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hyaluronidase 1

Summary

Gene Symbol

HYAL1

Aliases

HYAL-1
LUCA1

Organism

Homo sapiens (human)

External Links

NCBI Gene

3373

HGNC

5320

KEGG Gene ID

hsa:3373

PubChem

3373

Alliance of Genome Resources

HGNC:5320

Annotation

Keyword

3D-structure
Alternative splicing
Direct protein sequencing
Disease variant
Disulfide bond
EGF-like domain
Glycoprotein
Glycosidase
Lysosome
Mucopolysaccharidosis
Reference proteome
Secreted
Signal

Proteins

Displaying **all 4** entries
UniProt	Protein Name
A0A024R2X3
A0A0S2Z3Q0
B3KUI5
Q12794	Hyaluronoglucosaminidase-1 Lung carcinoma protein 1

Annotation information from UniProt.

Gene Ontology (GO)

Displaying entries **1 - 5** of 27 in total

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GO Term	Evidence Code	PMID
positive regulation of cell adhesion	IMP	20473947
glycosaminoglycan catabolic process	TAS
hyaluronan catabolic process	IBA IDA TAS	11296287 17170110 20473947 21695196 21699545
cellular response to fibroblast growth factor stimulus	IDA	19577615
carbohydrate metabolic process	IEA

GO Hierarchy

biological process

metabolic process [inference]

nitrogen compound metabolic process [inference]

organonitrogen compound metabolic process [inference]

aminoglycan metabolic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

glycosaminoglycan metabolic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

glycosaminoglycan catabolic process

mucopolysaccharide metabolic process [inference]

chondroitin sulfate metabolic process [inference]

chondroitin sulfate catabolic process

hyaluronan metabolic process

organonitrogen compound catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

organonitrogen compound biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

catabolic process [inference]

cellular catabolic process [inference]

sulfur compound catabolic process [inference]

chondroitin sulfate catabolic process

organic substance catabolic process [inference]

macromolecule catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

carbohydrate derivative catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

organonitrogen compound catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

biosynthetic process [inference]

cellular biosynthetic process [inference]

macromolecule biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

organic substance biosynthetic process [inference]

macromolecule biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

carbohydrate derivative biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

organonitrogen compound biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

cellular metabolic process [inference]

sulfur compound metabolic process [inference]

chondroitin sulfate metabolic process [inference]

chondroitin sulfate catabolic process

sulfur compound catabolic process [inference]

chondroitin sulfate catabolic process

cellular catabolic process [inference]

sulfur compound catabolic process [inference]

chondroitin sulfate catabolic process

cellular biosynthetic process [inference]

macromolecule biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

primary metabolic process [inference]

carbohydrate metabolic process

organic substance metabolic process [inference]

carbohydrate metabolic process

macromolecule metabolic process [inference]

aminoglycan metabolic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

glycosaminoglycan metabolic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

glycosaminoglycan catabolic process

mucopolysaccharide metabolic process [inference]

chondroitin sulfate metabolic process [inference]

chondroitin sulfate catabolic process

hyaluronan metabolic process

macromolecule catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

macromolecule biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

carbohydrate derivative metabolic process [inference]

aminoglycan metabolic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

glycosaminoglycan metabolic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

glycosaminoglycan catabolic process

mucopolysaccharide metabolic process [inference]

chondroitin sulfate metabolic process [inference]

chondroitin sulfate catabolic process

hyaluronan metabolic process

carbohydrate derivative catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

carbohydrate derivative biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

organonitrogen compound metabolic process [inference]

aminoglycan metabolic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

glycosaminoglycan metabolic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

glycosaminoglycan catabolic process

mucopolysaccharide metabolic process [inference]

chondroitin sulfate metabolic process [inference]

chondroitin sulfate catabolic process

hyaluronan metabolic process

organonitrogen compound catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

organonitrogen compound biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

organic substance catabolic process [inference]

