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Aliases for CST4 Gene

Aliases for CST4 Gene

  • Cystatin S 2 3 5
  • Salivary Acidic Protein 1 3 4
  • Cystatin-SA-III 3 4
  • Cystatin 4 3
  • Cystatin-S 3
  • Cystatin-4 4

External Ids for CST4 Gene

Previous GeneCards Identifiers for CST4 Gene

  • GC20M023654
  • GC20M023661
  • GC20M023614
  • GC20M023666
  • GC20M023626

Summaries for CST4 Gene

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. Among its related pathways are Salivary secretion. Gene Ontology (GO) annotations related to this gene include cysteine-type endopeptidase inhibitor activity. An important paralog of this gene is CST2.

UniProtKB/Swiss-Prot for CST4 Gene

  • 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

Additional gene information for CST4 Gene

No data available for CIViC summary , Tocris Summary , PharmGKB "VIP" Summary , fRNAdb sequence ontologies and piRNA Summary for CST4 Gene

Genomics for CST4 Gene

GeneHancer (GH) Regulatory Elements for CST4 Gene

Promoters and enhancers for CST4 Gene
GeneHancer (GH) Identifier GH Type GH
GH Sources Gene Association Score Total Score TSS distance (kb) Number of Genes Away Size (kb) Transcription Factor
Binding Sites
Gene Targets
GH20J023689 Promoter 0.5 EPDnew 650.7 0.0 -18 0.1 CST4 CST2P1
GH20J023692 Enhancer 0.7 Ensembl ENCODE 0.7 -4.2 -4210 1.3 FOSL1 JUN MNT FOS CST9 CST1 CST3 CST4 CST2P1
GH20J023704 Enhancer 0.2 FANTOM5 0.4 -15.4 -15359 0 CST9 CST2P1 CST4
- Elite GeneHancer and/or Elite GeneHancer-gene association Download GeneHancer data from 2017 publication | Request up-to-date GeneHancer data (full dataset)

GeneHancers around CST4 on UCSC Golden Path with GeneCards custom track

Top Transcription factor binding sites by QIAGEN in the CST4 gene promoter:
  • c-Myb
  • POU2F1a
  • POU2F1
  • Zic1
  • SREBP-1c
  • SREBP-1b
  • SREBP-1a
  • Nkx5-1
  • ER-alpha
  • NF-kappaB1

Genomic Locations for CST4 Gene

Genomic Locations for CST4 Gene
3,401 bases
Minus strand
3,401 bases
Minus strand

Genomic View for CST4 Gene

Genes around CST4 on UCSC Golden Path with GeneCards custom track

Cytogenetic band:
CST4 Gene in genomic location: bands according to Ensembl, locations according to GeneLoc (and/or Entrez Gene and/or Ensembl if different)
Genomic Location for CST4 Gene
GeneLoc Logo Genomic Neighborhood Exon StructureGene Density

RefSeq DNA sequence for CST4 Gene

Proteins for CST4 Gene

  • Protein details for CST4 Gene (UniProtKB/Swiss-Prot)

    Protein Symbol:
    Recommended name:
    Protein Accession:
    Secondary Accessions:
    • Q9UBI5
    • Q9UCS9

    Protein attributes for CST4 Gene

    141 amino acids
    Molecular mass:
    16214 Da
    Quaternary structure:
    No Data Available

neXtProt entry for CST4 Gene

Post-translational modifications for CST4 Gene

Other Protein References for CST4 Gene

ENSEMBL proteins:
REFSEQ proteins:

No data available for DME Specific Peptides for CST4 Gene

Domains & Families for CST4 Gene

Gene Families for CST4 Gene

Human Protein Atlas (HPA):
  • Predicted secreted proteins

Suggested Antigen Peptide Sequences for CST4 Gene

GenScript: Design optimal peptide antigens:

Graphical View of Domain Structure for InterPro Entry



  • Belongs to the cystatin family.
  • Belongs to the cystatin family.
genes like me logo Genes that share domains with CST4: view

