Reviewed, UniProtKB/Swiss-Prot Q12794 (HYAL1_HUMAN)

Names and origin

Protein names

Recommended name:
    Hyaluronidase-1
      Short name=Hyal-1
    EC=3.2.1.35
Alternative name(s):
    Hyaluronoglucosaminidase-1
      Short name=LUCA-1

Gene names

Name:	HYAL1
Synonyms:	LUCA1

Organism

Homo sapiens (Human) [Complete proteome]

Taxonomic identifier

9606 [NCBI]

Taxonomic lineage

Eukaryota › Metazoa › Chordata › Craniata › Vertebrata › Euteleostomi › Mammalia › Eutheria › Euarchontoglires › Primates › Haplorrhini › Catarrhini › Hominidae › Homo

Protein attributes

Sequence length	435 AA.
Sequence status	Complete.
Sequence processing	The displayed sequence is further processed into a mature form.
Protein existence	Evidence at protein level.

General annotation (Comments)

Function	May have a role in promoting tumor progression. May block the TGFB1-enhanced cell growth. Ref.4
Catalytic activity	Random hydrolysis of (1->4)-linkages between N-acetyl-beta-D-glucosamine and D-glucuronate residues in hyaluronate.
Subcellular location	Secreted. Lysosome. Ref.2
Tissue specificity	Highly expressed in the liver, kidney and heart. Weakly expressed in lung, placenta and skeletal muscle. No expression detected in adult brain. Isoform 1 is expressed only in bladder and prostate cancer cells, G2/G3 bladder tumor tissues and lymph node specimens showing tumor invasive tumors cells. Isoform 3, isoform 4, isoform 5 and isoform 6 are expressed in normal bladder and bladder tumor tissues. Ref.4 Ref.2 Ref.10
Involvement in disease	Defects in HYAL1 are the cause of mucopolysaccharidosis type 9 (MPS9) [MIM:601492]; also called hyaluronidase deficiency. MPS9 is a lysosomal storage disease characterized by high hyaluronan (HA) concentration in the serum. The clinical features are periarticular soft tissue masses, mild short stature and acetabular erosions, and absence of neurological or visceral involvement. Ref.10
Sequence similarities	Belongs to the glycosyl hydrolase 56 family. Contains 1 EGF-like domain.
Biophysicochemical properties	pH dependence: Optimum pH is about 3.8.

Ontologies

Keywords
Cellular component	Lysosome Secreted
Coding sequence diversity	Alternative splicing
Disease	Disease mutation Mucopolysaccharidosis
Domain	EGF-like domain Signal
Molecular function	Glycosidase Hydrolase
PTM	Disulfide bond Glycoprotein
Technical term	3D-structure Complete proteome Direct protein sequencing
Gene Ontology (GO)
Biological process	glycosaminoglycan metabolic process Inferred from electronic annotation. Source: UniProtKB-KW
Cellular component	extracellular space Ref.2 Traceable author statement. Source: ProtInc lysosome Inferred from electronic annotation. Source: UniProtKB-SubCell
Molecular function	hyalurononglucosaminidase activity Ref.2 Traceable author statement. Source: ProtInc
Complete GO annotation...

Alternative products

This entry describes 7 isoforms produced by alternative splicing. [Align] [Select]

Isoform 1 (identifier: Q12794-1)

This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

Isoform 2 (identifier: Q12794-2)

Also known as: HYAl1v1;

The sequence of this isoform differs from the canonical sequence as follows:
     301-330: Missing.

Note: Enzymatically inactive.

Isoform 3 (identifier: Q12794-3)

Also known as: HYAl1v2;

The sequence of this isoform differs from the canonical sequence as follows:
     1-182: Missing.

Note: Enzymatically inactive.

Isoform 4 (identifier: Q12794-4)

Also known as: HYAl1v3;

The sequence of this isoform differs from the canonical sequence as follows:
     208-209: YN → SG
     210-435: Missing.

Note: Enzymatically inactive.

Isoform 5 (identifier: Q12794-5)

Also known as: HYAl1v4;

The sequence of this isoform differs from the canonical sequence as follows:
     1-259: Missing.