macromolecule catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

carbohydrate derivative catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

organonitrogen compound catabolic process [inference]

aminoglycan catabolic process [inference]

glycosaminoglycan catabolic process

organic substance biosynthetic process [inference]

macromolecule biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

carbohydrate derivative biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

organonitrogen compound biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

cellular process [inference]

cellular metabolic process [inference]

sulfur compound metabolic process [inference]

chondroitin sulfate metabolic process [inference]

chondroitin sulfate catabolic process

sulfur compound catabolic process [inference]

chondroitin sulfate catabolic process

cellular catabolic process [inference]

sulfur compound catabolic process [inference]

chondroitin sulfate catabolic process

cellular biosynthetic process [inference]

macromolecule biosynthetic process [inference]

aminoglycan biosynthetic process [inference]

glycosaminoglycan biosynthetic process [inference]

hyaluronan biosynthetic process

cellular response to stimulus [inference]

cellular response to chemical stimulus [inference]

cellular response to organic substance [inference]

cellular response to cytokine stimulus [inference]

cellular response to environmental stimulus [inference]

cellular response to abiotic stimulus [inference]

cellular response to pH

cellular response to radiation [inference]

cellular response to light stimulus [inference]

cellular response to UV [inference]

cellular response to UV-B

developmental process [inference]

anatomical structure morphogenesis [inference]

animal organ morphogenesis [inference]

embryonic organ morphogenesis [inference]

embryonic skeletal system morphogenesis [inference]

embryonic skeletal joint morphogenesis

skeletal system morphogenesis [inference]

embryonic skeletal system morphogenesis [inference]

embryonic skeletal joint morphogenesis

embryonic morphogenesis [inference]

embryonic organ morphogenesis [inference]

embryonic skeletal system morphogenesis [inference]

embryonic skeletal joint morphogenesis

anatomical structure development [inference]

animal organ development [inference]

cartilage development

biological process involved in interspecies interaction between organisms [inference]

response to other organism [inference]

response to virus

response to stimulus [inference]

response to stress [inference]

defense response [inference]

inflammatory response

response to oxidative stress [inference]

response to reactive oxygen species

response to external stimulus [inference]

response to external biotic stimulus [inference]

response to other organism [inference]

response to virus

response to biotic stimulus [inference]

response to external biotic stimulus [inference]

response to other organism [inference]

response to virus

response to abiotic stimulus [inference]

response to pH [inference]

cellular response to pH

response to radiation [inference]

response to light stimulus [inference]

response to UV [inference]

response to UV-B [inference]

cellular response to UV-B

cellular response to UV [inference]

cellular response to UV-B

cellular response to light stimulus [inference]

cellular response to UV [inference]

cellular response to UV-B

cellular response to radiation [inference]

cellular response to light stimulus [inference]

cellular response to UV [inference]

cellular response to UV-B

cellular response to abiotic stimulus [inference]

cellular response to pH

cellular response to radiation [inference]

cellular response to light stimulus [inference]

cellular response to UV [inference]

cellular response to UV-B

response to endogenous stimulus [inference]

response to growth factor [inference]

response to platelet-derived growth factor [inference]

cellular response to platelet-derived growth factor stimulus

cellular response to growth factor stimulus [inference]

response to fibroblast growth factor [inference]

cellular response to fibroblast growth factor stimulus

cellular response to endogenous stimulus [inference]

cellular response to growth factor stimulus [inference]

response to chemical [inference]

response to organic substance [inference]

response to cytokine [inference]

response to tumor necrosis factor [inference]

cellular response to tumor necrosis factor

response to interleukin-1 [inference]

cellular response to interleukin-1

cellular response to cytokine stimulus [inference]

cellular response to organic substance [inference]

cellular response to cytokine stimulus [inference]

response to antibiotic

cellular response to chemical stimulus [inference]

cellular response to organic substance [inference]

cellular response to cytokine stimulus [inference]

response to oxygen-containing compound [inference]

