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Q9V730

- EXT1_DROME

UniProt

Q9V730 - EXT1_DROME

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Protein

Exostosin-1

Gene

ttv

Organism
Drosophila melanogaster (Fruit fly)
Status
Reviewed - Annotation score: 5 out of 5- Experimental evidence at protein leveli

Functioni

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.9 Publications

Catalytic activityi

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.
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.

Cofactori

Manganese.By similarity

Pathwayi

Sites

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Metal bindingi583 – 5831Manganese; catalyticBy similarity
Binding sitei611 – 6111SubstrateBy similarity
Active sitei670 – 6701By similarity

GO - Molecular functioni

  1. acetylglucosaminyltransferase activity Source: FlyBase
  2. glucuronosyl-N-acetylglucosaminyl-proteoglycan 4-alpha-N-acetylglucosaminyltransferase activity Source: UniProtKB
  3. glucuronosyltransferase activity Source: FlyBase
  4. N-acetylglucosaminyl-proteoglycan 4-beta-glucuronosyltransferase activity Source: UniProtKB

GO - Biological processi

  1. axon guidance Source: FlyBase
  2. decapentaplegic signaling pathway Source: UniProtKB
  3. germ cell migration Source: FlyBase
  4. glycosaminoglycan biosynthetic process Source: UniProtKB
  5. heparan sulfate proteoglycan biosynthetic process Source: UniProtKB
  6. heparan sulfate proteoglycan biosynthetic process, polysaccharide chain biosynthetic process Source: FlyBase
  7. heparin biosynthetic process Source: FlyBase
  8. intracellular distribution of mitochondria Source: FlyBase
  9. N-acetylglucosamine metabolic process Source: UniProtKB
  10. protein glycosylation Source: UniProtKB-UniPathway
  11. regulation of smoothened signaling pathway Source: UniProtKB
  12. smoothened signaling pathway Source: UniProtKB
  13. synapse organization Source: FlyBase
  14. synaptic vesicle endocytosis Source: FlyBase
  15. Wnt signaling pathway Source: UniProtKB
Complete GO annotation...

Keywords - Molecular functioni

Developmental protein, Glycosyltransferase, Transferase

Keywords - Biological processi

Wnt signaling pathway

Keywords - Ligandi

Manganese, Metal-binding

Enzyme and pathway databases

ReactomeiREACT_180263. HS-GAG biosynthesis.
SignaLinkiQ9V730.
UniPathwayiUPA00378.
UPA00756.
UPA00862.

Protein family/group databases

CAZyiGT47. Glycosyltransferase Family 47.
GT64. Glycosyltransferase Family 64.

Names & Taxonomyi

Protein namesi
Recommended name:
Exostosin-1 (EC:2.4.1.224, EC:2.4.1.225)
Alternative name(s):
Protein tout-velu
Gene namesi
Name:ttv
Synonyms:DEXT1
ORF Names:CG10117
OrganismiDrosophila melanogaster (Fruit fly)
Taxonomic identifieri7227 [NCBI]
Taxonomic lineageiEukaryotaMetazoaEcdysozoaArthropodaHexapodaInsectaPterygotaNeopteraEndopterygotaDipteraBrachyceraMuscomorphaEphydroideaDrosophilidaeDrosophilaSophophora
ProteomesiUP000000803: Chromosome 2R

Organism-specific databases

FlyBaseiFBgn0265974. ttv.

Subcellular locationi

GO - Cellular componenti

  1. endoplasmic reticulum Source: UniProtKB
  2. Golgi apparatus Source: UniProtKB
  3. integral component of membrane Source: UniProtKB
  4. intrinsic component of endoplasmic reticulum membrane Source: InterPro
Complete GO annotation...

Keywords - Cellular componenti

Endoplasmic reticulum, Golgi apparatus, Membrane

Pathology & Biotechi

Disruption phenotypei

According to some authors (PubMed:10549295) ttv mutants have no effect on wg signaling, while according to others (PubMed:14645127, PubMed:14729575 and PubMed:15056609) wg signaling is affected. Such discrepancy may be explained by the fact that the absence of ttv could be partially compensated by the intact sotv protein.4 Publications

Mutagenesis

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Mutagenesisi447 – 4471G → D in ttv205; induces defects in wing patterning due to impaired movement of morphogens. 1 Publication

PTM / Processingi

Molecule processing

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Chaini1 – 760760Exostosin-1PRO_0000149664Add
BLAST

Amino acid modifications

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Glycosylationi71 – 711N-linked (GlcNAc...)Sequence Analysis
Glycosylationi327 – 3271N-linked (GlcNAc...)Sequence Analysis
Glycosylationi476 – 4761N-linked (GlcNAc...)Sequence Analysis
Disulfide bondi668 ↔ 716By similarity

Keywords - PTMi

Disulfide bond, Glycoprotein

Proteomic databases

PaxDbiQ9V730.
PRIDEiQ9V730.

