FB2025_05 , released December 11, 2025
Gene: Dmel\ttv
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General Information
Symbol
Dmel\ttv
Species
D. melanogaster
Name
tout-velu
Annotation Symbol
CG10117
Feature Type
protein_coding_gene
FlyBase ID
FBgn0265974
Gene Model Status
Current
Stock Availability
39 publicly available
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 Summary
tout-velu (ttv) encodes a glycosyltransferase that forms a complex with sotv to catalyze the alternating transfer of glucuronic acid (GlcA) and N-acetylglucosamine (GlcNAc) to their respective acceptors, GlcNAc and GlcA, during the elongation of the heparan sulfate glycosaminoglycan chain. [Date last reviewed: 2024-12-19] (FlyBase Gene Snapshot)
All Summaries
Also Known As

l(2)05282, DEXT1, EXT1, l(2)k03617, l(2)00681

Key Links
Genomic Location
Cytogenetic map
51A7-51B4
Sequence location
Recombination map
2-72
RefSeq locus
NT_033778 REGION:14526267..14587917
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 (14 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
contributes_to glucuronosyl-N-acetylglucosaminyl-proteoglycan 4-alpha-N-acetylglucosaminyltransferase activity
inferred from direct assay
(Izumikawa et al., 2006)
contributes_to N-acetylglucosaminyl-proteoglycan 4-beta-glucuronosyltransferase activity
inferred from direct assay
(Izumikawa et al., 2006)
enables protein binding
inferred from physical interaction with UniProtKB:Q9Y169
(Izumikawa et al., 2006)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
enables acetylglucosaminyltransferase activity
inferred from biological aspect of ancestor with PANTHER:PTN000869020
(GO Reference Genome Project, 2011-)
enables glucuronosyltransferase activity
inferred from biological aspect of ancestor with PANTHER:PTN000869020
(GO Reference Genome Project, 2011-)
enables glycosyltransferase activity
inferred from electronic annotation with InterPro:IPR004263, InterPro:IPR015338
(InterPro Project Members, 2004-)
Biological Process (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
involved_in germ cell migration
inferred from mutant phenotype
(Deshpande et al., 2007)
involved_in glycosaminoglycan biosynthetic process
inferred from mutant phenotype
(Toyoda et al., 2000, Toyoda et al., 2000)
involved_in heparan sulfate proteoglycan biosynthetic process
inferred from mutant phenotype
(Ren et al., 2009, Bornemann et al., 2004, Han et al., 2004, Kim et al., 2002)
inferred from mutant phenotype
(Takei et al., 2004, The et al., 1999)
inferred from direct assay
(Izumikawa et al., 2006)
NOT involved_in germ cell migration
inferred from mutant phenotype
(Renault et al., 2009)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in glycoprotein biosynthetic process
inferred from electronic annotation with InterPro:IPR004263
(InterPro Project Members, 2004-)
involved_in heparan sulfate proteoglycan biosynthetic process
inferred from sequence or structural similarity with UniProtKB:O77783
(Gene Ontology Curators, 2002-)
Cellular Component (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
located_in endoplasmic reticulum
inferred from direct assay
(Han et al., 2004, The et al., 1999)
part_of EXT1-EXT2 complex
inferred from direct assay
(Han et al., 2004)
located_in Golgi apparatus
inferred from direct assay
(Han et al., 2004)
inferred from direct assay
(The et al., 1999)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in Golgi apparatus
inferred from biological aspect of ancestor with PANTHER:PTN000115158
(GO Reference Genome Project, 2011-)
located_in membrane
inferred from electronic annotation with InterPro:IPR015338
(InterPro Project Members, 2004-)
non-traceable author statement
(Bellaiche et al., 1998)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
Belongs to the glycosyltransferase 47 family. (Q9V730)
Protein Signatures (InterPro)
Catalytic Activity (EC/Rhea)
glucuronosyltransferase activity
glucuronate acceptor + UDP-alpha-D-glucuronate = acceptor beta-D- glucuronoside + UDP + H(+) (2.4.1.17)
RHEA 21032: glucuronosyl-N-acetylglucosaminyl-proteoglycan 4-alpha-N-acetylglucosaminyltransferase activity
3-O-{[(1->4)-beta-D-GlcA-(1->4)-alpha-D-GlcNAc](n)-(1->4)-beta-D-GlcA- (1->3)-beta-D-Gal-(1->3)-beta-D-Gal-(1->4)-beta-D-Xyl}-L-seryl-[protein] + UDP-N-acetyl-alpha-D-glucosamine = 3-O-{alpha-D-GlcNAc-[(1->4)-beta-D- GlcA-(1->4)-alpha-D-GlcNAc](n)-(1->4)-beta-D-GlcA-(1->3)-beta-D-Gal- (1->3)-beta-D-Gal-(1->4)-beta-D-Xyl}-L-seryl-[protein] + UDP + H(+) (2.4.1.224)
RHEA 16213: N-acetylglucosaminyl-proteoglycan 4-beta-glucuronosyltransferase activity
3-O-{alpha-D-GlcNAc-[(1->4)-beta-D-GlcA-(1->4)-alpha-D-GlcNAc](n)-(1->4)- beta-D-GlcA-(1->3)-beta-D-Gal-(1->3)-beta-D-Gal-(1->4)-beta-D-Xyl}-L- seryl-[protein] + UDP-alpha-D-glucuronate = 3-O-{[(1->4)-beta-D-GlcA- (1->4)-alpha-D-GlcNAc](n+1)-(1->4)-beta-D-GlcA-(1->3)-beta-D-Gal-(1->3)- beta-D-Gal-(1->4)-beta-D-Xyl}-L-seryl-[protein] + UDP + H(+) (2.4.1.225)
RHEA 20908:
Summaries
Gene Snapshot
tout-velu (ttv) encodes a glycosyltransferase that forms a complex with sotv to catalyze the alternating transfer of glucuronic acid (GlcA) and N-acetylglucosamine (GlcNAc) to their respective acceptors, GlcNAc and GlcA, during the elongation of the heparan sulfate glycosaminoglycan chain. [Date last reviewed: 2024-12-19]
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.)
EXT1-EXT2 COMPLEX -
The EXT1-EXT2 complex is a heterodimer required for the elongation of heparan sulfate proteoglycan (HSPG) chains. HSPGs are implicated in regulating the signalling activities of secreted morphogen molecules including Wingless, Hedgehog and Decapentaplegic. Each protein monomer within the complex contains distinct glycosyltransferase domains with different activities. (Adapted from PMID:36593275 and FBrf0174560.)
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)
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
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q9V730)

