FB2025_04 , released October 2, 2025
Gene: Dmel\Hs6st
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General Information
Symbol
Dmel\Hs6st
Species
D. melanogaster
Name
Heparan sulfate 6-O-sulfotransferase
Annotation Symbol
CG4451
Feature Type
protein_coding_gene
FlyBase ID
FBgn0038755
Gene Model Status
Current
Stock Availability
14 publicly available
Gene Summary
Heparan sulfate 6-O-sulfotransferase (Hs6st) encodes a heparan sulfate modifying enzyme, which transfers a sulfate group to the 6-O position of glucosamine residues of heparan sulfate. Loss of Hs6st eliminates 6-O sulfation, but increases 2-O sulfation performed by the product of Hs2st. Simultaneous elimination of Hs6st and Hs2st disrupts FGF, Wg and Dpp signaling pathways. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Also Known As

dHS6ST, 6-O sulfotransferase

Key Links
Genomic Location
Cytogenetic map
92B8-92C1
Sequence location
Recombination map
3-67
RefSeq locus
NT_033777 REGION:19929277..20008512
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 (10 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
enables heparan sulfate 6-sulfotransferase activity
inferred from mutant phenotype
(Kamimura et al., 2006)
enables protein binding
inferred from physical interaction with UniProtKB:A1Z6M8
(Dejima et al., 2013)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables heparan sulfate 6-sulfotransferase activity
inferred from biological aspect of ancestor with PANTHER:PTN000301790
(GO Reference Genome Project, 2011-)
enables sulfotransferase activity
inferred from electronic annotation with InterPro:IPR005331, InterPro:IPR010635
(InterPro Project Members, 2004-)
Biological Process (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
involved_in chaeta development
inferred from mutant phenotype
(Kamimura et al., 2011)
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 imaginal disc-derived wing margin morphogenesis
inferred from mutant phenotype
(Kamimura et al., 2011)
involved_in open tracheal system development
inferred from mutant phenotype
(Kamimura et al., 2006)
inferred from genetic interaction with FLYBASE:Hs2st; FB:FBgn0024230
(Kamimura et al., 2006)
involved_in regulation of imaginal disc growth
inferred from mutant phenotype
(Kamimura et al., 2011)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
involved_in heparan sulfate proteoglycan biosynthetic process
inferred from biological aspect of ancestor with PANTHER:PTN000301790
(GO Reference Genome Project, 2011-)
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:Q9QYK5
(Gene Ontology Curators, 2002-)
located_in membrane
inferred from electronic annotation with InterPro:IPR005331, InterPro:IPR010635
(InterPro Project Members, 2004-)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Catalytic Activity (EC/Rhea)
heparan sulfate 6-sulfotransferase activity
RHEA 56604:
Summaries
Gene Snapshot
Heparan sulfate 6-O-sulfotransferase (Hs6st) encodes a heparan sulfate modifying enzyme, which transfers a sulfate group to the 6-O position of glucosamine residues of heparan sulfate. Loss of Hs6st eliminates 6-O sulfation, but increases 2-O sulfation performed by the product of Hs2st. Simultaneous elimination of Hs6st and Hs2st 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.)
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\Hs6st 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 Q9VDR6)

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

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Hs6st-RA
FBtr0083846
NM_079685
2657
462
Hs6st-RB
FBtr0335240
NM_001275821
3100
462
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
Hs6st-PA
FBpp0083255
52.7
462
8.12
Q9VDR6
NP_524409
AAF55724
Hs6st-PB
FBpp0307227
52.7
462
8.12
Q9VDR6
NP_001262750
AGB96130
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

462 aa isoforms: Hs6st-PA, Hs6st-PB
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
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\Hs6st using the Feature Mapper tool.

External Data
Crossreferences
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).