Note: Enzymatically inactive.

Isoform 6 (identifier: Q12794-6)

Also known as: HYAl1v5;

The sequence of this isoform differs from the canonical sequence as follows:
     1-339: Missing.

Note: Enzymatically inactive.

Isoform 7 (identifier: Q12794-7)

The sequence of this isoform differs from the canonical sequence as follows:
     331-336: ESCQAI → VSLGLA
     337-435: Missing.

Note: No experimental confirmation available.

Sequence annotation (Features)

Feature key

Position(s)

Length

Description

Graphical view

Feature identifier

Molecule processing

Signal peptide

1 – 21

21

Potential

Chain

22 – 435

414

Hyaluronidase-1

PRO_0000042622

Regions

Domain

354 – 430

77

EGF-like

Sites

Active site

131

1

Proton donor Probable

Amino acid modifications

Glycosylation

99

1

N-linked (GlcNAc...) Ref.9

Glycosylation

216

1

N-linked (GlcNAc...) Ref.9

Glycosylation

350

1

N-linked (GlcNAc...) Ref.9

Disulfide bond

43 ↔ 333

Ref.9

Disulfide bond

207 ↔ 221

Ref.9

Disulfide bond

358 ↔ 369

Ref.9

Disulfide bond

363 ↔ 418

Ref.9

Disulfide bond

420 ↔ 429

Ref.9

Natural variations

Alternative sequence

1 – 339

339

Missing in isoform 6.

VSP_015915

Alternative sequence

1 – 259

259

Missing in isoform 5.

VSP_015916

Alternative sequence

1 – 182

182

Missing in isoform 3.

VSP_015917

Alternative sequence

208 – 209

2

YN → SG in isoform 4.

VSP_015918

Alternative sequence

210 – 435

226

Missing in isoform 4.

VSP_015919

Alternative sequence

301 – 330

30

Missing in isoform 2.

VSP_015920

Alternative sequence

331 – 336

6

ESCQAI → VSLGLA in isoform 7.

VSP_015921

Alternative sequence

337 – 435

99

Missing in isoform 7.

VSP_015922

Natural variant

268

1

E → K in MPS9. Ref.10

VAR_023643

Experimental info

Sequence conflict

3

1

A → G in CAG46731. Ref.6

Sequence conflict

191

1

R → G in AAD53277. Ref.1

Sequence conflict

300

1

L → Q in AAD24460. Ref.3

Secondary structure

1

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

435

Helix Strand Turn

Details...

