FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Hs2st
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General Information
Symbol
Dmel\Hs2st
Species
D. melanogaster
Name
Heparan sulfate 2-O-sulfotransferase
Annotation Symbol
CG10234
Feature Type
protein_coding_gene
FlyBase ID
FBgn0024230
Gene Model Status
Current
Stock Availability
9 publicly available
Gene Summary
Heparan sulfate 2-O-sulfotransferase (Hs2st) encodes a heparan sulfate modifying enzyme, which transfers a sulfate group to 2-O position of glucuronic or iduronic acid residues of heparan sulfate. Loss of Hs2st function eliminates 2-O sulfation, but increases 6-O sulfation performed by the product of Hs6st. Simultaneous elimination of Hs6st and Hs2st functions disrupts FGF, Wg and Dpp signaling pathways. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Key Links
Genomic Location
Cytogenetic map
37E1-37E1
Sequence location
Recombination map
2-54
RefSeq locus
NT_033779 REGION:19442833..19444750
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (9 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
enables heparan sulfate 2-sulfotransferase activity
inferred from direct assay
(Xu et al., 2007, Kamimura et al., 2006)
enables protein binding
inferred from physical interaction with UniProtKB:A1Z6M8
(Dejima et al., 2013)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
enables heparan sulfate 2-sulfotransferase activity
inferred from biological aspect of ancestor with PANTHER:PTN000951108
(GO Reference Genome Project, 2011-)
enables sulfotransferase activity
inferred from electronic annotation with InterPro:IPR005331, InterPro:IPR007734
(InterPro Project Members, 2004-)
NOT enables chondroitin 2-sulfotransferase activity
traceable author statement
(Xu et al., 2007)
Biological Process (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
involved_in heparan sulfate proteoglycan biosynthetic process
inferred from mutant phenotype
(Nakato et al., 2019, Kamimura et al., 2011, Kamimura et al., 2006)
involved_in open tracheal system development
inferred from mutant phenotype
(Kamimura et al., 2006)
inferred from genetic interaction with FLYBASE:Hs6st; FB:FBgn0038755
(Kamimura et al., 2006)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in heparan sulfate proteoglycan biosynthetic process
inferred from biological aspect of ancestor with PANTHER:PTN000951108
(GO Reference Genome Project, 2011-)
involved_in signal transduction
non-traceable author statement
(Kusano et al., 2003)
Cellular Component (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in Golgi apparatus
inferred from sequence or structural similarity with UniProtKB:Q8R3H7
(Gene Ontology Curators, 2002-)
located_in membrane
inferred from electronic annotation with InterPro:IPR005331, InterPro:IPR007734
(InterPro Project Members, 2004-)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
Belongs to the sulfotransferase 3 family. (P25722)
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Heparan sulfate 2-O-sulfotransferase (Hs2st) encodes a heparan sulfate modifying enzyme, which transfers a sulfate group to 2-O position of glucuronic or iduronic acid residues of heparan sulfate. Loss of Hs2st function eliminates 2-O sulfation, but increases 6-O sulfation performed by the product of Hs6st. Simultaneous elimination of Hs6st and Hs2st functions disrupts FGF, Wg and Dpp signaling pathways. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
HEPARAN SULFATE SULFOTRANSFERASES -
Heparan sulfate sulfotransferases transfers the sulfo group from 3'-phosphoadenosine 5'-phosphosulfate on to heparan sulfate. (Adapted from FBrf0202009).
Pathway (FlyBase)
HEPARAN SULFATE PROTEOGLYCAN BIOSYNTHESIS -
Heparan sulfate proteoglycans (HSPGs) are glycoconjugates that consist of a core protein linked to one or more linear heparan sulfate (HS) chains. The chains are composed of alternating N-acetyl-D-glucosamine and glucuronic acid or iduronic acid that can be variably N- and O-sulfated. HS synthesis occurs in the Golgi apparatus. It starts with the formation of a tetrasaccharide linker on the core protein followed by the alternating addition of glucuronic acid and N-acetylglucosamine residues. Glucuronic acid can undergo epimerization into iduronic acid, which together with N-acetylglucosamine residues, can be sulfated. HSPGs are then transported to the plasma membrane where they can be processed further by endosulfatases such as Sulf1 that remove sulfate groups from specific sites. (Adapted from FBrf0232460 and FBrf0228609.)
Protein Function (UniProtKB)
Catalyzes the transfer of sulfate to the C2-position of selected hexuronic acid residues within the maturing heparan sulfate (HS).
(UniProt, P25722)
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\Hs2st for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry P25722)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.47

Gene model reviewed during 5.55

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Hs2st-RA
FBtr0081288
NM_057991
1652
349
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
Hs2st-PA
FBpp0080826
41.3
349
9.55
P25722
NP_477339
AAF53800
Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Homotrimer.

(UniProt, P25722)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\Hs2st using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-1.25

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4
organism
(Herr and Harris, 2004)
embryonic stage 4 -- 6
organism
(Fisher et al., 2012)
embryonic stage 7 -- 8
organism | ubiquitous
(Fisher et al., 2012)
third instar larval stage
lamina furrow
(Dejima et al., 2013)
Additional Descriptive Data

