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Submissions

heparanase

Summary

Gene Symbol

HPSE

Aliases

HPA
HPSE1
HSE1

Organism

Homo sapiens (human)

External Links

NCBI Gene

10855

HGNC

5164

KEGG Gene ID

hsa:10855

PubChem

10855

Alliance of Genome Resources

HGNC:5164

Annotation

Keyword

3D-structure
Alternative splicing
Calcium
Cell adhesion
Direct protein sequencing
Disulfide bond
Glycoprotein
Hydrolase
Lysosome
Magnesium
Membrane
Nucleus
Reference proteome
Secreted
Signal

Proteins

Displaying 1 entry
UniProt	Protein Name
Q9Y251	Endo-glucoronidase Heparanase-1

Annotation information from UniProt.

Gene Ontology (GO)

Displaying entries **1 - 5** of 15 in total

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GO Term	Evidence Code	PMID
protein transmembrane transport	IEA
response to organic substance	IEA
positive regulation of blood coagulation	IDA	20634491
establishment of endothelial barrier	IEA
positive regulation of vascular endothelial growth factor production	IDA	16452201

GO Hierarchy

biological process

metabolic process [inference]

nitrogen compound metabolic process [inference]

organonitrogen compound metabolic process [inference]

aminoglycan metabolic process [inference]

glycosaminoglycan metabolic process [inference]

mucopolysaccharide metabolic process [inference]

heparin metabolic process

protein metabolic process [inference]

glycoprotein metabolic process [inference]

proteoglycan metabolic process

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

organonitrogen compound catabolic process [inference]

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

catabolic process [inference]

cellular catabolic process [inference]

sulfur compound catabolic process [inference]

heparan sulfate proteoglycan catabolic process

organic substance catabolic process [inference]

macromolecule catabolic process [inference]

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

carbohydrate derivative catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

organonitrogen compound catabolic process [inference]

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

cellular metabolic process [inference]

sulfur compound metabolic process [inference]

heparin metabolic process

sulfur compound catabolic process [inference]

heparan sulfate proteoglycan catabolic process

cellular catabolic process [inference]

sulfur compound catabolic process [inference]

heparan sulfate proteoglycan catabolic process

primary metabolic process [inference]

protein metabolic process [inference]

glycoprotein metabolic process [inference]

proteoglycan metabolic process

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

organic substance metabolic process [inference]

macromolecule metabolic process [inference]

aminoglycan metabolic process [inference]

glycosaminoglycan metabolic process [inference]

mucopolysaccharide metabolic process [inference]

heparin metabolic process

macromolecule catabolic process [inference]

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

protein metabolic process [inference]

glycoprotein metabolic process [inference]

proteoglycan metabolic process

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

carbohydrate derivative metabolic process [inference]

aminoglycan metabolic process [inference]

glycosaminoglycan metabolic process [inference]

mucopolysaccharide metabolic process [inference]

heparin metabolic process

glycoprotein metabolic process [inference]

proteoglycan metabolic process

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

carbohydrate derivative catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

organonitrogen compound metabolic process [inference]

aminoglycan metabolic process [inference]

glycosaminoglycan metabolic process [inference]

mucopolysaccharide metabolic process [inference]

heparin metabolic process

protein metabolic process [inference]

glycoprotein metabolic process [inference]

proteoglycan metabolic process

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

organonitrogen compound catabolic process [inference]

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

organic substance catabolic process [inference]

macromolecule catabolic process [inference]

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

carbohydrate derivative catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

organonitrogen compound catabolic process [inference]

protein catabolic process [inference]

glycoprotein catabolic process [inference]

proteoglycan catabolic process [inference]

heparan sulfate proteoglycan catabolic process

cellular process [inference]

cell adhesion [inference]

cell-substrate adhesion [inference]

cell-matrix adhesion

cellular metabolic process [inference]

sulfur compound metabolic process [inference]

heparin metabolic process

sulfur compound catabolic process [inference]

heparan sulfate proteoglycan catabolic process

cellular catabolic process [inference]

sulfur compound catabolic process [inference]

heparan sulfate proteoglycan catabolic process

cellular developmental process [inference]

cell development [inference]

epithelial cell development [inference]

endothelial cell development [inference]

establishment of endothelial barrier

cellular localization [inference]

cellular macromolecule localization [inference]

protein localization [inference]

establishment of protein localization [inference]

protein transport [inference]

protein transmembrane transport

transmembrane transport [inference]

protein transmembrane transport

developmental process [inference]

anatomical structure formation involved in morphogenesis [inference]

angiogenesis [inference]

angiogenesis involved in wound healing

vascular wound healing

anatomical structure development [inference]

cell development [inference]

epithelial cell development [inference]

endothelial cell development [inference]

establishment of endothelial barrier

cellular developmental process [inference]