Function for CST4 Gene

Molecular function for CST4 Gene

UniProtKB/Swiss-Prot 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:
cystatin S,cysteine proteinase inhibitor,expressed at high levels in saliva,tears,seminal plasma,type 2 family

Phenotypes From GWAS Catalog for CST4 Gene

Gene Ontology (GO) - Molecular Function for CST4 Gene

GO ID Qualified GO term Evidence PubMed IDs
GO:0002020 protease binding IBA --
GO:0004869 cysteine-type endopeptidase inhibitor activity IDA 3488317
GO:0030414 peptidase inhibitor activity IEA --
genes like me logo Genes that share ontologies with CST4: view
genes like me logo Genes that share phenotypes with CST4: view

Animal Model Products

CRISPR Products

miRNA for CST4 Gene

miRTarBase miRNAs that target CST4

Inhibitory RNA Products

  • Search GeneCopoeia for shRNA, lentivirus and/or AAV clone products for CST4

No data available for Enzyme Numbers (IUBMB) , Human Phenotype Ontology , Animal Models , Transcription Factor Targets and HOMER Transcription for CST4 Gene

Localization for CST4 Gene

Subcellular locations from UniProtKB/Swiss-Prot for CST4 Gene


Subcellular locations from

Extracellular space Cytosol Plasma membrane Cytoskeleton Lysosome Endosome Peroxisome ER Golgi Apparatus Nucleus Mitochondrion 0 1 2 3 4 5 Confidence
COMPARTMENTS Subcellular localization image for CST4 gene
Compartment Confidence
extracellular 5
lysosome 2

Gene Ontology (GO) - Cellular Components for CST4 Gene

GO ID Qualified GO term Evidence PubMed IDs
GO:0005576 extracellular region IEA --
GO:0005615 extracellular space IDA 3488317
GO:0070062 extracellular exosome HDA 19199708
genes like me logo Genes that share ontologies with CST4: view

No data available for Subcellular locations from the Human Protein Atlas (HPA) for CST4 Gene

Pathways & Interactions for CST4 Gene

SuperPathway Contained pathways
1 Salivary secretion
genes like me logo Genes that share pathways with CST4: view

Pathways by source for CST4 Gene

1 KEGG pathway for CST4 Gene

Gene Ontology (GO) - Biological Process for CST4 Gene

GO ID Qualified GO term Evidence PubMed IDs
GO:0001580 detection of chemical stimulus involved in sensory perception of bitter taste IDA 24248522
GO:0001895 retina homeostasis HEP 23580065
GO:0010466 negative regulation of peptidase activity IEA --
GO:0045861 negative regulation of proteolysis IDA 3488317
GO:2000117 negative regulation of cysteine-type endopeptidase activity IBA --
genes like me logo Genes that share ontologies with CST4: view

No data available for SIGNOR curated interactions for CST4 Gene

Drugs & Compounds for CST4 Gene

No Compound Related Data Available

Transcripts for CST4 Gene

mRNA/cDNA for CST4 Gene

(1) REFSEQ mRNAs :
(4) Additional mRNA sequences :
(198) Selected AceView cDNA sequences:
(1) Ensembl transcripts including schematic representations, and UCSC links where relevant :

Unigene Clusters for CST4 Gene

Cystatin S:
Representative Sequences:

CRISPR Products

Inhibitory RNA Products

  • Search GeneCopoeia for shRNA, lentivirus and/or AAV clone products for CST4

Alternative Splicing Database (ASD) splice patterns (SP) for CST4 Gene

No ASD Table

Relevant External Links for CST4 Gene

GeneLoc Exon Structure for
ECgene alternative splicing isoforms for

Expression for CST4 Gene

mRNA expression in normal human tissues from GTEx, Illumina, BioGPS, and CGAP SAGE for CST4 Gene

mRNA differential expression in normal tissues according to GTEx for CST4 Gene

This gene is overexpressed in Thyroid (x21.3), Prostate (x19.3), and Uterus (x5.8).

