud-Clone Corp., and/or eBioscience, Antibodies by EMD Millipore, R&D Systems, Cell Signaling Technology, OriGene, Novus Biologicals, Thermo Fisher Scientific, LSBio, Abcam, Cloud-Clone Corp, and/or others.)
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UniProtKB/Swiss-Prot: CYTS_HUMAN, P01036 (See protein sequence)
Recommended Name: Cystatin-S precursor  
Size: 141 amino acids; 16214 Da
Mass spectrometry: Mass=14175.8569; Mass_error=0.0564; Method=Electrospray; Range=21-141; Source=PubMed:20189825;
Mass spectrometry: Mass=14255.8567; Mass_error=0.0899; Method=Electrospray; Range=21-141; Note=Monophosphorylated
at Ser-23, also called form S1; Source=PubMed:20189825;
Mass spectrometry: Mass=14335.811; Mass_error=0.0775; Method=Electrospray; Range=21-141; Note=Diphosphorylated at
Ser-21 and Ser-23, also called form S2; Source=PubMed:20189825;
Secondary accessions: Q9UBI5 Q9UCS9

Explore the universe of human proteins at neXtProt for CST4: NX_P01036

Explore proteomics data for CST4 at MOPED

Post-translational modifications: 

  • Phosphorylated at both its N- and C-terminal regions1
  • Modification sites at neXtProt
  • Modification sites at PhosphoSitePlus

  • See CST4 Protein Expression from SPIRE MOPED, PaxDB, and MaxQB

    REFSEQ proteins: NP_001890.1  
    ENSEMBL proteins: 
     ENSP00000217423  

    CST4 Human Recombinant Protein Products:

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    (According to HGNC, IUPHAR, InterPro, ProtoNet, UniProtKB, and/or BLOCKS, Sets of similar genes according to GeneDecks)
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    3 InterPro protein domains:
     IPR018073 Prot_inh_cystat_CS
     IPR000010 Prot_inh_cystat
     IPR027214 Cystatin

    Graphical View of Domain Structure for InterPro Entry P01036

    ProtoNet protein and cluster: P01036

    2 Blocks protein domains:
    IPB000010 Cysteine protease inhibitor
    IPB003243 Proteinase inhibitor I25A and I25B


    UniProtKB/Swiss-Prot: CYTS_HUMAN, P01036
    Similarity: Belongs to the cystatin family


    Find genes that share domains with CST4           About GenesLikeMe


    (According to 1UniProtKB, Genatlas, LifeMap Discovery™, IUBMB, and/or 2DME, Human phenotypes from GenomeRNAi, Animal models from MGI Mar 06 2013, inGenious Targeting Laboratory, genOway,
    transcription factor targeting from QIAGEN and/or HOMER, miRNA Gene Targets from miRTarBase, shRNA from OriGene, siRNAs from OriGene, QIAGEN, microRNA from QIAGEN, SwitchGear Genomics, Gene Editing from DNA2.0, Clones from OriGene, GenScript, Sino Biological, DNA2.0, Vector BioLabs and/or Addgene, Cell Lines from GenScript, ESI BIO, In Situ Hybridization Assays from Advanced Cell Diagnostics, Flow cytometry from eBioscience, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene.)
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    Molecular Function:

         UniProtKB/Swiss-Prot Summary: CYTS_HUMAN, P01036
    Function: This protein strongly inhibits papain and ficin, partially inhibits stem bromelain and bovine cathepsin
    C, but does not inhibit porcine cathepsin B or clostripain. Papain is inhibited non-competitively

         Genatlas biochemistry entry for CST4:
    cystatin S,cysteine proteinase inhibitor,expressed at high levels in saliva,tears,seminal plasma,type 2 family

         Gene Ontology (GO): 1 molecular function term:    About this table

    GO IDQualified GO termEvidencePubMed IDs
    GO:0004869cysteine-type endopeptidase inhibitor activity IEA--
         
    Find genes that share ontologies with CST4           About GenesLikeMe


    Phenotypes:
         1 GenomeRNAi human phenotype for CST4:
     Synthetic lethal with imatinib 