cell development [inference]

epithelial cell development [inference]

endothelial cell development [inference]

establishment of endothelial barrier

response to stimulus [inference]

response to chemical [inference]

response to organic substance

localization [inference]

macromolecule localization [inference]

cellular macromolecule localization [inference]

protein localization [inference]

establishment of protein localization [inference]

protein transport [inference]

protein transmembrane transport

establishment of localization [inference]

transport [inference]

transmembrane transport [inference]

protein transmembrane transport

organic substance transport [inference]

protein transport [inference]

protein transmembrane transport

nitrogen compound transport [inference]

protein transport [inference]

protein transmembrane transport

establishment of protein localization [inference]

protein transport [inference]

protein transmembrane transport

cellular localization [inference]

cellular macromolecule localization [inference]

protein localization [inference]

establishment of protein localization [inference]

protein transport [inference]

protein transmembrane transport

biological regulation [inference]

regulation of biological process [inference]

regulation of metabolic process [inference]

regulation of biosynthetic process [inference]

positive regulation of biosynthetic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of cellular biosynthetic process [inference]

regulation of macromolecule biosynthetic process [inference]

regulation of gene expression [inference]

regulation of cytokine production [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of vascular endothelial growth factor production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of metabolic process [inference]

positive regulation of biosynthetic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of macromolecule metabolic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of cellular metabolic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of cellular metabolic process [inference]

positive regulation of cellular metabolic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of cellular biosynthetic process [inference]

regulation of macromolecule biosynthetic process [inference]

regulation of gene expression [inference]

regulation of cytokine production [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of vascular endothelial growth factor production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of macromolecule metabolic process [inference]

regulation of macromolecule biosynthetic process [inference]

regulation of gene expression [inference]

regulation of cytokine production [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of vascular endothelial growth factor production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of macromolecule metabolic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of signaling [inference]

regulation of signal transduction [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

regulation of intracellular signal transduction [inference]

regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of signaling [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of biological process [inference]

positive regulation of metabolic process [inference]

positive regulation of biosynthetic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of macromolecule metabolic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of cellular metabolic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of signaling [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of cellular process [inference]

positive regulation of cell population proliferation [inference]

positive regulation of osteoblast proliferation

positive regulation of cell communication [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of cellular metabolic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of response to stimulus [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of response to wounding [inference]

positive regulation of wound healing [inference]

positive regulation of blood coagulation

positive regulation of developmental process [inference]

positive regulation of hair follicle development

positive regulation of multicellular organismal process [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of coagulation [inference]

positive regulation of blood coagulation

positive regulation of hair follicle development

positive regulation of hemostasis [inference]

positive regulation of blood coagulation

regulation of response to stimulus [inference]

regulation of signal transduction [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

regulation of intracellular signal transduction [inference]

regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

regulation of response to external stimulus [inference]

regulation of blood coagulation [inference]

positive regulation of blood coagulation

positive regulation of response to stimulus [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of response to wounding [inference]

positive regulation of wound healing [inference]

positive regulation of blood coagulation

regulation of response to stress [inference]

regulation of response to wounding [inference]

regulation of wound healing [inference]

regulation of blood coagulation [inference]

positive regulation of blood coagulation

positive regulation of wound healing [inference]

positive regulation of blood coagulation

positive regulation of response to wounding [inference]

positive regulation of wound healing [inference]

positive regulation of blood coagulation

regulation of developmental process [inference]

positive regulation of developmental process [inference]

positive regulation of hair follicle development

regulation of multicellular organismal development [inference]

regulation of hair follicle development

positive regulation of hair follicle development

regulation of cellular process [inference]

regulation of cell communication [inference]

regulation of signal transduction [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

regulation of intracellular signal transduction [inference]

regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of cell communication [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

regulation of cellular metabolic process [inference]

positive regulation of cellular metabolic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of cellular biosynthetic process [inference]

regulation of macromolecule biosynthetic process [inference]

regulation of gene expression [inference]

regulation of cytokine production [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of vascular endothelial growth factor production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of cell population proliferation [inference]

positive regulation of cell population proliferation [inference]

positive regulation of osteoblast proliferation

regulation of osteoblast proliferation [inference]

positive regulation of osteoblast proliferation

positive regulation of cellular process [inference]

positive regulation of cell population proliferation [inference]

positive regulation of osteoblast proliferation

positive regulation of cell communication [inference]

positive regulation of signal transduction [inference]

positive regulation of intracellular signal transduction [inference]

positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction

positive regulation of cellular metabolic process [inference]

positive regulation of cellular biosynthetic process [inference]

positive regulation of macromolecule biosynthetic process [inference]

positive regulation of gene expression [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of multicellular organismal process [inference]

regulation of cytokine production [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

regulation of vascular endothelial growth factor production [inference]