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

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.

Stop-codon suppression (UAG) postulated; FBrf0234051.

Gene model reviewed during 6.25

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
ttv-RA
FBtr0087495
NM_057883
3994
760
ttv-RB
FBtr0333164
NM_001274042
6899
760
ttv-RC
FBtr0333165
NM_001274043
2351
299
ttv-RD
FBtr0474168
NM_001369957
3994
772
Additional Transcript Data and Comments
Reported size (kB)

3.8 (longest cDNA)

(Bellaiche et al., 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
ttv-PA
FBpp0086624
87.3
760
9.37
Q9V730
NP_477231
AAF58236
ttv-PB
FBpp0305367
87.3
760
9.37
Q9V730
NP_001260971
AGB93503
ttv-PC
FBpp0305368
34.4
299
9.71
D5SHU8
NP_001260972
AGB93504
ttv-PD
FBpp0423167
88.7
772
9.51
QCD26199
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)

(Bellaiche et al., 1998)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with sau (PubMed:23720043).

(UniProt, Q9V730)
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\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
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).

-0.78

Transcript Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
organism
(The et al., 1999)
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.

(The et al., 1999)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in endoplasmic reticulum
inferred from direct assay
(Han et al., 2004, The et al., 1999)
part_of EXT1-EXT2 complex
inferred from direct assay
(Han et al., 2004)
located_in Golgi apparatus
inferred from direct assay
(Han et al., 2004)
inferred from direct assay
(The et al., 1999)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\ttv 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
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
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 70 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 12 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ttv
Transgenic constructs containing regulatory region of ttv
Aberrations (Deficiencies and Duplications) ( 10 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal, with Scer\GAL4Act5C.PU
lethal (with ttv00681b)
lethal (with ttvG00158)
lethal (with ttvk11904)
lethal (with ttvNP3088)
lethal (with ttvSH0631)
lethal (with ttvUM-8373-3)
lethal | embryonic stage | rescuable maternal effect
lethal | recessive
lethal - all die before end of pupal stage
lethal - all die before end of pupal stage, with Scer\GAL4da.G32, ttv00681b
lethal - all die before end of pupal stage, with Scer\GAL4da.G32, ttvUAS.Tag:MYC
lethal - all die before end of pupal stage, with Scer\GAL4elav-C155, ttv00681b
lethal - all die before end of pupal stage, with Scer\GAL4elav-C155, ttvUAS.Tag:MYC
lethal - all die before end of pupal stage, with Scer\GAL4G14, ttv00681b
lethal - all die before end of pupal stage, with Scer\GAL4G14, ttvUAS.Tag:MYC
lethal - all die before end of pupal stage (with ttv205)
lethal - all die before end of pupal stage (with ttv524)
lethal - all die during embryonic stage | germline clone | maternal effect
lethal - all die during embryonic stage | rescuable maternal effect
lethal - all die during P-stage
partially lethal - majority die
partially lethal - majority die, with Scer\GAL4pnr-MD237
partially lethal - majority die | recessive
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4tin.CΔ4
viable (with ttv00681b)
viable (with ttvEP2199)
viable (with ttvEX3)
viable (with ttvEX5)
viable (with ttvEX16)
viable (with ttvEX187)
viable (with ttvEX265)
viable (with ttvEX296)
viable (with ttvEX364)
Other Phenotypes
Allele
abnormal locomotor behavior | pupal stage
abnormal neuroanatomy
abnormal neuroanatomy | cell non-autonomous | somatic clone | third instar larval stage
abnormal neuroanatomy | cell non-autonomous | somatic clone | third instar larval stage, with Scer\GAL4Act5C.PI
abnormal neuroanatomy | cell non-autonomous | third instar larval stage, with Scer\GAL4A58
abnormal neuroanatomy | cell non-autonomous | third instar larval stage, with Scer\GAL4GMR10C12
abnormal neuroanatomy | cell non-autonomous | third instar larval stage, with Scer\GAL4GMR16D01
abnormal neuroanatomy | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
abnormal neurophysiology | pupal stage
increased cell number | adult stage, with Scer\GAL4esg.PU
increased occurrence of cell division | adult stage, with Scer\GAL4esg.PU
long lived | dominant