    Animal Models:
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    hsa-mir-335-5p (MIRT017577)

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    (According to UniProtKB, COMPARTMENTS Subcellular localization database, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene.)
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    Subcellular locations from UniProtKB/Swiss-Prot
    CYTS_HUMAN, P01036: Secreted
    Subcellular locations from COMPARTMENTS: 

    CompartmentConfidence
    extracellular5
    lysosome1
    vacuole1

    Gene Ontology (GO): 3 cellular component terms:    About this table

    GO IDQualified GO termEvidencePubMed IDs
    GO:0005576extracellular region ----
    GO:0005615extracellular space IDA--
    GO:0070062extracellular vesicular exosome IDA19199708

    Find genes that share ontologies with CST4           About GenesLikeMe


    (SuperPaths according to PathCards, Pathways according to R&D Systems, Cell Signaling Technology, KEGG, PharmGKB, BioSystems, Sino Biological, Reactome, Tocris Bioscience, GeneGo (Thomson Reuters), QIAGEN, and/or UniProtKB, Sets of similar genes according to GeneDecks, Interaction Networks according to QIAGEN, and/or STRING, Interactions according to 1UniProtKB, 2MINT, 3I2D, and/or 4STRING, with links to IntAct and Ensembl, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene).
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    SuperPaths for CST4 About    
    See pathways by source

    SuperPathContained pathways About
    1Salivary secretion
    Salivary secretion0.34


    Find genes that share SuperPaths with CST4           About GenesLikeMe

    Pathways by source                                                                                                                                                                 See SuperPaths
    Show all pathways




    1 Kegg Pathway  (Kegg details for CST4):
        Salivary secretion

        Custom Pathway & Disease-focused RT2 Profiler PCR Arrays for CST4
    Interactions:

        GeneGlobe Interaction Network for CST4

    STRING Interaction Network Preview (showing 2 interactants - click image to see more details)

    2 Interacting proteins for CST4 (P010363 ENSP000002174234) via UniProtKB, MINT, STRING, and/or I2D
    InteractantInteraction Details
    GeneCardExternal ID(s)
    SOD2P041793, ENSP000003371274I2D: score=1 STRING: ENSP00000337127
    KNCNENSP000003796074STRING: ENSP00000379607
    About this table

    Gene Ontology (GO): 2 biological process terms:    About this table

    GO IDQualified GO termEvidencePubMed IDs
    GO:0001580detection of chemical stimulus involved in sensory perception of bitter taste IDA--
    GO:0001895retina homeostasis IEP--

    Find genes that share ontologies with CST4           About GenesLikeMe



    (Chemical Compounds according to UniProtKB, Enzo Life Sciences, EMD Millipore, Tocris Bioscience, ApexBio, HMDB, BitterDB, and/or Novoseek, Ligands according to IUPHAR, and Drugs according to DrugBank, Enzo Life Sciences, and/or PharmGKB)
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    Browse Small Molecules at EMD Millipore
       Browse drugs & compounds from Enzo Life Sciences
      Browse compounds at ApexBio 

    Browse Tocris compounds for CST4 (CYTS)



    (Secondary structures according to fRNAdb,
    GenBank/EMBL/DDBJ Accessions according to
    Unigene (Build 236 Homo sapiens; Apr 25 2013) or GenBank,
    RefSeq according to Entrez Gene,
    DOTS (version 10), and/or AceView, transcript ids from Ensembl with links to UCSC,
    exon structure from GeneLoc, alternative splicing isoforms according to ASD and/or ECgene,
    siRNAs from OriGene, QIAGEN, shRNA from OriGene, microRNA from QIAGEN, SwitchGear Genomics,
    Tagged/untagged cDNA clones from OriGene, GenScript, DNA2.0, Vector BioLabs, and/or Addgene, Primers from OriGene, and/or QIAGEN, Flow cytometry from eBioscience )
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    REFSEQ mRNAs for CST4 gene: 
    NM_001899.2  