Beta strand

24 – 27

4

Beta strand

30 – 38

9

Helix

40 – 47

8

Beta strand

56 – 59

4

Beta strand

69 – 74

6

Beta strand

78 – 80

3

Helix

97 – 99

3

Helix

102 – 116

15

Beta strand

124 – 128

5

Helix

137 – 139

3

Helix

142 – 144

3

Helix

145 – 158

14

Beta strand

159 – 161

3

Helix

164 – 193

30

Beta strand

197 – 202

6

Helix

223 – 230

8

Helix

233 – 238

6

Beta strand

240 – 243

4

Helix

250 – 252

3

Helix

258 – 275

18

Helix

300 – 304

5

Helix

307 – 312

6

Beta strand

316 – 321

6

Helix

324 – 326

3

Beta strand

327 – 329

3

Helix

330 – 342

13

Helix

344 – 362

19

Beta strand

366 – 371

6

Turn

384 – 386

3

Beta strand

387 – 391

5

Helix

393 – 395

3

Beta strand

398 – 402

5

Helix

406 – 415

10

Beta strand

416 – 420

5

Turn

426 – 429

4

Sequences

	Sequence		Length	Mass (Da)
	Isoform 1 [UniParc]. Last modified March 1, 2001. Version 2. Checksum: 9C2B2D8DB361E0BB	FASTA	435	48,368
	10 20 30 40 50 60 MAAHLLPICA LFLTLLDMAQ GFRGPLLPNR PFTTVWNANT QWCLERHGVD VDVSVFDVVA 70 80 90 100 110 120 NPGQTFRGPD MTIFYSSQLG TYPYYTPTGE PVFGGLPQNA SLIAHLARTF QDILAAIPAP 130 140 150 160 170 180 DFSGLAVIDW EAWRPRWAFN WDTKDIYRQR SRALVQAQHP DWPAPQVEAV AQDQFQGAAR 190 200 210 220 230 240 AWMAGTLQLG RALRPRGLWG FYGFPDCYNY DFLSPNYTGQ CPSGIRAQND QLGWLWGQSR 250 260 270 280 290 300 ALYPSIYMPA VLEGTGKSQM YVQHRVAEAF RVAVAAGDPN LPVLPYVQIF YDTTNHFLPL 310 320 330 340 350 360 DELEHSLGES AAQGAAGVVL WVSWENTRTK ESCQAIKEYM DTTLGPFILN VTSGALLCSQ 370 380 390 400 410 420 ALCSGHGRCV RRTSHPKALL LLNPASFSIQ LTPGGGPLSL RGALSLEDQA QMAVEFKCRC 430 YPGWQAPWCE RKSMW
	Isoform 2 (HYAl1v1). Checksum: C24DAD360D9CF032	FASTA	405	45,145
	10 20 30 40 50 60 MAAHLLPICA LFLTLLDMAQ GFRGPLLPNR PFTTVWNANT QWCLERHGVD VDVSVFDVVA 70 80 90 100 110 120 NPGQTFRGPD MTIFYSSQLG TYPYYTPTGE PVFGGLPQNA SLIAHLARTF QDILAAIPAP 130 140 150 160 170 180 DFSGLAVIDW EAWRPRWAFN WDTKDIYRQR SRALVQAQHP DWPAPQVEAV AQDQFQGAAR 190 200 210 220 230 240 AWMAGTLQLG RALRPRGLWG FYGFPDCYNY DFLSPNYTGQ CPSGIRAQND QLGWLWGQSR 250 260 270 280 290 300 ALYPSIYMPA VLEGTGKSQM YVQHRVAEAF RVAVAAGDPN LPVLPYVQIF YDTTNHFLPL 310 320 330 340 350 360 ESCQAIKEYM DTTLGPFILN VTSGALLCSQ ALCSGHGRCV RRTSHPKALL LLNPASFSIQ 370 380 390 400 LTPGGGPLSL RGALSLEDQA QMAVEFKCRC YPGWQAPWCE RKSMW