-0.86

Transcript Expression
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\Hs6st 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
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 4-6
  • Stages(s) 11-12
  • Stages(s) 13-16
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 23 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 6 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Hs6st
Transgenic constructs containing regulatory region of Hs6st
Aberrations (Deficiencies and Duplications) ( 0 )
Inferred from experimentation ( 0 )
Inferred from location ( 4 )
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) (3)
Hsap\HS6ST1
13 of 14
Yes
Yes
1  
Hsap\HS6ST2
12 of 14
No
Yes
1  
Hsap\HS6ST3
12 of 14
No
Yes
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) (3)
Rnor\Hs6st1
13 of 14
Yes
Yes
Rnor\Hs6st2
12 of 14
No
Yes
Rnor\Hs6st3
1 of 14
No
Yes
Mus musculus (laboratory mouse) (3)
Mmus\Hs6st1
12 of 14
Yes
Yes
Mmus\Hs6st2
12 of 14
Yes
Yes
Mmus\Hs6st3
12 of 14
Yes
Yes
Xenopus tropicalis (Western clawed frog) (8)
Xtro\hs6st1
10 of 13
Yes
Yes
Xtro\hs6st2
8 of 13
No
Yes
Xtro\crhbp
3 of 13
No
No
Xtro\hs6st3
3 of 13
No
Yes
Xtro\eny2
2 of 13
No
No
Xtro\cdk19
1 of 13
No
No
Xtro\gal3st2
1 of 13
No
Yes
Xtro\nr1d1
1 of 13
No
No
Danio rerio (Zebrafish) (5)
Drer\hs6st1b
13 of 14
Yes
Yes
Drer\hs6st1a
12 of 14
No
Yes
Drer\hs6st2
12 of 14
No
Yes
Drer\hs6st3b
12 of 14
No
Yes
Drer\hs6st3a
11 of 14
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (1)
Cele\hst-6
13 of 14
Yes
Yes
Anopheles gambiae (African malaria mosquito) (2)
Agam\AgaP_AGAP001444
12 of 12
Yes
Yes
Agam\AgaP_AGAP001439
5 of 12
No
Yes
Arabidopsis thaliana (thale-cress) (1)
Atha\TPST
4 of 13
Yes
Yes
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:Hs6st. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
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 ( 2 )
    Modifiers Based on Experimental Evidence ( 2 )
    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?
    HS6ST1; heparan sulfate 6-O-sulfotransferase 1
    13 of 14
    604846
     
    HS6ST2; heparan sulfate 6-O-sulfotransferase 2
    12 of 14
    300545
     
    HS6ST3; heparan sulfate 6-O-sulfotransferase 3
    12 of 14
    609401
        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
        Hs6st
        enhanceable
        Hsepi
        (Dejima et al., 2013)
        Hs6st
        suppressible
        Hsepi
        (Kamimura et al., 2011)
        Hs6st
        enhanceable
        Papst2
        (Goda et al., 2006)
        Hs6st
        enhanceable
        β3GalTII
        (Ueyama et al., 2008)
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        Hsepi
        suppressible
        Hs6st
        (Dejima et al., 2013)
        Sulf1
        suppressible
        Hs6st
        (Kleinschmit et al., 2010)
        β3GalTII
        enhanceable
        Hs6st
        (Ueyama et al., 2008)
        External Data
        Linkouts
        BioGRID - A database of protein and genetic interactions.
        DroID - A comprehensive database of gene and protein interactions.
        MIST (genetic) - An integrated Molecular Interaction Database
        MIST (protein-protein) - An integrated Molecular Interaction Database
        Pathways
        Signaling Pathways (FlyBase)
        Metabolic Pathways
        FlyBase
        HEPARAN SULFATE PROTEOGLYCAN BIOSYNTHESIS
        External Links
        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)
        3R
        Recombination map
        3-67
        Cytogenetic map
        92B8-92C1
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        92B8-92C1
        ; Limits computationally determined from genome sequence between P{PZ}l(3)1058510585 and P{EP}SyndEP409
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        Experimentally Determined Recombination Data
        Location

        3-67

        (FlyBase, 2016)

        3-69.8

        (Comeron et al., 2012)
        Left of (cM)
        Right of (cM)
        Notes
        Stocks and Reagents
        Stocks (14)
        Genomic Clones (52)
        cDNA Clones (129)
         

        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

            Levels of 2-O sulfated disaccharides are strikingly elevated in Hs6st mutants. 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)

            Shows particularly robust cycling of transcription in adult heads, as assessed by expression analysis using high density oligonucleotide arrays with probe generated during three 12-point time course experiments over the course of 6 days.