segment polarity phenotype | germline clone | rescuable maternal effect
some die during embryonic stage | germline clone | maternal effect | maternal effect
some die during pupal stage
some die during pupal stage, with Scer\GAL4da.G32, ttv00681b
some die during pupal stage, with Scer\GAL4da.G32, ttvUAS.Tag:MYC
some die during pupal stage, with Scer\GAL4elav-C155, ttv00681b
some die during pupal stage, with Scer\GAL4elav-C155, ttvUAS.Tag:MYC
some die during pupal stage, with Scer\GAL4G14, ttv00681b
some die during pupal stage, with Scer\GAL4G14, ttvUAS.Tag:MYC
some die during pupal stage, with Scer\GAL4pnr-MD237
some die during pupal stage (with ttv205)
some die during pupal stage (with ttv524)
visible, with Scer\GAL4pnr-MD237
visible | adult stage, with Scer\GAL4Mef2.PR
visible | recessive | somatic clone
wild-type
Phenotype manifest in
Allele
adult head, with Scer\GAL4elav-C155
adult midgut epithelium, with Scer\GAL4esg.PU
anterior fascicle & bouton
bouton
bouton, with Scer\GAL4da.G32, ttv00681b
bouton, with Scer\GAL4da.G32, ttvUAS.Tag:MYC
cuticle
dendritic tree | cell non-autonomous | somatic clone | third instar larval stage, with Scer\GAL4Act5C.PI
dendritic tree | cell non-autonomous | third instar larval stage
dendritic tree | cell non-autonomous | third instar larval stage, with Scer\GAL4A58
dendritic tree | cell non-autonomous | third instar larval stage, with Scer\GAL4GMR10C12
dendritic tree | cell non-autonomous | third instar larval stage, with Scer\GAL4GMR16D01
dendritic tree | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
denticle | ectopic | germline clone | rescuable maternal effect
denticle | germline clone | maternal effect
distal dendrite | cell non-autonomous | somatic clone | increased number | third instar larval stage, with Scer\GAL4Act5C.PI
distal dendrite | cell non-autonomous | third instar larval stage
distal dendrite | cell non-autonomous | third instar larval stage, with Scer\GAL4GMR10C12
distal dendrite | cell non-autonomous | third instar larval stage, with Scer\GAL4GMR16D01
distal dendrite | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
embryonic/first instar larval cuticle | germline clone
embryonic/first instar larval cuticle | germline clone | maternal effect
embryonic/first instar larval cuticle | germline clone | rescuable maternal effect
embryonic/larval salivary gland, with Scer\GAL4fkh.PH
eye | P-stage (with ttv205)
eye | P-stage (with ttv524)
eye | somatic clone
germline cell, with Scer\GAL4elav.PLu
germline cell, with Scer\GAL4ptc-559.1
germline cell, with Scer\GAL4wg.PM
germline cell | non-rescuable maternal effect
imaginal disc | larval stage (with ttv205)
imaginal disc | larval stage (with ttv524)
larval dorsal multidendritic neuron ddaA | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
larval dorsal multidendritic neuron ddaB | cell non-autonomous | third instar larval stage, with Scer\GAL4A58
larval dorsal multidendritic neuron ddaD | cell non-autonomous | third instar larval stage, with Scer\GAL4A58
larval dorsal multidendritic neuron ddaE | cell non-autonomous | third instar larval stage, with Scer\GAL4A58
larval multidendritic class IV neuron | cell non-autonomous | somatic clone | third instar larval stage, with Scer\GAL4Act5C.PI
larval multidendritic class IV neuron | cell non-autonomous | third instar larval stage
larval multidendritic class IV neuron | cell non-autonomous | third instar larval stage, with Scer\GAL4A58
larval multidendritic class IV neuron | cell non-autonomous | third instar larval stage, with Scer\GAL4GMR10C12
larval multidendritic class IV neuron | cell non-autonomous | third instar larval stage, with Scer\GAL4GMR16D01
larval multidendritic class IV neuron | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
leg | P-stage (with ttv205)
leg | P-stage (with ttv524)
macrochaeta
synapse
trichogen cell, with Scer\GAL4pnr-MD237
ventral denticle belt
wing, with Scer\GAL4Mef2.PR
wing | somatic clone
wing disc
wing disc | larval stage (with ttv205)
wing disc | larval stage (with ttv524)
wing margin | somatic clone
wing margin bristle | ectopic | somatic clone
wing vein | ectopic, with Scer\GAL4A9
wing vein | ectopic, with Scer\GAL4Bx-MS1096
wing vein | ectopic, with Scer\GAL4sd-SG29.1
wing vein | somatic clone
wing vein L3 | somatic clone
wing vein L4 | somatic clone
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) (5)
Hsap\EXT1
13 of 14
Yes
Yes
 