	Isoform 3 (HYAl1v2). Checksum: 441C0DB43301E03F	FASTA	253	27,958
	10 20 30 40 50 60 MAGTLQLGRA LRPRGLWGFY GFPDCYNYDF LSPNYTGQCP SGIRAQNDQL GWLWGQSRAL 70 80 90 100 110 120 YPSIYMPAVL EGTGKSQMYV QHRVAEAFRV AVAAGDPNLP VLPYVQIFYD TTNHFLPLDE 130 140 150 160 170 180 LEHSLGESAA QGAAGVVLWV SWENTRTKES CQAIKEYMDT TLGPFILNVT SGALLCSQAL 190 200 210 220 230 240 CSGHGRCVRR TSHPKALLLL NPASFSIQLT PGGGPLSLRG ALSLEDQAQM AVEFKCRCYP 250 GWQAPWCERK SMW
	Isoform 4 (HYAl1v3). Checksum: 5C1046F8199F3BFF	FASTA	209	23,338
	10 20 30 40 50 60 MAAHLLPICA LFLTLLDMAQ GFRGPLLPNR PFTTVWNANT QWCLERHGVD VDVSVFDVVA 70 80 90 100 110 120 NPGQTFRGPD MTIFYSSQLG TYPYYTPTGE PVFGGLPQNA SLIAHLARTF QDILAAIPAP 130 140 150 160 170 180 DFSGLAVIDW EAWRPRWAFN WDTKDIYRQR SRALVQAQHP DWPAPQVEAV AQDQFQGAAR 190 200 AWMAGTLQLG RALRPRGLWG FYGFPDCSG
	Isoform 5 (HYAl1v4). Checksum: 75F674283B175FDD	FASTA	176	19,367
	10 20 30 40 50 60 MYVQHRVAEA FRVAVAAGDP NLPVLPYVQI FYDTTNHFLP LDELEHSLGE SAAQGAAGVV 70 80 90 100 110 120 LWVSWENTRT KESCQAIKEY MDTTLGPFIL NVTSGALLCS QALCSGHGRC VRRTSHPKAL 130 140 150 160 170 LLLNPASFSI QLTPGGGPLS LRGALSLEDQ AQMAVEFKCR CYPGWQAPWC ERKSMW
	Isoform 6 (HYAl1v5). Checksum: 70AD570E4305A4E8	FASTA	96	10,435
	10 20 30 40 50 60 MDTTLGPFIL NVTSGALLCS QALCSGHGRC VRRTSHPKAL LLLNPASFSI QLTPGGGPLS 70 80 90 LRGALSLEDQ AQMAVEFKCR CYPGWQAPWC ERKSMW
	Isoform 7. Checksum: 7D19178001F20E51	FASTA	336	37,439
	10 20 30 40 50 60 MAAHLLPICA LFLTLLDMAQ GFRGPLLPNR PFTTVWNANT QWCLERHGVD VDVSVFDVVA 70 80 90 100 110 120 NPGQTFRGPD MTIFYSSQLG TYPYYTPTGE PVFGGLPQNA SLIAHLARTF QDILAAIPAP 130 140 150 160 170 180 DFSGLAVIDW EAWRPRWAFN WDTKDIYRQR SRALVQAQHP DWPAPQVEAV AQDQFQGAAR 190 200 210 220 230 240 AWMAGTLQLG RALRPRGLWG FYGFPDCYNY DFLSPNYTGQ CPSGIRAQND QLGWLWGQSR 250 260 270 280 290 300 ALYPSIYMPA VLEGTGKSQM YVQHRVAEAF RVAVAAGDPN LPVLPYVQIF YDTTNHFLPL 310 320 330 DELEHSLGES AAQGAAGVVL WVSWENTRTK VSLGLA