Expressioni

Tissue specificityi

Ubiquitously expressed in early embryos. Later (in stage 10 embryos), it is expressed at higher level in the nervous system. Ubiquitously expressed in wing imaginal disk.2 Publications

Developmental stagei

Expressed both maternally and zygotically.1 Publication

Gene expression databases

BgeeiQ9V730.
ExpressionAtlasiQ9V730. differential.

Interactioni

Subunit structurei

Forms a homo/heterooligomeric complex with sotv. May interact with rti.

Binary interactionsi

WithEntry#Exp.IntActNotes
Ext2Q9Y1693EBI-166374,EBI-142791

Protein-protein interaction databases

BioGridi62362. 2 interactions.
IntActiQ9V730. 4 interactions.
MINTiMINT-7953341.

Structurei

3D structure databases

ProteinModelPortaliQ9V730.
SMRiQ9V730. Positions 562-734.
ModBaseiSearch...
MobiDBiSearch...

Topological domain

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Topological domaini1 – 66CytoplasmicSequence Analysis
Topological domaini26 – 760735LumenalSequence AnalysisAdd
BLAST

Transmembrane

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Transmembranei7 – 2519Helical; Signal-anchor for type II membrane proteinSequence AnalysisAdd
BLAST

Family & Domainsi

Region

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Regioni561 – 5655Substrate bindingBy similarity
Regioni581 – 5833Substrate bindingBy similarity
Regioni666 – 6705Substrate bindingBy similarity

Sequence similaritiesi

Belongs to the glycosyltransferase 47 family.Curated

Keywords - Domaini

Signal-anchor, Transmembrane, Transmembrane helix

Phylogenomic databases

eggNOGiNOG272619.
GeneTreeiENSGT00550000074496.
InParanoidiQ9V730.
KOiK02366.
OMAiWEAYFNS.
PhylomeDBiQ9V730.

Family and domain databases

Gene3Di3.90.550.10. 1 hit.
InterProiIPR004263. Exostosin.
IPR015338. HexNAc_Trfase_a.
IPR029044. Nucleotide-diphossugar_trans.
[Graphical view]
PANTHERiPTHR11062. PTHR11062. 1 hit.
PfamiPF03016. Exostosin. 1 hit.
PF09258. Glyco_transf_64. 1 hit.
[Graphical view]
SUPFAMiSSF53448. SSF53448. 1 hit.

Sequencei

Sequence statusi: Complete.

Q9V730-1 [UniParc]FASTAAdd to Basket

« Hide

        10         20         30         40         50
MQAKKRYILV FVSCAFLAYA YFGGYRLKVS PLRPRRAQHE SAKDGGVQPH
60 70 80 90 100
EQLPSFLGAH DMQELQLLQS NQSKSLDSSK HLVTRKPDCR METCFDFTRC
110 120 130 140 150
YDRFLVYIYP PEPLNSLGAA PPTSANYQKI LTAIQESRYY TSDPTAACLF
160 170 180 190 200
VLGIDTLDRD SLSEDYVRNV PSRLARLPYW NNGRNHIIFN LYSGTWPDYA
210 220 230 240 250
ENSLGFDAGE AILAKASMGV LQLRHGFDVS IPLFHKQFPL RAGATGTVQS
260 270 280 290 300
NNFPANKKYL LAFKGKRYVH GIGSETRNSL FHLHNGRDMV LVTTCRHGKS
310 320 330 340 350
WRELQDNRCD EDNREYDRYD YETLLQNSTF CLVPRGRRLG SFRFLEALQA
360 370 380 390 400
GCIPVLLSNA WVLPFESKID WKQAAIWADE RLLLQVPDIV RSIPAERIFA
410 420 430 440 450
LRQQTQVLWE RYFGSIEKIV FTTFEIIRER LPDYPVRSSL VWNSSPGALL
460 470 480 490 500
TLPTFADSSR YMPFLLNSMG AEPRHNYTAV IYVQIGAALG PNAALYKLVR
510 520 530 540 550
TITKSQFVER ILVLWAADRP LPLKKRWPPT SHIPLHVISL GGSTRSQGAG
560 570 580 590 600
PTSQTTEGRP SISQRFLPYD EIQTDAVLSL DEDAILNTDE LDFAYTVWRD
610 620 630 640 650
FPERIVGYPA RAHFWDDSKN AWGYTSKWTN YYSIVLTGAA FYHRYYNYLY
660 670 680 690 700
TNWLSLLLLK TVQQSSNCED ILMNLLVSHV TRKPPIKVTQ RKGYKDRETG
710 720 730 740 750
RSPWNDPDHF IQRQSCLNTF AAVFGYMPLI RSNLRMDPML YRDPVSNLRK
760
KYRQIELVGS
Length:760
Mass (Da):87,309
Last modified:May 1, 2000 - v1
Checksum:iF6369F0BA206DEA9
GO