response to reactive oxygen species

cellular response to stimulus [inference]

cellular response to chemical stimulus [inference]

cellular response to organic substance [inference]

cellular response to cytokine stimulus [inference]

cellular response to environmental stimulus [inference]

cellular response to abiotic stimulus [inference]

cellular response to pH

cellular response to radiation [inference]

cellular response to light stimulus [inference]

cellular response to UV [inference]

cellular response to UV-B

biological regulation [inference]

regulation of biological process [inference]

regulation of growth [inference]

regulation of cell growth [inference]

negative regulation of growth [inference]

negative regulation of cell growth

positive regulation of growth

positive regulation of cell growth

regulation of locomotion [inference]

positive regulation of locomotion [inference]

positive regulation of cell motility [inference]

positive regulation of cell migration [inference]

positive regulation of epithelial cell migration

regulation of cell motility [inference]

regulation of cell migration [inference]

regulation of epithelial cell migration [inference]

positive regulation of epithelial cell migration

positive regulation of cell migration [inference]

positive regulation of epithelial cell migration

positive regulation of cell motility [inference]

positive regulation of cell migration [inference]

positive regulation of epithelial cell migration

positive regulation of biological process [inference]

positive regulation of locomotion [inference]

positive regulation of cell motility [inference]

positive regulation of cell migration [inference]

positive regulation of epithelial cell migration

positive regulation of growth

positive regulation of cell growth

positive regulation of cellular process [inference]

positive regulation of cell population proliferation [inference]

positive regulation of epithelial cell proliferation

positive regulation of cell growth

positive regulation of cellular component biogenesis [inference]

positive regulation of hyaluranon cable assembly

positive regulation of cell adhesion

positive regulation of cell cycle [inference]

positive regulation of mitotic cell cycle [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle process [inference]

positive regulation of cell cycle phase transition [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle G1/S phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cellular component organization [inference]

positive regulation of hyaluranon cable assembly

positive regulation of cell motility [inference]

positive regulation of cell migration [inference]

positive regulation of epithelial cell migration

positive regulation of developmental process [inference]

positive regulation of vasculature development [inference]

positive regulation of angiogenesis

positive regulation of multicellular organismal process [inference]

positive regulation of vasculature development [inference]

positive regulation of angiogenesis

negative regulation of biological process [inference]

negative regulation of growth [inference]

negative regulation of cell growth

negative regulation of cellular process [inference]

negative regulation of cell growth

regulation of developmental process [inference]

regulation of anatomical structure morphogenesis [inference]

regulation of angiogenesis [inference]

positive regulation of angiogenesis

positive regulation of developmental process [inference]

positive regulation of vasculature development [inference]

positive regulation of angiogenesis

regulation of multicellular organismal development [inference]

regulation of vasculature development [inference]

regulation of angiogenesis [inference]

positive regulation of angiogenesis

positive regulation of vasculature development [inference]

positive regulation of angiogenesis

regulation of cellular process [inference]

regulation of cell adhesion [inference]

positive regulation of cell adhesion

regulation of cell population proliferation [inference]

positive regulation of cell population proliferation [inference]

positive regulation of epithelial cell proliferation

regulation of epithelial cell proliferation [inference]

positive regulation of epithelial cell proliferation

regulation of cellular component biogenesis [inference]

positive regulation of cellular component biogenesis [inference]

positive regulation of hyaluranon cable assembly

regulation of hyaluranon cable assembly [inference]

positive regulation of hyaluranon cable assembly

positive regulation of cellular process [inference]

positive regulation of cell population proliferation [inference]

positive regulation of epithelial cell proliferation

positive regulation of cell growth

positive regulation of cellular component biogenesis [inference]

positive regulation of hyaluranon cable assembly

positive regulation of cell adhesion

positive regulation of cell cycle [inference]

positive regulation of mitotic cell cycle [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle process [inference]

positive regulation of cell cycle phase transition [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle G1/S phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cellular component organization [inference]