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"Construction of a 600-kilobase cosmid clone contig and generation of a transcriptional map surrounding the lung cancer tumor suppressor gene (TSG) locus on human chromosome 3p21.3: progress toward the isolation of a lung cancer TSG." Wei M.H., Latif F., Bader S., Kashuba V., Chen J.Y., Duh F.-M., Sekido Y., Lee C.C., Geil L., Kuzmin I., Zabarovsky E., Klein G., Zbar B., Minna J.D., Lerman M.I. Cancer Res. 56:1487-1492(1996) [PubMed: 8603390] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1). Tissue: Heart and Lung cancer.
[2]	"Purification, cloning, and expression of human plasma hyaluronidase." Frost G.I., Csoka A.B., Wong T., Stern R. Biochem. Biophys. Res. Commun. 236:10-15(1997) [PubMed: 9223416] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1), PROTEIN SEQUENCE OF 22-37; 72-84; 248-259 AND 261-273, SUBCELLULAR LOCATION, TISSUE SPECIFICITY. Tissue: Plasma.
[3]	"HYAL1(LUCA-1), a candidate tumor suppressor gene on chromosome 3p21.3, is inactivated in head and neck squamous cell carcinomas by aberrant splicing of pre-mRNA." Frost G.I., Mohapatra G., Wong T.M., Csoka A.B., Gray J.W., Stern R. Oncogene 19:870-877(2000) [PubMed: 10702795] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1).
[4]	"Regulation of hyaluronidase activity by alternative mRNA splicing." Lokeshwar V.B., Schroeder G.L., Carey R.I., Soloway M.S., Iida N. J. Biol. Chem. 277:33654-33663(2002) [PubMed: 12084718] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORMS 2; 3; 4; 5 AND 6), FUNCTION, TISSUE SPECIFICITY.
[5]	Forgacs E., Sekido Y., Bader S., Cundiff S., Compton L., Latif F., Lerman M.I., Minna J.D. Submitted (JUL-1999) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1).
[6]	"Cloning of human full open reading frames in Gateway(TM) system entry vector (pDONR201)." Ebert L., Schick M., Neubert P., Schatten R., Henze S., Korn B. Submitted (JUN-2004) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 2).
[7]	"The DNA sequence, annotation and analysis of human chromosome 3." Muzny D.M., Scherer S.E., Kaul R., Wang J., Yu J., Sudbrak R., Buhay C.J., Chen R., Cree A., Ding Y., Dugan-Rocha S., Gill R., Gunaratne P., Harris R.A., Hawes A.C., Hernandez J., Hodgson A.V., Hume J. , Jackson A., Khan Z.M., Kovar-Smith C., Lewis L.R., Lozado R.J., Metzker M.L., Milosavljevic A., Miner G.R., Morgan M.B., Nazareth L.V., Scott G., Sodergren E., Song X.-Z., Steffen D., Wei S., Wheeler D.A., Wright M.W., Worley K.C., Yuan Y., Zhang Z., Adams C.Q., Ansari-Lari M.A., Ayele M., Brown M.J., Chen G., Chen Z., Clendenning J., Clerc-Blankenburg K.P., Chen R., Chen Z., Davis C., Delgado O., Dinh H.H., Dong W., Draper H., Ernst S., Fu G., Gonzalez-Garay M.L., Garcia D.K., Gillett W., Gu J., Hao B., Haugen E., Havlak P., He X., Hennig S., Hu S., Huang W., Jackson L.R., Jacob L.S., Kelly S.H., Kube M., Levy R., Li Z., Liu B., Liu J., Liu W., Lu J., Maheshwari M., Nguyen B.-V., Okwuonu G.O., Palmeiri A., Pasternak S., Perez L.M., Phelps K.A., Plopper F.J., Qiang B., Raymond C., Rodriguez R., Saenphimmachak C., Santibanez J., Shen H., Shen Y., Subramanian S., Tabor P.E., Verduzco D., Waldron L., Wang J., Wang J., Wang Q., Williams G.A., Wong G.K.-S., Yao Z., Zhang J., Zhang X., Zhao G., Zhou J., Zhou Y., Nelson D., Lehrach H., Reinhardt R., Naylor S.L., Yang H., Olson M., Weinstock G., Gibbs R.A. Nature 440:1194-1198(2006) [PubMed: 16641997] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
[8]	"The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC)." The MGC Project Team Genome Res. 14:2121-2127(2004) [PubMed: 15489334] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORMS 1 AND 7). Tissue: Colon and Pancreas.
[9]	"Structure of human hyaluronidase-1, a hyaluronan hydrolyzing enzyme involved in tumor growth and angiogenesis." Chao K.L., Muthukumar L., Herzberg O. Biochemistry 46:6911-6920(2007) [PubMed: 17503783] [Abstract] Cited for: X-RAY CRYSTALLOGRAPHY (2.0 ANGSTROMS) OF 20-435; MASS SPECTROMETRY, GLYCOSYLATION AT ASN-99; ASN-216 AND ASN-350, DISULFIDE BONDS.
[10]	"Mutations in HYAL1, a member of a tandemly distributed multigene family encoding disparate hyaluronidase activities, cause a newly described lysosomal disorder, mucopolysaccharidosis IX." Triggs-Raine B., Salo T.J., Zhang H., Wicklow B.A., Natowicz M.R. Proc. Natl. Acad. Sci. U.S.A. 96:6296-6300(1999) [PubMed: 10339581] [Abstract] Cited for: VARIANT MPS9 LYS-268, TISSUE SPECIFICITY.
+	Additional computationally mapped references.