Sequence cautioni

The sequence AAC32397.1 differs from that shown. Reason: Frameshift at positions 694 and 702.

Sequence databases

Select the link destinations:
EMBL
GenBank
DDBJ
Links Updated
AF083889 mRNA. Translation: AAC32397.1. Frameshift.
AB221351 mRNA. Translation: BAE78509.1.
AE013599 Genomic DNA. Translation: AAF58236.1.
BT021403 mRNA. Translation: AAX33551.1.
RefSeqiNP_001260971.1. NM_001274042.1.
NP_477231.1. NM_057883.3.
UniGeneiDm.2966.

Genome annotation databases

EnsemblMetazoaiFBtr0087495; FBpp0086624; FBgn0020245.
FBtr0333164; FBpp0305367; FBgn0020245.
GeneIDi36614.
KEGGidme:Dmel_CG10117.

Cross-referencesi

Sequence databases

Select the link destinations:
EMBL
GenBank
DDBJ
Links Updated
AF083889 mRNA. Translation: AAC32397.1 . Frameshift.
AB221351 mRNA. Translation: BAE78509.1 .
AE013599 Genomic DNA. Translation: AAF58236.1 .
BT021403 mRNA. Translation: AAX33551.1 .
RefSeqi NP_001260971.1. NM_001274042.1.
NP_477231.1. NM_057883.3.
UniGenei Dm.2966.

3D structure databases

ProteinModelPortali Q9V730.
SMRi Q9V730. Positions 562-734.
ModBasei Search...
MobiDBi Search...

Protein-protein interaction databases

BioGridi 62362. 2 interactions.
IntActi Q9V730. 4 interactions.
MINTi MINT-7953341.

Protein family/group databases

CAZyi GT47. Glycosyltransferase Family 47.
GT64. Glycosyltransferase Family 64.

Proteomic databases

PaxDbi Q9V730.
PRIDEi Q9V730.

Protocols and materials databases

Structural Biology Knowledgebase Search...

Genome annotation databases

EnsemblMetazoai FBtr0087495 ; FBpp0086624 ; FBgn0020245 .
FBtr0333164 ; FBpp0305367 ; FBgn0020245 .
GeneIDi 36614.
KEGGi dme:Dmel_CG10117.

Organism-specific databases

CTDi 36614.
FlyBasei FBgn0265974. ttv.

Phylogenomic databases

eggNOGi NOG272619.
GeneTreei ENSGT00550000074496.
InParanoidi Q9V730.
KOi K02366.
OMAi WEAYFNS.
PhylomeDBi Q9V730.

Enzyme and pathway databases

UniPathwayi UPA00378 .
UPA00756 .
UPA00862 .
Reactomei REACT_180263. HS-GAG biosynthesis.
SignaLinki Q9V730.

Miscellaneous databases

ChiTaRSi EXT1. drosophila.
GenomeRNAii 36614.
NextBioi 799500.
PROi Q9V730.

Gene expression databases

Bgeei Q9V730.
ExpressionAtlasi Q9V730. differential.

Family and domain databases

Gene3Di 3.90.550.10. 1 hit.
InterProi IPR004263. Exostosin.
IPR015338. HexNAc_Trfase_a.
IPR029044. Nucleotide-diphossugar_trans.
[Graphical view ]
PANTHERi PTHR11062. PTHR11062. 1 hit.
Pfami PF03016. Exostosin. 1 hit.
PF09258. Glyco_transf_64. 1 hit.
[Graphical view ]
SUPFAMi SSF53448. SSF53448. 1 hit.
ProtoNeti Search...

