Open Close
General Information
Symbol
Dmel\ttv
Species
D. melanogaster
Name
tout-velu
Annotation Symbol
CG10117
Feature Type
FlyBase ID
FBgn0265974
Gene Model Status
Stock Availability
Enzyme Name (EC)
Glucuronosyltransferase (2.4.1.17)
Glucuronosyl-N-acetylglucosaminyl-proteoglycan 4-alpha-N- acetylglucosaminyltransferase (2.4.1.224)
N-acetylglucosaminyl-proteoglycan 4-beta-glucuronosyltransferase (2.4.1.225)
Gene Snapshot
Also Known As

DEXT1, l(2)05282, l(2)00681, l(2)k03617

Key Links
Genomic Location
Cytogenetic map
Sequence location
2R:14,526,267..14,587,917 [+]
Recombination map

2-71

RefSeq locus
NT_033778 REGION:14526267..14587917
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (20 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000869020
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000869020
(assigned by GO_Central )
inferred from sequence or structural similarity with UniProtKB:O77783
(assigned by UniProt )
inferred from biological aspect of ancestor with PANTHER:PTN000115158
(assigned by GO_Central )
Biological Process (11 terms)
Terms Based on Experimental Evidence (9 terms)
CV Term
Evidence
References
inferred from genetic interaction with FLYBASE:Sdc; FB:FBgn0010415
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
(assigned by UniProt )
inferred from genetic interaction with UniProtKB:Q9Y169
(assigned by UniProt )
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
(assigned by UniProt )
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from sequence or structural similarity with UniProtKB:O77783
(assigned by UniProt )
inferred from electronic annotation with InterPro:IPR004263
(assigned by InterPro )
Cellular Component (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000115158
(assigned by GO_Central )
inferred from electronic annotation with InterPro:IPR015338
(assigned by InterPro )
non-traceable author statement
(assigned by UniProt )
Protein Family (UniProt)
Belongs to the glycosyltransferase 47 family. (Q9V730)
Catalytic Activity (EC)
Experimental Evidence
UDP-alpha-D-glucuronate + N-acetyl-alpha-D-glucosaminyl-(1->4)-beta-D- glucuronosyl-proteoglycan = UDP + beta-D-glucuronosyl-(1->4)-N-acetyl- alpha-D-glucosaminyl-(1->4)-beta-D-glucuronosyl-proteoglycan (2.4.1.225)
Predictions / Assertions
UDP-glucuronate + acceptor = UDP + acceptor beta-D-glucuronoside (2.4.1.17)
UDP-N-acetyl-D-glucosamine + beta-D-glucuronosyl-(1->4)-N-acetyl-alpha-D- glucosaminyl-proteoglycan = UDP + N-acetyl-alpha-D-glucosaminyl-(1->4)- beta-D-glucuronosyl-(1->4)-N-acetyl-alpha-D-glucosaminyl-proteoglycan (2.4.1.224)
UDP-alpha-D-glucuronate + N-acetyl-alpha-D-glucosaminyl-(1->4)-beta-D- glucuronosyl-proteoglycan = UDP + beta-D-glucuronosyl-(1->4)-N-acetyl- alpha-D-glucosaminyl-(1->4)-beta-D-glucuronosyl-proteoglycan (2.4.1.225)
Summaries
Gene Group (FlyBase)
PROTEOGLYCAN N-ACETYLGLUCOSAMINYLTRANSFERASES -
Proteoglycan N-acetylglucosaminyltransferases catalyze the transfer of N-acetylglucosamine (GlcNAc) to proteoglycans, such as heparan sulfate proteoglycans (HSPGs). HSPGs consist of a protein core to which heparan sulfate (HS) glycosaminoglycan (GAG) chains are attached. The biosynthesis of HS GAG chains is initiated by the formation of a GAG-protein linkage region consisting of a tetrasaccharide attached to specific serine residues in a proteoglycan core protein. Both the attachment of the first GlcNAc to the GAG-protein linkage region and the subsequent polymer formation are catalyzed by members of the hereditary multiple exostoses (EXT) gene family of tumor suppressors, or their orthologs. HSPGs are implicated in regulating the signalling activities of secreted morphogen molecules including Wingless (Wg), Hedgehog (Hh) and Decapentaplegic (Dpp). (Adapted from FBrf0174560.)
GLUCURONOSYLTRANSFERASES -
Glucuronosyltransferases catalyze the conjugation of glucuronic acid from UDP-glucuronate to an acceptor, typically a small lipophilic molecule. (Adapted from PMID:26453144.)
Protein Function (UniProtKB)
Glycosyltransferase required for the biosynthesis of heparan-sulfate and responsible for the alternating addition of beta-1-4-linked glucuronic acid (GlcA) and alpha-1-4-linked N-acetylglucosamine (GlcNAc) units to nascent heparan sulfate chains. Botv is the trigger of heparan sulfate chain initiation and polymerization takes place by a complex of ttv and sotv. Plays a central role in the diffusion of morphogens hedgehog (hh), wingless (wg) and decapentaplegic (dpp) via its role in heparan sulfate proteoglycans (HSPGs) biosynthesis which are required for movement of hh, dpp and wg morphogens.
(UniProt, Q9V730)
Summary (Interactive Fly)