Hs2st is coexpressed with hob

(Herr and Harris, 2004)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Hs2st in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 5 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 11 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Hs2st
Transgenic constructs containing regulatory region of Hs2st
Aberrations (Deficiencies and Duplications) ( 5 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (2)
Hsap\HS2ST1
14 of 14
Yes
Yes
1  
Hsap\UST
3 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (2)
Rnor\Hs2st1
14 of 14
Yes
Yes
Rnor\Ust
3 of 14
No
No
Mus musculus (laboratory mouse) (2)
Mmus\Hs2st1
13 of 14
Yes
Yes
Mmus\Ust
3 of 14
No
No
Xenopus tropicalis (Western clawed frog) (3)
Xtro\hs2st1
11 of 13
Yes
Yes
Xtro\ttc3
1 of 13
No
Yes
Xtro\ust
1 of 13
No
No
Danio rerio (Zebrafish) (4)
Drer\hs2st1a
13 of 14
Yes
Yes
Drer\hs2st1b
13 of 14
Yes
Yes
Drer\usta
3 of 14
No
No
Drer\LOC100331876
2 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (1)
Cele\hst-2
13 of 14
Yes
Yes
Anopheles gambiae (African malaria mosquito) (2)
Agam\AgaP_AGAP008166
12 of 12
Yes
Yes
Agam\AgaP_AGAP006058
3 of 12
No
No
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Hs2st. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (1)
pip
5 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Interaction
    References
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    HS2ST1; heparan sulfate 2-O-sulfotransferase 1
    14 of 14
    604844
       
      UST; uronyl 2-sulfotransferase
      3 of 14
      610752
          Functional Complementation Data
          Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
          Interactions
          Summary of Physical Interactions
          Summary of Genetic Interactions
          Interaction Browsers
          View in FlyBase Interactions Browser   View in MIST tool (external link)
          Please look at the allele data for full details of the genetic interactions
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          Hs2st
          enhanceable
          Hsepi
          (Dejima et al., 2013)
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          sfl
          suppressible
          Hs2st
          (Kamimura et al., 2011)
          External Data
          Subunit Structure (UniProtKB)
          Homotrimer.
          (UniProt, P25722 )
          Linkouts
          DroID - A comprehensive database of gene and protein interactions.
          MIST (protein-protein) - An integrated Molecular Interaction Database
          Pathways
          Signaling Pathways (FlyBase)
          Metabolic Pathways
          FlyBase
          HEPARAN SULFATE PROTEOGLYCAN BIOSYNTHESIS
          External Links
          FlyCyc Pathways - Pathways from a BioCyc PGDB for Dmel
          KEGG Metabolic Pathways - A collection of manually drawn metabolic pathway maps representing knowledge of molecular interaction, reaction and relation networks.
          Reactome Metabolic Pathways - An open-source, open access, manually curated and peer-reviewed metabolic pathway database.
          External Data
          Linkouts
          KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
          Class of Gene
          Genomic Location and Detailed Mapping Data
          Chromosome (arm)
          2L
          Recombination map
          2-54
          Cytogenetic map
          37E1-37E1
          Sequence location
          FlyBase Computed Cytological Location
          Cytogenetic map
          Evidence for location
          37E1-37E1
          Limits computationally determined from genome sequence between P{EP}dntEP2158 and P{EP}EP623&P{PZ}spi01068
          Experimentally Determined Cytological Location
          Cytogenetic map
          Notes
          References
          Experimentally Determined Recombination Data
          Location

          2-54

          (FlyBase, 2016)

          2-53.7

          (Comeron et al., 2012)
          Left of (cM)
          Right of (cM)
          Notes
          Stocks and Reagents
          Stocks (9)
          Genomic Clones (17)
           

          Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

          cDNA Clones (30)
           

          Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see JBrowse for alignment of the cDNAs and ESTs to the gene model.

          cDNA clones, fully sequenced
          BDGP DGC clones
          Other clones
          Drosophila Genomics Resource Center cDNA clones

          For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

          cDNA Clones, End Sequenced (ESTs)
          BDGP DGC clones
          Other clones
          RNAi and Array Information
          Linkouts
          DRSC - Results frm RNAi screens
          Antibody Information
          Laboratory Generated Antibodies
           
          Commercially Available Antibodies
           
          Cell Line Information
          Publicly Available Cell Lines
           
            Other Stable Cell Lines
             
              Other Comments

              Heparan sulfate samples from Hs2st mutants show a loss of 2-O sulfated disaccharide units, as well as an increase in 6-O sulfated disaccharides. The level of total sulfate groups on heparan sulfate is not affected, and the total charge of heparan sulfate is almost wild-type.

              (Kamimura et al., 2006)

              dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.

              (Kiger et al., 2003)
              Relationship to Other Genes
              Source for database merge of

              Source for merge of: Hs2st CG10234

              (Bayraktaroglu, 2000.4.13)
              Additional comments
              Nomenclature History
              Source for database identify of
              Nomenclature comments
              Etymology
              Synonyms and Secondary IDs (14)
              Datasets (0)
              Study focus (0)
              Experimental Role
              Project
              Project Type
              Title
              Study result (0)
              Result
              Result Type
              Title
              External Crossreferences and Linkouts ( 43 )
              Sequence Crossreferences
              NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
              GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
              GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
              RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
              UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
              UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
              Other crossreferences
              AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
              BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
              DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
              EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
              FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
              FlyMine - An integrated database for Drosophila genomics
              InterPro - A database of protein families, domains and functional sites
              KEGG Genes - Molecular building blocks of life in the genomic space.
              MARRVEL_MODEL - MARRVEL (model organism gene)
              Linkouts
              Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
              DroID - A comprehensive database of gene and protein interactions.
              DRSC - Results frm RNAi screens
              Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
              FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
              FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
              FlyCyc Pathways - Pathways from a BioCyc PGDB for Dmel
              Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
              Flygut - An atlas of the Drosophila adult midgut
              FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
              iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
              KEGG Metabolic Pathways - A collection of manually drawn metabolic pathway maps representing knowledge of molecular interaction, reaction and relation networks.
              KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
              MIST (protein-protein) - An integrated Molecular Interaction Database
              Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
              Reactome Metabolic Pathways - An open-source, open access, manually curated and peer-reviewed metabolic pathway database.
              References (56)