Publicationsi

« Hide 'large scale' publications
  1. "Tout-velu is a Drosophila homologue of the putative tumour suppressor EXT-1 and is needed for Hh diffusion."
    Bellaiche Y., The I., Perrimon N.
    Nature 394:85-88(1998) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [MRNA], FUNCTION, MEMBRANE TOPOLOGY, DEVELOPMENTAL STAGE.
  2. Cited for: NUCLEOTIDE SEQUENCE [MRNA], FUNCTION, INTERACTION WITH SOTV.
  3. "The genome sequence of Drosophila melanogaster."
    Adams M.D., Celniker S.E., Holt R.A., Evans C.A., Gocayne J.D., Amanatides P.G., Scherer S.E., Li P.W., Hoskins R.A., Galle R.F., George R.A., Lewis S.E., Richards S., Ashburner M., Henderson S.N., Sutton G.G., Wortman J.R., Yandell M.D.
    , Zhang Q., Chen L.X., Brandon R.C., Rogers Y.-H.C., Blazej R.G., Champe M., Pfeiffer B.D., Wan K.H., Doyle C., Baxter E.G., Helt G., Nelson C.R., Miklos G.L.G., Abril J.F., Agbayani A., An H.-J., Andrews-Pfannkoch C., Baldwin D., Ballew R.M., Basu A., Baxendale J., Bayraktaroglu L., Beasley E.M., Beeson K.Y., Benos P.V., Berman B.P., Bhandari D., Bolshakov S., Borkova D., Botchan M.R., Bouck J., Brokstein P., Brottier P., Burtis K.C., Busam D.A., Butler H., Cadieu E., Center A., Chandra I., Cherry J.M., Cawley S., Dahlke C., Davenport L.B., Davies P., de Pablos B., Delcher A., Deng Z., Mays A.D., Dew I., Dietz S.M., Dodson K., Doup L.E., Downes M., Dugan-Rocha S., Dunkov B.C., Dunn P., Durbin K.J., Evangelista C.C., Ferraz C., Ferriera S., Fleischmann W., Fosler C., Gabrielian A.E., Garg N.S., Gelbart W.M., Glasser K., Glodek A., Gong F., Gorrell J.H., Gu Z., Guan P., Harris M., Harris N.L., Harvey D.A., Heiman T.J., Hernandez J.R., Houck J., Hostin D., Houston K.A., Howland T.J., Wei M.-H., Ibegwam C., Jalali M., Kalush F., Karpen G.H., Ke Z., Kennison J.A., Ketchum K.A., Kimmel B.E., Kodira C.D., Kraft C.L., Kravitz S., Kulp D., Lai Z., Lasko P., Lei Y., Levitsky A.A., Li J.H., Li Z., Liang Y., Lin X., Liu X., Mattei B., McIntosh T.C., McLeod M.P., McPherson D., Merkulov G., Milshina N.V., Mobarry C., Morris J., Moshrefi A., Mount S.M., Moy M., Murphy B., Murphy L., Muzny D.M., Nelson D.L., Nelson D.R., Nelson K.A., Nixon K., Nusskern D.R., Pacleb J.M., Palazzolo M., Pittman G.S., Pan S., Pollard J., Puri V., Reese M.G., Reinert K., Remington K., Saunders R.D.C., Scheeler F., Shen H., Shue B.C., Siden-Kiamos I., Simpson M., Skupski M.P., Smith T.J., Spier E., Spradling A.C., Stapleton M., Strong R., Sun E., Svirskas R., Tector C., Turner R., Venter E., Wang A.H., Wang X., Wang Z.-Y., Wassarman D.A., Weinstock G.M., Weissenbach J., Williams S.M., Woodage T., Worley K.C., Wu D., Yang S., Yao Q.A., Ye J., Yeh R.-F., Zaveri J.S., Zhan M., Zhang G., Zhao Q., Zheng L., Zheng X.H., Zhong F.N., Zhong W., Zhou X., Zhu S.C., Zhu X., Smith H.O., Gibbs R.A., Myers E.W., Rubin G.M., Venter J.C.
    Science 287:2185-2195(2000) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
    Strain: Berkeley.
  4. Cited for: GENOME REANNOTATION.
    Strain: Berkeley.
  5. Stapleton M., Carlson J.W., Chavez C., Frise E., George R.A., Pacleb J.M., Park S., Wan K.H., Yu C., Rubin G.M., Celniker S.E.
    Submitted (MAR-2005) to the EMBL/GenBank/DDBJ databases
    Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
    Strain: Berkeley.
    Tissue: Embryo.
  6. "Hedgehog movement is regulated through tout velu-dependent synthesis of a heparan sulfate proteoglycan."
    The I., Bellaiche Y., Perrimon N.