2  
Hsap\EXTL1
8 of 14
No
Yes
Hsap\EXT2
3 of 14
No
No
Hsap\EXTL3
3 of 14
No
No
Hsap\EXTL2
2 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) (5)
Rnor\Ext1
13 of 14
Yes
Yes
Rnor\Extl1
8 of 14
No
Yes
Rnor\Ext2
3 of 14
No
No
Rnor\Extl3
3 of 14
No
No
Rnor\Extl2
2 of 14
No
No
Mus musculus (laboratory mouse) (5)
Mmus\Ext1
13 of 14
Yes
Yes
Mmus\Extl1
8 of 14
No
Yes
Mmus\Extl3
3 of 14
No
No
Mmus\Ext2
2 of 14
No
No
Mmus\Extl2
2 of 14
No
No
Xenopus tropicalis (Western clawed frog) (6)
Xtro\ext1
11 of 13
Yes
Yes
Xtro\ext2
2 of 13
No
No
Xtro\extl3
2 of 13
No
No
Xtro\extl2
1 of 13
No
Yes
Xtro\myo7b
1 of 13
No
No
Xtro\skiv2l
1 of 13
No
No
Danio rerio (Zebrafish) (10)
Drer\ext1b
12 of 14
Yes
Yes
Drer\ext1a
11 of 14
No
Yes
Drer\ext1c
11 of 14
No
Yes
Drer\ext2
3 of 14
No
No
Drer\extl3
3 of 14
No
No
Drer\extl2
2 of 14
No
Yes
Drer\LOC108179064
2 of 14
No
Yes
Drer\LOC101885478
1 of 14
No
Yes
Drer\si:dkey-13e3.1
1 of 14
No
Yes
Drer\si:rp71-23d18.4
1 of 14
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (2)
Cele\rib-1
11 of 14
Yes
Yes
Cele\rib-2
3 of 14
No
No
Anopheles gambiae (African malaria mosquito) (3)
Agam\AgaP_AGAP010388
12 of 12
Yes
Yes
Agam\AgaP_AGAP000081
3 of 12
No
No
Agam\AgaP_AGAP001688
3 of 12
No
No
Arabidopsis thaliana (thale-cress) (42)
Atha\AT3G07620
6 of 13
Yes
Yes
Atha\AT3G42180
6 of 13
Yes
Yes
Atha\AT4G16745
6 of 13
Yes
Yes
Atha\AT4G32790
6 of 13
Yes
Yes
Atha\AT4G38040
6 of 13
Yes
Yes
Atha\AT5G03795
6 of 13
Yes
Yes
Atha\AT5G11130
6 of 13
Yes
Yes
Atha\AT5G11610
6 of 13
Yes
Yes
Atha\AT5G19670
6 of 13
Yes
Yes
Atha\AT5G20260
6 of 13
Yes
Yes
Atha\AT5G25310
6 of 13
Yes
Yes
Atha\AT5G25820
6 of 13
Yes
Yes
Atha\AT5G37000
6 of 13
Yes
Yes
Atha\XGD1
6 of 13
Yes
Yes
Atha\F8H
5 of 13
No
Yes
Atha\FRA8
5 of 13
No
Yes
Atha\GUT1
5 of 13
No
Yes
Atha\GUT2
5 of 13
No
Yes
Atha\EPC1
4 of 13
No
No
Atha\ARAD1
3 of 13
No
Yes
Atha\ARAD2
3 of 13
No
Yes
Atha\AT1G21480
3 of 13
No
Yes
Atha\AT1G34270
3 of 13
No
Yes
Atha\AT1G67410
3 of 13
No
Yes
Atha\AT1G68470
3 of 13
No
Yes
Atha\AT1G80290
3 of 13
No
Yes
Atha\AT2G31990
3 of 13
No
Yes
Atha\AT2G32750
3 of 13
No
Yes
Atha\AT4G22580
3 of 13
No
Yes
Atha\AT5G16890
3 of 13
No
Yes
Atha\AT5G41250
3 of 13
No
Yes
Atha\MUR3
3 of 13
No
Yes
Atha\RHS8
3 of 13
No
Yes
Atha\AT1G74680
2 of 13
No
Yes
Atha\AT2G29040
2 of 13
No
Yes
Atha\AT3G45400
2 of 13
No
Yes
Atha\AT3G57630
2 of 13
No
Yes
Atha\AT4G13990
2 of 13
No
Yes
Atha\EDA5
2 of 13
No
Yes
Atha\GT13
2 of 13
No
Yes
Atha\GT18
2 of 13
No
Yes
Atha\AT5G04500
1 of 13
No
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:ttv. 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) (2)
botv
8 of 13
sotv
8 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 1 )
    Allele
    Disease
    Evidence
    References
    ttvRNAi.UAS.cUa
    model of  germ cell cancer
    CEA
    (Levings and Nakato, 2018)
    Potential Models Based on Orthology ( 3 )
    Modifiers Based on Experimental Evidence ( 3 )
    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?
    EXT1; exostosin glycosyltransferase 1
    13 of 14
    608177
     