acetylglucosaminyltransferase - involved in heparan sulfate proteoglycan biosynthesis - affects diffusion of Wingless, Hedgehog and Decapentaplegic

Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
3

Please see the JBrowse view of Dmel\ttv 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
Comments on Gene Model

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Gene model reviewed during 5.46

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

gene_with_stop_codon_read_through ; SO:0000697

Stop-codon suppression (UAG) postulated; FBrf0234051.

Gene model reviewed during 6.25

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0087495
3994
760
FBtr0333164
6899
760
FBtr0333165
2351
299
FBtr0474168
3994
772
Additional Transcript Data and Comments
Reported size (kB)

3.8 (longest cDNA)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0086624
87.3
760
9.37
FBpp0305367
87.3
760
9.37
FBpp0305368
34.4
299
9.71
FBpp0423167
88.7
772
9.51
Polypeptides with Identical Sequences

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

760 aa isoforms: ttv-PA, ttv-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

760 (aa); 80 (kD observed)

Comments
External Data
Subunit Structure (UniProtKB)

Interacts with sau (PubMed:23720043).

(UniProt, Q9V730)
Crossreferences
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\ttv 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
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

Epitope-tagged ttv protein colocalizes with Golgi apparatus marker βCop and endoplasmic reticulum marker Bip, but not with plasma membrane markers Arm or shg, indicating that it is localized to the secretory apparatus, and not to the plasma membrane.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\ttv in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 69 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 9 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ttv
Transgenic constructs containing regulatory region of ttv
Deletions and Duplications ( 6 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
anterior fascicle & bouton
Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (7)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
13 of 15
Yes
Yes
 
2  
6 of 15
No
Yes
1 of 15
No
No
5  
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (4)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
14 of 15
Yes
Yes
7 of 15
No
Yes
2 of 15
No
Yes
1 of 15
No
No
Rattus norvegicus (Norway rat) (6)
12 of 13
Yes
Yes
7 of 13
No
Yes
2 of 13
No
Yes
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (4)
8 of 12
Yes
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (10)
12 of 15
Yes
Yes
11 of 15
No
Yes
10 of 15
No
Yes
8 of 15
No
Yes
2 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (2)
10 of 15
Yes
Yes
2 of 15
No
No
Arabidopsis thaliana (thale-cress) (38)
3 of 9
Yes
Yes
3 of 9
Yes
Yes
3 of 9
Yes
Yes
3 of 9
Yes
Yes
3 of 9
Yes
Yes
3 of 9
Yes
Yes
3 of 9
Yes
Yes
3 of 9
Yes
Yes
3 of 9
Yes
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
No
2 of 9
No
No
2 of 9
No
No
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
Yes
2 of 9
No
No
2 of 9
No
Yes
2 of 9
No
No
2 of 9
No
No
2 of 9
No
Yes
2 of 9
No
No
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG0919030F )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG0915065W )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0678 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Rhodnius prolixus
Kissing bug
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Zootermopsis nevadensis
Nevada dampwood termite
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X062P )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0BDR )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (2)
4 of 10
4 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 3 )
    Modifiers Based on Experimental Evidence ( 2 )
    Disease Associations of Human Orthologs (via DIOPT v8.0 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    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.
    Dmel gene
    Ortholog showing functional complementation
    Supporting References
    Interactions
    Summary of Physical Interactions
    esyN Network Diagram
    Show neighbor-neighbor interactions:
    Select Layout:
    Legend:
    Protein
    RNA
    Selected Interactor(s)
    Interactions Browser