            (Claridge-Chang et al., 2001)
            Relationship to Other Genes
            Source for database merge of

            Source for merge of: Hs6st CG4451 CG17188

            (Misra, 2001.8.10)

            Source for merge of: CG11683 CG18533

            (FlyBase, 1996-)
            Additional comments

            Source for merge of CG11683 CG18533 was sequence comparison ( date:001104 ).

            (FlyBase, 1996-)

            Source for identity of Hs6st CG11683 CG4451 was sequence comparison ( date:010522 ).

            (FlyBase, 1996-)
            Nomenclature History
            Source for database identify of

            Source for identity of: Hs6st CG11683 CG4451

            (FlyBase, 1996-)
            Nomenclature comments
            Etymology
            Synonyms and Secondary IDs (14)
            Reported As
            Symbol Synonym
            CG11683
            CG17188
            (Misra, 2001.8.10)
            CG18533
            CG4451
            (Yamamoto-Hino et al., 2015, Dani et al., 2012, Keleman et al., 2009.8.5, Perkins et al., 2009, Quinones-Coello, 2007, Häcker et al., 2005)
            DmHs6st
            (Nybakken and Perrimon, 2002)
            HS6ST
            (Kamimura and Maeda, 2017)
            Hs6st
            (Miao et al., 2024, Owings and Chow, 2024, Kamimura and Maeda, 2021, van der Graaf et al., 2021, Knudsen et al., 2020, Rushton et al., 2020, Theis et al., 2020, Meltzer et al., 2019, Nakato et al., 2019, Al-Ramahi et al., 2018, Gene Disruption Project members, 2018-, Takemura and Nakato, 2017, Levings et al., 2016, Nakato and Li, 2016, Ou et al., 2016, Duff et al., 2015, Gene Disruption Project members, 2015-, Ashwal-Fluss et al., 2014, Chandran et al., 2014, Dejima et al., 2013, Sekine et al., 2013, Dani et al., 2012, Kamimura et al., 2011, Zhou et al., 2011, Kleinschmit et al., 2010, Quinones-Coello, 2007, Kamimura et al., 2006, Häcker et al., 2005)
            dHS6ST
            (Yamamoto-Hino et al., 2015, Ueyama et al., 2008, Goda et al., 2006, Butler et al., 2003, Ghabrial et al., 2003, Ebner et al., 2002, Nakato and Kimata, 2002, Kamimura et al., 2001, Kamimura, 2000.11.29)
            hs6st
            (Nishihara, 2020, Dani et al., 2012, Gistelinck et al., 2012, Shulman et al., 2011, Brennan, 2007, Sandmann et al., 2007)
            Name Synonyms
            6-O sulfotransferase
            (Kamimura et al., 2011, Kamimura et al., 2006)
            6-O-sulfotransferase
            (Nishihara, 2010)
            HS 6-O-sulfotransferase
            (Nishihara, 2020)
            Heparan sulfate 6-O-sulfotransferase
            (Kamimura et al., 2001)
            heparan sulfate 6-O-sulfotransferase
            (Shulman et al., 2011)
            Secondary FlyBase IDs
            • FBgn0038754
            • FBgn0038759
            • FBgn0042209
            Datasets (0)
            Study focus (0)
            Experimental Role
            Project
            Project Type
            Title
            Study result (0)
            Result
            Result Type
            Title
            External Crossreferences and Linkouts ( 55 )
            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/TrEMBL - Automatically annotated and unreviewed 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
            BioGRID - A database of protein and genetic interactions.
            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
            FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
            FlyCyc Genes - Genes 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 (genetic) - An integrated Molecular Interaction Database
            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 (86)