    Mol. Cell 4:633-639(1999) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION IN HH PATHWAY, ENZYME ACTIVITY, SUBCELLULAR LOCATION, TISSUE SPECIFICITY, DISRUPTION PHENOTYPE.
  7. "Structural analysis of glycosaminoglycans in Drosophila and Caenorhabditis elegans and demonstration that tout-velu, a Drosophila gene related to EXT tumor suppressors, affects heparan sulfate in vivo."
    Toyoda H., Kinoshita-Toyoda A., Selleck S.B.
    J. Biol. Chem. 275:2269-2275(2000) [PubMed] [Europe PMC] [Abstract]
    Cited for: ENZYME ACTIVITY.
  8. "Structural analysis of glycosaminoglycans in animals bearing mutations in sugarless, sulfateless, and tout-velu. Drosophila homologues of vertebrate genes encoding glycosaminoglycan biosynthetic enzymes."
    Toyoda H., Kinoshita-Toyoda A., Fox B., Selleck S.B.
    J. Biol. Chem. 275:21856-21861(2000) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION IN HH PATHWAY.
  9. "Cholesterol modification of hedgehog is required for trafficking and movement, revealing an asymmetric cellular response to hedgehog."
    Gallet A., Rodriguez R., Ruel L., Therond P.P.
    Dev. Cell 4:191-204(2003) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION IN HH PATHWAY.
  10. "Three Drosophila EXT genes shape morphogen gradients through synthesis of heparan sulfate proteoglycans."
    Takei Y., Ozawa Y., Sato M., Watanabe A., Tabata T.
    Development 131:73-82(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, TISSUE SPECIFICITY, DISRUPTION PHENOTYPE, MUTAGENESIS OF GLY-447.
  11. "Drosophila glypicans control the cell-to-cell movement of Hedgehog by a dynamin-independent process."
    Han C., Belenkaya T.Y., Wang B., Lin X.
    Development 131:601-611(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION IN HH PATHWAY, DISRUPTION PHENOTYPE.
  12. "Distinct and collaborative roles of Drosophila EXT family proteins in morphogen signalling and gradient formation."
    Han C., Belenkaya T.Y., Khodoun M., Tauchi M., Lin X., Lin X.
    Development 131:1563-1575(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION.
  13. "Abrogation of heparan sulfate synthesis in Drosophila disrupts the Wingless, Hedgehog and Decapentaplegic signaling pathways."
    Bornemann D.J., Duncan J.E., Staatz W., Selleck S., Warrior R.
    Development 131:1927-1938(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE.
  14. "The Drosophila GOLPH3 homolog regulates the biosynthesis of heparan sulfate proteoglycans by modulating the retrograde trafficking of exostosins."
    Chang W.L., Chang C.W., Chang Y.Y., Sung H.H., Lin M.D., Chang S.C., Chen C.H., Huang C.W., Tung K.S., Chou T.B.
    Development 140:2798-2807(2013) [PubMed] [Europe PMC] [Abstract]
    Cited for: INTERACTION WITH RTI, SUBCELLULAR LOCATION.

Entry informationi

Entry nameiEXT1_DROME
AccessioniPrimary (citable) accession number: Q9V730
Secondary accession number(s): O76796, Q5BI21
Entry historyi
Integrated into UniProtKB/Swiss-Prot: March 15, 2005
Last sequence update: May 1, 2000
Last modified: October 29, 2014
This is version 98 of the entry and version 1 of the sequence. [Complete history]
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programDrosophila annotation project

Miscellaneousi

Miscellaneous

'Tout velu' means 'very hairy' in French.

Keywords - Technical termi

Complete proteome, Reference proteome

Documents

  1. Drosophila
    Drosophila: entries, gene names and cross-references to FlyBase
  2. PATHWAY comments
    Index of metabolic and biosynthesis pathways
  3. SIMILARITY comments
    Index of protein domains and families

External Data

Dasty 3