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    esyN Network Diagram
    esyN Network Key:
    Suppression
    Enhancement

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Subunit Structure (UniProtKB)
    Interacts with sau (PubMed:23720043).
    (UniProt, Q9V730 )
    Linkouts
    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
    External Data
    Linkouts
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map

    2-71

    Cytogenetic map
    Sequence location
    2R:14,526,267..14,587,917 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    51A7-51B1
    Left limit from in situ hybridisation (FBrf0157097) Right limit from in situ hybridisation (FBrf0157097)
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    51B1-51B1
    51B5-51B5
    (determined by in situ hybridisation)
    51B1-51B5
    (determined by in situ hybridisation)
    51A-51A
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes

    36 recombinants in 31,272 progeny.

    Stocks and Reagents
    Stocks (37)
    Genomic Clones (46)
    cDNA Clones (65)
     

    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 GBrowse 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)
    RNAi and Array Information
    Linkouts
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    Antibody Information
    Laboratory Generated Antibodies
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of

    Source for merge of: ttv l(2)00681

    Source for merge of: ttv l(2)05282 l(2)k03617

    Source for merge of: ttv l(2)SH0631

    Additional comments

    l(2)k03906b may correspond to ttv: the P{lacW}l(2)k03906bk03906b insertion maps within the transcription unit.

    Other Comments

    ttv is required for normal germ cell migration in the developing embryo.

    ttv is necessary for normal levels of heparan sulphate glycosaminoglycan modification of dally and dlp gene products.

    ttv is required for transport of morphogens (hh, wg, and dpp) in the developing imaginal discs.

    8 alleles of l(2)SH0631 recovered in a P-insertion screen.

    ttv is involved in heparan sulfate proteoglycan synthesis.

    ttv is required for the diffusion of the hh gene product. ttv is a member of the EXT gene family.

    Mutants show maternal effect segment polarity phenotypes.

    Mutants isolated in a screen of the second chromosome identifying genes affecting disc morphology.

    Isolated in a screen for lethal mutations affecting segmentation.

    The autosomal "FLP-DFS" technique (using the P{ovoD1-18} P{FRT(whs)} P{hsFLP} chromosomes) has been used to identify the specific maternal effect phenotype for the zygotic lethal mutation.

    Origin and Etymology
    Discoverer
    Etymology

    'tout-velu' means 'all hair' in French.

    Identification
    External Crossreferences and Linkouts ( 58 )
    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/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
    UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
    Other crossreferences
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
    KEGG Genes - Molecular building blocks of life in the genomic space.
    Linkouts
    ApoDroso - Functional genomic database for photoreceptor development, survival and function
    DroID - A comprehensive database of gene and protein interactions.
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    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.
    Synonyms and Secondary IDs (21)
    Reported As
    Symbol Synonym
    l(2)SH0631
    l(2)SH2 0631
    l(2)k00115
    Secondary FlyBase IDs
    • FBgn0065697
    • FBgn0020245
    • FBgn0010469
    • FBgn0010592
    • FBgn0024670
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (189)