positive regulation of hyaluranon cable assembly

positive regulation of cell motility [inference]

positive regulation of cell migration [inference]

positive regulation of epithelial cell migration

negative regulation of cellular process [inference]

negative regulation of cell growth

regulation of cellular component organization [inference]

regulation of cell growth [inference]

positive regulation of cellular component organization [inference]

positive regulation of hyaluranon cable assembly

regulation of hyaluranon cable assembly [inference]

positive regulation of hyaluranon cable assembly

regulation of cell cycle [inference]

regulation of mitotic cell cycle [inference]

positive regulation of mitotic cell cycle [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of mitotic cell cycle phase transition [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of G1/S transition of mitotic cell cycle [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of cell cycle process [inference]

positive regulation of cell cycle process [inference]

positive regulation of cell cycle phase transition [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle G1/S phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of cell cycle phase transition [inference]

positive regulation of cell cycle phase transition [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle G1/S phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of mitotic cell cycle phase transition [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of G1/S transition of mitotic cell cycle [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of cell cycle G1/S phase transition [inference]

positive regulation of cell cycle G1/S phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of G1/S transition of mitotic cell cycle [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle [inference]

positive regulation of mitotic cell cycle [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle process [inference]

positive regulation of cell cycle phase transition [inference]

positive regulation of mitotic cell cycle phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

positive regulation of cell cycle G1/S phase transition [inference]

positive regulation of G1/S transition of mitotic cell cycle

regulation of cell motility [inference]

regulation of cell migration [inference]

regulation of epithelial cell migration [inference]

positive regulation of epithelial cell migration

positive regulation of cell migration [inference]

positive regulation of epithelial cell migration

positive regulation of cell motility [inference]

positive regulation of cell migration [inference]

positive regulation of epithelial cell migration

regulation of multicellular organismal process [inference]

regulation of epithelial cell migration [inference]

positive regulation of epithelial cell migration

positive regulation of multicellular organismal process [inference]

positive regulation of vasculature development [inference]

positive regulation of angiogenesis

regulation of multicellular organismal development [inference]

regulation of vasculature development [inference]

regulation of angiogenesis [inference]

positive regulation of angiogenesis

positive regulation of vasculature development [inference]

positive regulation of angiogenesis

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GO Term	Evidence Code	PMID
cytoplasm	IDA
hyaluranon cable	IDA	16900089
extracellular space	IDA	17170110 21695196 9223416
lysosomal lumen	TAS
extracellular exosome	HDA	23533145

GO Hierarchy

cellular component

cellular anatomical entity [inference]

membrane-enclosed lumen [inference]

organelle lumen [inference]

intracellular organelle lumen [inference]

vacuolar lumen [inference]

lysosomal lumen

hyaluranon cable

organelle [inference]

membrane-bounded organelle [inference]

vesicle [inference]

intracellular vesicle [inference]

cytoplasmic vesicle

extracellular vesicle [inference]

extracellular exosome

intracellular membrane-bounded organelle [inference]

vacuole [inference]

lytic vacuole [inference]

lysosome

intracellular vesicle [inference]

cytoplasmic vesicle

extracellular membrane-bounded organelle [inference]

extracellular vesicle [inference]

extracellular exosome

intracellular organelle [inference]

intracellular membrane-bounded organelle [inference]

vacuole [inference]

lytic vacuole [inference]

lysosome

intracellular vesicle [inference]

cytoplasmic vesicle

extracellular organelle [inference]

extracellular membrane-bounded organelle [inference]

extracellular vesicle [inference]

extracellular exosome

Displaying **all 4** entries
GO Term	Evidence Code	PMID
virus receptor activity	IDA	11296287
hyaluronan synthase activity	IDA
hyalurononglucosaminidase activity	IDA TAS	11296287 11944887 17170110 18390475 19478093 20473947 20572808 21695196 21699545 9223416
chondroitin hydrolase activity	IEA