    EXTL1; exostosin like glycosyltransferase 1
    8 of 14
    601738
        EXT2; exostosin glycosyltransferase 2
        3 of 14
        608210
        EXTL3; exostosin like glycosyltransferase 3
        3 of 14
        605744
          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
          ttv
          Hsap\EXT1
          (Dasgupta et al., 2007)
          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
          ttv
          enhanceable
          β3GalTII
          (Ueyama et al., 2008)
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          hh
          suppressible
          ttv
          (Gallet et al., 2003)
          hh
          enhanceable
          ttv
          (Deshpande et al., 2007)
          Psn
          suppressible
          ttv
          (Reynolds-Peterson et al., 2020)
          Sdc
          enhanceable
          ttv
          (Chanana et al., 2009)
          trol
          enhanceable
          ttv
          (Park et al., 2003)
          β3GalTII
          enhanceable
          ttv
          (Ueyama et al., 2008)
          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
          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)
          2R
          Recombination map
          2-72
          Cytogenetic map
          51A7-51B4
          Sequence location
          FlyBase Computed Cytological Location
          Cytogenetic map
          Evidence for location
          51A7-51B4
          Experimentally Determined Cytological Location
          Cytogenetic map
          Notes
          References
          51B1-51B1
          (Oh et al., 2003)
          51B5-51B5
          (determined by in situ hybridisation)
          (Misra, 2000.8.25)
          51A2-51B2
          (Bellaiche, 1998.8.13)
          51B1-51B5
          (determined by in situ hybridisation)
          (Spradling et al., 1999, Perrimon et al., 1996)
          51A-51A
          (determined by in situ hybridisation)
          (Bellaiche et al., 1998)
          51B1-51B2
          51B7--8
          (BDGP Project Members, 1994-1999)
          Experimentally Determined Recombination Data
          Location