    Unigene Cluster for CST4:

    Cystatin S
    Hs.654549  [show with all ESTs]
    Unigene Representative Sequence: CB959353
    1 Ensembl transcript including schematic representation, and UCSC links where relevant:
    ENST00000217423(uc002wto.1)
    miRNA
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    Additional mRNA sequence: 

    BC065714.1 BC074952.2 BC074953.2 X54667.1 

    4 DOTS entries:

    DT.103103  DT.120823043  DT.97759874  DT.91721790 

    Selected AceView cDNA sequences (see all 198):

    NM_001898 BC074953 BC074952 BX485701 CD721927 NM_001899 AI648499 CD721796 
    CK429248 CB959252 BP334429 BP333995 BV203402 BP334029 BV203396 BV203393 
    BP334885 BP334805 CK429873 BP334108 BP334742 BP333755 CB986215 BP334945 

    GeneLoc Exon Structure


    (RNA expression data according to H-InvDB, NONCODE, miRBase, and RNAdb, Expression images according to data from BioGPS, Illumina Human BodyMap, and CGAP SAGE, Sets of similar genes according to GeneDecks, in vivo and in vitro expression data from LifeMap Discovery™, Protein expression images according to data from SPIRE 1MOPED, 2PaxDb, and 3MaxQB, plus additional links to SOURCE, and/or BioGPS, and/or UniProtKB,
    PCR Arrays from QIAGEN, Primers from OriGene, and/or QIAGEN, In Situ Hybridization Assays from Advanced Cell Diagnostics)
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    CST4 expression in normal human tissues (normalized intensities)
    See probesets specificity/sensitivity at GeneAnnot
    About this imageBioGPS <intensity>2/3
    CGAP TAG: --
    CST4 Expression
    About this image

    CST4 Protein expression data from MOPED1, PaxDb2 and MaxQB3    About this image

    CST4 Protein Expression

    SOURCE GeneReport for Unigene cluster: Hs.654549

    UniProtKB/Swiss-Prot: CYTS_HUMAN, P01036
    Tissue specificity: Expressed in submandibular and sublingual saliva but not in parotid saliva (at protein level).
    Expressed in saliva, tears, urine and seminal fluid

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    In Situ
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    (Orthologs according to 1,2HomoloGene (2older version, for species not in 1newer version), 3euGenes, 4SGD , 5MGI Mar 06 2013, with possible further links to Flybase and/or WormBase, and/or 6Ensembl pan taxonomic compara , Gene Trees according to Ensembl and TreeFam)
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    This gene was present in the common ancestor of chordates.

    Orthologs for CST4 gene from Selected species (see all 9)    About this table
    Organism Taxonomic
    classification
    Gene Description Human
    Similarity
    Orthology
    Type
    Details
    chicken
    (Gallus gallus)
    Aves gga-mir-35266
    Gallus gallus cystatin C (CST3), mRNA.
    36(a)
    1 → many
    3(14928612-14932913)
    lizard
    (Anolis carolinensis)
    Reptilia --
    Uncharacterized protein
    39(a)
    1 → many
    4(82204275-82217421)
    zebrafish
    (Danio rerio)
    Actinopterygii cst36
    cystatin C (amyloid angiopathy and cerebral hemorr...
    30(a)
    many ↔ many
    13(33900031-33908388) ENSDARG00000007795
            Species with no ortholog for CST4

    ENSEMBL Gene Tree for CST4 (if available)
    TreeFam Gene Tree for CST4 (if available)

    (Paralogs according to 1HomoloGene,
    2Ensembl, and 3SIMAP, Pseudogenes according to 4Pseudogene.org Build 68)
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    Paralogs for CST4 gene
    CST22  CSTL12  CST82  CST12  CST32  CST52  CST112  
    9 SIMAP similar genes for CST4 using alignment to 1 protein entry:     CYTS_HUMAN:
    CST1    CST2    CST5    CST3    CST9LP1    CST11
    CST8    CSTL1    CST6