GO Hierarchy

molecular function

catalytic activity [inference]

transferase activity [inference]

glycosyltransferase activity [inference]

UDP-glycosyltransferase activity [inference]

hyaluronan synthase activity

hexosyltransferase activity [inference]

hyaluronan synthase activity

hydrolase activity [inference]

hydrolase activity, acting on glycosyl bonds [inference]

hydrolase activity, hydrolyzing O-glycosyl compounds [inference]

hexosaminidase activity [inference]

hyalurononglucosaminidase activity

endogalactosaminidase activity [inference]

chondroitin hydrolase activity

binding [inference]

protein binding [inference]

exogenous protein binding [inference]

virus receptor activity

Gene Ontology information from NCBI Gene and UniProt.

KEGG BRITE Database

Orthology

K01197

Name

hyaluronoglucosaminidase [EC:3.2.1.35]

References

Reactions

Displaying **all 3** entries
KEGG Reaction	Enzyme	Acceptor	Product
R07825	3.2.1.35	G34586BV	G92988PL
R07824	3.2.1.35	G81393WR	G92988PL
R10905	3.2.1.35	G16235VG	G16235VG G93263XO

Disease

Disease Ontology

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DO ID	Disease Name	Source
DOID:0050809	mucopolysaccharidosis IX	DisGeNET Alliance of Genome Resources
DOID:0050865	tongue squamous cell carcinoma	DisGeNET
DOID:0080199	colorectal carcinoma	DisGeNET
DOID:10283	prostate cancer	DisGeNET DisGeNET DisGeNET
DOID:10286	prostate carcinoma	DisGeNET DisGeNET DisGeNET
DOID:11054	urinary bladder cancer	DisGeNET DisGeNET DisGeNET
DOID:11934	head and neck cancer	DisGeNET
DOID:12798	mucopolysaccharidosis	DisGeNET DisGeNET Alliance of Genome Resources
DOID:12799	mucopolysaccharidosis II	DisGeNET DisGeNET
DOID:12800	mucopolysaccharidosis VI	DisGeNET DisGeNET

The Human Phenotype Ontology

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HPO ID	HPO Term
HP:0000007	Autosomal recessive inheritance
HP:0000176	Submucous cleft hard palate
HP:0000193	Bifid uvula
HP:0000403	Recurrent otitis media
HP:0000934	Chondrocalcinosis
HP:0003170	Abnormal acetabulum morphology
HP:0003621	Juvenile onset
HP:0004322	Short stature
HP:0004626	Lumbar scoliosis
HP:0005072	Wrist hypermobility

Displaying **all 2** entries
Disease ID	Disease Name
ORPHA:67041	mucopolysaccharidosis type 9
OMIM:601492	mucopolysaccharidosis type 9

PubChem Disease

MedGen Diseases

Deficiency of hyaluronoglucosaminidase

OMIM Phenotypes

MUCOPOLYSACCHARIDOSIS, TYPE IX; MPS9

KEGG Diseases

Gene-Disease-Co-Occurrences-in-Literature (PubChem)

Ortholog

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Species	Gene ID	Alliance of Genome Resources	Orthologous MAtrix
Caenorhabditis elegans	174383	WB:WBGene00011923
Latimeria chalumnae	102358666		LATCH15912
Danio rerio	664772	ZFIN:ZDB-GENE-060312-42
Ictalurus punctatus	108271980		ICTPU02883
Electrophorus electricus	113571631		ELEEL00362
Salmo salar	106565982		SALSA62986
Salmo trutta	115207485		SALTR37346
Gadus morhua	115557423		GADMO08638
Oryzias latipes	101169103		ORYLA13848
Oreochromis niloticus	100699578		ORENI56824

GlyTouCan

GlyComb

GlycoPOST

UniCarb-DR

All Resources

Other Information

Tools

Search Tools

Summary

External Links

Annotation

KEGG BRITE Database

PubChem Disease

International Collaboration

Acknowledgements