          2-72

          (FlyBase, 2016)

          2-76.0

          (Comeron et al., 2012)

          2-

          (Dickson et al., 1995)
          Left of (cM)
          Right of (cM)
          (Dickson et al., 1995)
          Notes

          36 recombinants in 31,272 progeny.

          (Dickson et al., 1995)
          Stocks and Reagents
          Stocks (39)
          Genomic Clones (46)
          cDNA Clones (61)
           

          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
          Antibody Information
          Laboratory Generated Antibodies
           

          polyclonal

          (Chang et al., 2013, The et al., 1999)
          Commercially Available Antibodies
           
          Cell Line Information
          Publicly Available Cell Lines
           
            Other Stable Cell Lines
             
              Other Comments

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

              (Deshpande et al., 2007)

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

              (Han et al., 2004)

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

              (Takei et al., 2004)

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

              (Oh et al., 2003)

              ttv is involved in heparan sulfate proteoglycan synthesis.

              (The et al., 1999)

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

              (Bellaiche et al., 1998)

              Mutants show maternal effect segment polarity phenotypes.

              (Bellaiche et al., 1998)

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

              (Roch et al., 1998)

              Isolated in a screen for lethal mutations affecting segmentation.

              (Bellaiche et al., 1997)

              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.

              (Perrimon et al., 1996)
              Relationship to Other Genes
              Source for database merge of

              Source for merge of: ttv l(2)00681

              (Bellaiche et al., 1998)

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

              (Beaton, 1999.12.12)

              Source for merge of: ttv l(2)SH0631

              (Gurudatta et al., 2010)
              Additional comments

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

              (Gene Disruption Project members, 2015.10.8)
              Nomenclature History
              Source for database identify of
              Nomenclature comments
              Etymology