    Find genes that share paralogs with CST4           About GenesLikeMe


    1 Pseudogenes.org Pseudogene for CST4
    PGOHUM00000247469


    (SNPs/Variants according to the 1NCBI SNP Database, 2Ensembl, 3PupaSUITE, 4UniProtKB, and DNA2.0, Linkage Disequilibrium by HapMap, Structural Variations(CNVs/InDels/Inversions) from the Database of Genomic Variants, Mutations from the Human Gene Mutation Database (HGMD), the Human Cytochrome P450 Allele Nomenclature Database, and the Locus Specific Mutation Databases (LSDB), Blood group antigen gene mutations by BGMUT, Resequencing Primers, Cancer Mutation PCR Arrays and Assays, and Copy Number PCR Arrays from QIAGEN)
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    Selected SNPs for CST4 (see all 243)    About this table                                 

    Genomic DataTranscription Related DataAllele Frequencies
    SNP IDValidClinical
    significance
    Chr 20 posSequence#AA
    Chg
    TypeMore#Allele
    freq
    PopTotal
    sample
    More
    ----------
    VAR_0365494
    A breast cancer sample4--see VAR_0365492 T N mis40--------
    rs1898497191,2
    --23665793(+) CTGTGA/GAAAGA 1 -- ds50010--------
    rs1927933071,2
    --23665802(+) GAACTA/TCACGT 1 -- ds50010--------
    rs1133439841,2
    F--23665831(+) TGTCTA/TGGGGA 1 -- ds50012Minor allele frequency- T:0.02CSA WA 120
    rs1120454511,2
    F--23665839(+) GGATCT/AGATGA 1 -- ds50012Minor allele frequency- A:0.02CSA WA 120
    rs1414314841,2
    --23665898(+) ATCTAC/TGTCCT 1 -- ds50010--------
    rs1841158811,2
    --23665899(+) TCTACA/GTCCTG 1 -- ds50010--------
    rs1474794381,2
    --23665900(+) CTACGG/TCCTGG 1 -- ds50010--------
    rs1884071301,2
    --23665941(+) TGGACA/TCAACC 1 -- ds50010--------
    rs1807715161,2
    --23665946(+) ACAACA/CAGCCC 1 -- ds50010--------

    HapMap Linkage Disequilibrium report for CST4 (23666277 - 23669677 bp)

    Structural Variations
         Database of Genomic Variants (DGV) 5 variations for CST4:    About this table    
    Variant IDTypeSubtypePubMed ID
    esv2722303CNV Deletion23290073
    nsv458950CNV Loss19166990
    nsv482066CNV Gain20164927
    nsv458952CNV Gain19166990
    nsv912821CNV Gain+Loss21882294

    Site Specific Mutation Identification with PCR Assays
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    (in which this Gene is Involved, According to MalaCards, OMIM, UniProtKB, the University of Copenhagen DISEASES database, Genatlas, GeneTests, GAD, HuGE Navigator, and/or TGDB.)
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    OMIM gene information: 123857    OMIM disorders: --

    1 disease for CST4:    
    About MalaCards
    blepharitis

    1 disease from the University of Copenhagen DISEASES database for CST4:
    Periodontal disease

    Find genes that share disorders with CST4           About GenesLikeMe


    Export disorders for CST4 gene to outside databases

    (in PubMed. Associations of this gene to articles via 1Entrez Gene, 2UniProtKB/Swiss-Prot, 3HGNC, 4GAD, 5PharmGKB, 6HMDB, 7DrugBank, 8UniProtKB/TrEMBL, 9 Novoseek, and/or 10fRNAdb)
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    PubMed articles for CST4 gene, integrated from 10 sources (see all 31):
    (articles sorted by number of sources associating them with CST4)
        Utopia: connect your pdf to the dynamic
    world of online information