Protein differential expression in normal tissues from HIPED for CST4 Gene

This gene is overexpressed in Skin (32.6), Salivary gland (24.9), and Saliva (10.9).

Integrated Proteomics: protein expression in normal tissues and cell lines from ProteomicsDB and MOPED for CST4 Gene

Protein tissue co-expression partners for CST4 Gene

NURSA nuclear receptor signaling pathways regulating expression of CST4 Gene:


SOURCE GeneReport for Unigene cluster for CST4 Gene:


mRNA Expression by UniProt/SwissProt for CST4 Gene:

Tissue specificity: Expressed in submandibular and sublingual saliva but not in parotid saliva (at protein level). Expressed in saliva, tears, urine and seminal fluid.

Evidence on tissue expression from TISSUES for CST4 Gene

  • Saliva(4.5)
  • Thyroid gland(4)
  • Gall bladder(2.1)
genes like me logo Genes that share expression patterns with CST4: view

No data available for mRNA expression in embryonic tissues and stem cells from LifeMap Discovery and Phenotype-based relationships between genes and organs from Gene ORGANizer for CST4 Gene

Orthologs for CST4 Gene

This gene was present in the common ancestor of chordates.

Orthologs for CST4 Gene

Organism Taxonomy Gene Similarity Type Details
(Pan troglodytes)
Mammalia CST4 34
  • 94 (a)
(Canis familiaris)
Mammalia -- 34
  • 52 (a)
(Bos Taurus)
Mammalia CST3 34
  • 47 (a)
(Monodelphis domestica)
Mammalia -- 34
  • 39 (a)
(Mus musculus)
Mammalia Cst10 34
  • 36 (a)
(Ornithorhynchus anatinus)
Mammalia -- 34
  • 34 (a)
(Gallus gallus)
Aves gga-mir-3526 34
  • 36 (a)
(Anolis carolinensis)
Reptilia -- 34
  • 39 (a)
(Danio rerio)
Actinopterygii cst3 34
  • 30 (a)
Species where no ortholog for CST4 was found in the sources mined by GeneCards:
  • A. gosspyii yeast (Ashbya gossypii)
  • Actinobacteria (Mycobacterium tuberculosis)
  • African clawed frog (Xenopus laevis)
  • African malaria mosquito (Anopheles gambiae)
  • Alicante grape (Vitis vinifera)
  • alpha proteobacteria (Wolbachia pipientis)
  • amoeba (Dictyostelium discoideum)
  • Archea (Pyrococcus horikoshii)
  • baker's yeast (Saccharomyces cerevisiae)
  • barley (Hordeum vulgare)
  • beta proteobacteria (Neisseria meningitidis)
  • bread mold (Neurospora crassa)
  • Chromalveolata (Phytophthora infestans)
  • common water flea (Daphnia pulex)
  • corn (Zea mays)
  • E. coli (Escherichia coli)
  • filamentous fungi (Aspergillus nidulans)
  • Firmicute bacteria (Streptococcus pneumoniae)
  • fission yeast (Schizosaccharomyces pombe)
  • fruit fly (Drosophila melanogaster)
  • green algae (Chlamydomonas reinhardtii)
  • honey bee (Apis mellifera)
  • K. lactis yeast (Kluyveromyces lactis)
  • loblloly pine (Pinus taeda)
  • malaria parasite (Plasmodium falciparum)
  • medicago trunc (Medicago Truncatula)
  • moss (Physcomitrella patens)
  • orangutan (Pongo pygmaeus)
  • pig (Sus scrofa)
  • rainbow trout (Oncorhynchus mykiss)
  • rat (Rattus norvegicus)
  • rice (Oryza sativa)
  • rice blast fungus (Magnaporthe grisea)
  • schistosome parasite (Schistosoma mansoni)
  • sea anemone (Nematostella vectensis)
  • sea squirt (Ciona intestinalis)
  • sea squirt (Ciona savignyi)
  • sea urchin (Strongylocentrotus purpuratus)
  • sorghum (Sorghum bicolor)
  • soybean (Glycine max)
  • stem rust fungus (Puccinia graminis)
  • sugarcane (Saccharum officinarum)
  • thale cress (Arabidopsis thaliana)
  • tomato (Lycopersicon esculentum)
  • toxoplasmosis (Toxoplasma gondii)
  • Trichoplax (Trichoplax adhaerens)
  • tropical clawed frog (Silurana tropicalis)
  • wheat (Triticum aestivum)
  • worm (Caenorhabditis elegans)