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

              (Bellaiche et al., 1998)
              Synonyms and Secondary IDs (22)
              Reported As
              Symbol Synonym
              CG10117
              (National Institute of Genetics Fly Stocks, 2023-, National Institute of Genetics Fly Stocks, 2022-, Meadows, 2017.1.11, Butzlaff et al., 2015, Hosono et al., 2015, Yamamoto-Hino et al., 2015, Gurudatta et al., 2010, Ni et al., 2010.12.1, Izumikawa et al., 2006, Häcker et al., 2005, Norga et al., 2003, Kraut et al., 2001, Kraut et al., 2001, Mount, 2001.8.14)
              DEXT1
              (Nishihara, 2010, Izumikawa et al., 2006, Wilson, 2002)
              EXT1
              (Nishihara, 2020, Nakato and Li, 2016)
              Ext1
              (Wilson and Chuang, 2006)
              P1.15
              (Dickson et al., 1995)
              Ttv
              (Nishihara, 2020, Nagarajan et al., 2015, Yamamoto-Hino et al., 2015, Mbodj et al., 2013, Raftery and Umulis, 2012, Gallet et al., 2006)
              dExt1
              (Yamamoto-Hino et al., 2015)
              l(2)00681
              (Spradling et al., 1999)
              l(2)05282
              (BDGP Project Members, 1994-1999, Berkeley Drosophila Genome Project, 1993.6.1)
              l(2)SH0631
              l(2)SH2 0631
              (Oh et al., 2002.3.24)
              l(2)k00115
              l(2)k03617
              (BDGP Project Members, 1994-1999)
              ttv
              (Patterson et al., 2025, Jiménez-Jiménez et al., 2024, Deshpande et al., 2023, Gude et al., 2023, Aguirre-Tamaral et al., 2022, López-Varea et al., 2021, Parniewska and Stocker, 2020, Wei et al., 2020, Meltzer et al., 2019, Drelon et al., 2018, Gene Disruption Project members, 2018-, Kanai et al., 2018, Levings and Nakato, 2018, Levis, 2018.8.30, Poe et al., 2017, Reynolds-Peterson et al., 2017, Transgenic RNAi Project members, 2017-, Clandinin and Owens, 2016-, Jungreis et al., 2016, Levings et al., 2016, Nakato and Li, 2016, Butzlaff et al., 2015, Gene Disruption Project members, 2015-, Ashwal-Fluss et al., 2014, He et al., 2014, Chang et al., 2013, Deshpande et al., 2013, Gradilla and Guerrero, 2013, Sekine et al., 2013, Uchino et al., 2013, Cho et al., 2012, Rodriguez et al., 2012, Chang et al., 2011, Dejima et al., 2011, Toku et al., 2011, Wang et al., 2011, Gurudatta et al., 2010, Nishihara, 2010, Pospisilik et al., 2010, Baron et al., 2009, Bischoff et al., 2009, Chanana et al., 2009, Hayashi et al., 2009, Maybeck and Röper, 2009, Renault et al., 2009, Ren et al., 2009, Belenkaya et al., 2008, Bornemann et al., 2008, Christensen et al., 2008.12.28, Ueyama et al., 2008, Christensen and Cook, 2007.10.29, Dasgupta et al., 2007, Deshpande et al., 2007, Eugster et al., 2007, Callejo et al., 2006, Gallet et al., 2006, Izumikawa et al., 2006, Wendler et al., 2006, Wilson and Chuang, 2006, Yao et al., 2006, Briscoe and Therond, 2005, Glise et al., 2005, Gorfinkiel et al., 2005, Häcker et al., 2005, Torroja et al., 2005, Stanyon et al., 2004, The et al., 1999)
              Name Synonyms
              Tout velu
              (Gallet et al., 2006)
              Tout-velu
              (Parniewska and Stocker, 2020, Nakato and Li, 2016, Nagarajan et al., 2015)
              lethal (2) SH0631
              tout velour
              (Gradilla and Guerrero, 2013)
              tout velu
              (Dejima et al., 2011, Gurudatta et al., 2010, Bischoff et al., 2009, Maybeck and Röper, 2009, Dasgupta et al., 2007, Häcker et al., 2005, Torroja et al., 2005, Kuwabara and Labouesse, 2002, Mann and Beachy, 2000, The et al., 1998)
              tout-velo
              (He et al., 2014)
              tout-velu
              (Levings et al., 2016, Chanana et al., 2009, Hayashi et al., 2009, Bornemann et al., 2008, Eugster et al., 2007, Izumikawa et al., 2006, Wendler et al., 2006, Wilson and Chuang, 2006, Eldar and Barkai, 2005, Kraut et al., 2001, Kraut et al., 2001, The et al., 1999, Bellaiche, 1998.8.13, Bellaiche et al., 1997)
              toutvelu
              (Deshpande et al., 2007)
              Secondary FlyBase IDs
              • FBgn0065697
              • FBgn0020245
              • FBgn0010469
              • FBgn0010592
              • FBgn0024670
              Datasets (0)
              Study focus (0)
              Experimental Role
              Project
              Project Type
              Title
              Study result (0)
              Result
              Result Type
              Title
              External Crossreferences and Linkouts ( 71 )
              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
              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.
              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
              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.
              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
              FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
              iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
              Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
              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 (211)