positive regulation of vascular endothelial growth factor production

regulation of hair cycle [inference]

regulation of hair follicle development

positive regulation of hair follicle development

regulation of coagulation [inference]

regulation of blood coagulation [inference]

positive regulation of blood coagulation

positive regulation of coagulation [inference]

positive regulation of blood coagulation

positive regulation of multicellular organismal process [inference]

positive regulation of cytokine production [inference]

positive regulation of vascular endothelial growth factor production

positive regulation of coagulation [inference]

positive regulation of blood coagulation

positive regulation of hair follicle development

regulation of multicellular organismal development [inference]

regulation of hair follicle development

positive regulation of hair follicle development

regulation of hemostasis [inference]

regulation of blood coagulation [inference]

positive regulation of blood coagulation

positive regulation of hemostasis [inference]

positive regulation of blood coagulation

regulation of biological quality [inference]

regulation of body fluid levels [inference]

regulation of hemostasis [inference]

regulation of blood coagulation [inference]

positive regulation of blood coagulation

positive regulation of hemostasis [inference]

positive regulation of blood coagulation

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GO Term	Evidence Code	PMID
nucleus	IDA	20309870
extracellular matrix	IBA
intracellular membrane-bounded organelle	IDA
extracellular region	TAS
extracellular space	IBA

GO Hierarchy

cellular component

cellular anatomical entity [inference]

membrane [inference]

organelle membrane [inference]

bounding membrane of organelle [inference]

vacuolar membrane [inference]

lytic vacuole membrane [inference]

lysosomal membrane

membrane microdomain [inference]

membrane raft

external encapsulating structure [inference]

extracellular matrix

membrane-enclosed lumen [inference]

organelle lumen [inference]

vesicle lumen [inference]

cytoplasmic vesicle lumen [inference]

secretory granule lumen [inference]

specific granule lumen

intracellular organelle lumen [inference]

vacuolar lumen [inference]

lysosomal lumen

cytoplasmic vesicle lumen [inference]

secretory granule lumen [inference]

specific granule lumen

organelle [inference]

membrane-bounded organelle [inference]

intracellular membrane-bounded organelle

nucleus

vacuole [inference]

lytic vacuole [inference]

lysosome

intracellular organelle [inference]

intracellular membrane-bounded organelle

nucleus

vacuole [inference]

lytic vacuole [inference]

lysosome

Displaying **all 4** entries
GO Term	Evidence Code	PMID
syndecan binding	IDA	15292202
beta-glucuronidase activity	TAS	10395325
heparanase activity	IDA	12213822
protein binding	IPI	15292202 20561914 20634491

GO Hierarchy

molecular function

catalytic activity [inference]

hydrolase activity [inference]

hydrolase activity, acting on glycosyl bonds [inference]

hydrolase activity, hydrolyzing O-glycosyl compounds [inference]

binding [inference]

protein binding

proteoglycan binding [inference]

syndecan binding

carbohydrate derivative binding [inference]

proteoglycan binding [inference]

syndecan binding

Gene Ontology information from NCBI Gene and UniProt.

KEGG BRITE Database

Orthology

K07964

Name

heparanase [EC:3.2.1.166]

References

10405343

Disease

Disease Ontology

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DO ID	Disease Name	Source
DOID:0050387	nonpapillary renal cell carcinoma	DisGeNET
DOID:0050535	exudative vitreoretinopathy	DisGeNET
DOID:0050540	Charcot-Marie-Tooth disease type 3	DisGeNET
DOID:0050589	inflammatory bowel disease	DisGeNET DisGeNET
DOID:0050700	cardiomyopathy	DisGeNET
DOID:0050771	pheochromocytoma	DisGeNET
DOID:0050816	urofacial syndrome	DisGeNET
DOID:0050865	tongue squamous cell carcinoma	DisGeNET
DOID:0050894	ameloblastoma	DisGeNET
DOID:0050902	medulloblastoma	DisGeNET DisGeNET DisGeNET

Ortholog

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Species	Gene ID	Alliance of Genome Resources	Orthologous MAtrix
Drosophila melanogaster	42212	FB:FBgn0038611
Callorhinchus milii	103187960		CALMI37927
Latimeria chalumnae	102346905		LATCH11094
Danio rerio	563020	ZFIN:ZDB-GENE-041111-200	DANRE46327
Ictalurus punctatus	108278991		ICTPU11875
Electrophorus electricus	113570752		ELEEL08258
Salmo salar	106563622		SALSA12204
Salmo salar	106568158		SALSA68820
Salmo trutta	115149247		SALTR20017
Salmo trutta	115159329		SALTR70687

GlyTouCan

GlyComb

GlycoPOST

UniCarb-DR

All Resources

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Tools

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Summary

External Links

Annotation

KEGG BRITE Database

International Collaboration

Acknowledgements