Evolution for CST4 Gene

Gene Tree for CST4 (if available)
Gene Tree for CST4 (if available)
Evolutionary constrained regions (ECRs) for CST4: view image

Paralogs for CST4 Gene

Paralogs for CST4 Gene

(9) SIMAP similar genes for CST4 Gene using alignment to 1 proteins:

  • CYTS_HUMAN Pseudogenes for CST4 Gene

genes like me logo Genes that share paralogs with CST4: view

Variants for CST4 Gene

Sequence variations from dbSNP and Humsavar for CST4 Gene

SNP ID Clin Chr 20 pos Variation AA Info Type
VAR_036549 A breast cancer sample p.Thr77Asn
rs1000247380 -- 23,687,336(-) C/A intron_variant
rs1001248815 -- 23,686,615(-) C/T intron_variant
rs1001314195 -- 23,686,497(-) C/T intron_variant
rs1001399541 -- 23,690,693(-) T/G upstream_transcript_variant

Structural Variations from Database of Genomic Variants (DGV) for CST4 Gene

Variant ID Type Subtype PubMed ID
esv2722303 CNV deletion 23290073
esv3575365 CNV gain 25503493
esv3893322 CNV gain 25118596
nsv1056210 CNV gain 25217958
nsv1057975 CNV loss 25217958
nsv1112348 CNV deletion 24896259
nsv1129405 OTHER inversion 24896259
nsv458950 CNV loss 19166990
nsv458952 CNV gain 19166990
nsv482066 CNV gain 20164927
nsv585724 CNV loss 21841781
nsv585725 CNV gain 21841781
nsv965862 CNV duplication 23825009

Variation tolerance for CST4 Gene

Residual Variation Intolerance Score: 94% of all genes are more intolerant (likely to be disease-causing)
Gene Damage Index Score: 0.69; 14.61% of all genes are more intolerant (likely to be disease-causing)

Additional Variant Information for CST4 Gene

SNPedia medical, phenotypic, and genealogical associations of SNPs for

SNP Genotyping and Copy Number Assay Products

No data available for Polymorphic Variants from UniProtKB/Swiss-Prot for CST4 Gene

Disorders for CST4 Gene

Additional Disease Information for CST4

No disorders were found for CST4 Gene.

No data available for MalaCards , UniProtKB/Swiss-Prot and Genatlas for CST4 Gene

Publications for CST4 Gene

  1. Salivary cystatin SA-III, a potential precursor of the acquired enamel pellicle, is phosphorylated at both its amino- and carboxyl-terminal regions. (PMID: 1898055) Lamkin MS … Oppenheim FG (Archives of biochemistry and biophysics 1991) 3 4 22 58
  2. The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC). (PMID: 15489334) Gerhard DS … MGC Project Team (Genome research 2004) 3 4 58
  3. The DNA sequence and comparative analysis of human chromosome 20. (PMID: 11780052) Deloukas P … Rogers J (Nature 2001) 3 4 58
  4. Characterization of two members (CST4 and CST5) of the cystatin gene family and molecular evolution of cystatin genes. (PMID: 1334620) Saitoh E … Ohnishi K (Agents and actions. Supplements 1992) 3 4 58
  5. The human cystatin gene family: cloning of three members and evolutionary relationship between cystatins and Bowman-Birk type proteinase inhibitors. (PMID: 1801729) Saitoh E … Ohnishi K (Biomedica biochimica acta 1991) 2 3 58

Products for CST4 Gene

Sources for CST4 Gene

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