    1. Salivary cystatin SA-III, a potential precursor of the acquired enamel pellicle, is phosphorylated at both its amino- and carboxyl- terminal regions. (PubMed id 1898055)1, 2, 9 Lamkin M.S.... Oppenheim F.G. (Arch. Biochem. Biophys. 1991)
    2. The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC). (PubMed id 15489334)1, 2 Gerhard D.S.... Malek J. (Genome Res. 2004)
    3. The DNA sequence and comparative analysis of human chromosome 20. (PubMed id 11780052)1, 2 Deloukas P....Rogers J. (Nature 2001)
    4. Characterization of two members (CST4 and CST5) of the cystatin gene family and molecular evolution of cystatin genes. (PubMed id 1334620)1, 2 Saitoh E.... Ohnishi K. (Agents Actions 1992)
    5. Large-scale purification and characterization of the major phosphoproteins and mucins of human submandibular-sublingual saliva. (PubMed id 1747107)1, 2 Ramasubbu N....Levine M.J. (Biochem. J. 1991)
    6. Human salivary cystatin S. Cloning, sequence analysis, hybridization in situ and immunocytochemistry. (PubMed id 1898352)1, 2 Bobek L.A.... Levine M.J. (Biochem. J. 1991)
    7. The human cystatin gene family: cloning of three members and evolutionary relationship between cystatins and Bowman-Birk type proteinase inhibitors. (PubMed id 1801729)1, 3 Saitoh E....Ohnishi K. (Biomed. Biochim. Acta 1991)
    8. The effects of human salivary cystatins and statherin on hydroxyapatite crystallization. (PubMed id 1741693)1, 2 Johnsson M....Nancollas G.H. (Arch. Oral Biol. 1991)
    9. Identification of full-sized forms of salivary (S-type) cystatins (cystatin SN, cystatin SA, cystatin S, and two phosphorylated forms of cystatin S) in human whole saliva and determination of phosphorylation sites of cystatin S. (PubMed id 1778989)1, 2 Isemura S.... Minakata K. (J. Biochem. 1991)
    10. Identification of a long form of cystatin from human saliva by rapid microbore HPLC mapping. (PubMed id 3496880)1, 2 Hawke D.H.... Hunkapiller M.W. (Biochem. Biophys. Res. Commun. 1987)

    (in PubMed, OMIM, and NCBI Bookshelf)
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      Query String
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    (According to Entrez Gene, HGNC, AceView, euGenes, Ensembl, miRBase, ECgene, Kegg, and/or H-InvDB)
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    Entrez Gene: 1472 HGNC: 2476 AceView: CST1andCST4 Ensembl:ENSG00000101441 euGenes: HUgn1472
    ECgene: CST4 Kegg: 1472 H-InvDB: CST4

    (According to HUGE)
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      --

    (According to PharmGKB, ATLAS, HORDE, IMGT, LEIDEN, UniProtKB/Swiss-Prot, UniProtKB/TrEMBL, and/or others, e.g. Wikipedia and GeneReviews, via UniProtKB/Swiss-Prot)
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    NameDescription
    PharmGKB entry for CST4 Pharmacogenomics, SNPs, Pathways

    (Patent information from GeneIP,
    Licensable technologies from WIS Yeda, Salk, Tufts,
    IP news from LifeMap Sciences, Inc.)
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    Patent Information for CST4 gene:
    Search GeneIP for patents involving CST4

    GeneCards and IP:
    Japan Patent Office Licenses GeneCards     European Patent Office Licenses GeneCards     Improving the IP Search



    (Antibodies, recombinant proteins, and assays from EMD Millipore, R&D Systems, OriGene, QIAGEN, GenScript, Cell Signaling Technology, Novus Biologicals, Sino Biological, Enzo Life Sciences, Abcam, ProSpec, Cloud-Clone Corp., Thermo Fisher Scientific, LSBio, eBioscience, and/or others, Gene Editing from DNA2.0. Clones from OriGene, GenScript, Sino Biological, DNA2.0, SwitchGear Genomics, Vector BioLabs, Addgene, Cell lines from GenScript, and ESI BIO, Flow cytometery from eBioscience, PCR Arrays from QIAGEN, Drugs and/or compounds from EMD Millipore, Tocris Bioscience, Enzo Life Sciences, and/or ApexBio, In Situ Hybridization Assays from
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