Web resources

Atlas of Genetics and Cytogenetics in Oncology and Haematology

GeneReviews

Cross-references

Sequence databases

EMBLGenBankDDBJ

U03056 mRNA. Translation: AAD09137.2.
U96078 mRNA. Translation: AAD04190.1.
AF118821 mRNA. Translation: AAD24460.1.
AF502904 mRNA. Translation: AAM60770.1.
AF502905 mRNA. Translation: AAM60771.1.
AF502906 mRNA. Translation: AAM60772.1.
AF502907 mRNA. Translation: AAM60773.1.
AF502908 mRNA. Translation: AAM60774.1.
AF173154 mRNA. Translation: AAD53277.1.
CR541933 mRNA. Translation: CAG46731.1.
AC002455 Genomic DNA. Translation: AAB67046.1.
U73167 Genomic DNA. Translation: AAC02730.1.
BC025774 mRNA. Translation: AAH25774.1. Different initiation.
BC035695 mRNA. Translation: AAH35695.1.

IPI

IPI00168847.
IPI00170520.
IPI00170521.
IPI00170522.
IPI00170523.
IPI00178140.
IPI00654773.

PIR

JC5584.

RefSeq

NP_009296.1.
NP_149349.2.
NP_695013.1.
NP_695014.1.
NP_695015.1.
NP_695017.1.

UniGene

Hs.75619

3D structure databases

PDBPDBe

Entry	Method	Resolution (Å)	Chain	Positions	PDBsum
2PE4	X-ray	2.00	A	22-435	[»]

ModBase

Search...

Protein family/group databases

CAZy

GH56. Glycoside Hydrolase Family 56.

Proteomic databases

PRIDE

Q12794.

Genome annotation databases

Ensembl

ENSG00000114378. Homo sapiens. [Contig view]

GeneID

3373.

KEGG

hsa:3373.

UCSC

uc003czm.1. human.
uc003czn.1. human.
uc003czp.1. human.
uc003czq.1. human.
uc003czt.2. human.

Organism-specific databases

GeneCards

GC03M050312.

H-InvDB

HIX0003315.

HGNC

HGNC:5320. HYAL1.

HPA

HPA002112.

MIM

601492. phenotype.
607071. gene.

Orphanet

67041. Hyaluronidase deficiency.

PharmGKB

PA29571.

GenAtlas

Search...

Phylogenomic databases

HOVERGEN

Q12794.

OMA

Q12794. ESCQAIK.

Enzyme and pathway databases

BRENDA

3.2.1.35. 247.

Gene expression databases

ArrayExpress

Q12794.

Bgee

Q12794.

GermOnline

ENSG00000114378. Homo sapiens.

Family and domain databases

InterPro

IPR013785. Aldolase_TIM.
IPR006210. EGF-like.
IPR013032. EGF-like_reg_CS.
IPR017430. Glyco_hydro_56_Hyaluronidase.
IPR001968. Glycoside_hydrolase_family_56.
IPR018155. Hyaluronidase.
[Graphical view]

Gene3D

G3DSA:3.20.20.70. Aldolase_TIM. 1 hit.

PANTHER

PTHR11769. Glyco_hydro_56. 1 hit.

Pfam

PF01630. Glyco_hydro_56. 1 hit.
[Graphical view]

PIRSF

PIRSF038193. Hyaluronidase. 1 hit.

PRINTS

PR00846. GLHYDRLASE56.

ProDom

PD003549. Glyco_hydro_56. 1 hit.
[Graphical view] [Entries sharing at least one domain]

SMART

SM00181. EGF. 1 hit.
[Graphical view]

PROSITE

PS00022. EGF_1. False negative.
PS01186. EGF_2. 1 hit.
PS50026. EGF_3. False negative.
[Graphical view]

ProtoNet

Search...

Other Resources

DrugBank

DB00070. Hyaluronidase.

NextBio

13338.

SOURCE

Search...

Entry information

Entry name

HYAL1_HUMAN

Accession

Primary (citable) accession number: Q12794
Secondary accession number(s): Q6FH23 , Q6PIZ6, Q7KYU2, Q7LE34, Q8NFK5, Q8NFK6, Q8NFK7, Q8NFK8, Q8NFK9, Q93013, Q9UKD5, Q9UNI8

Entry history

Integrated into UniProtKB/Swiss-Prot:	October 25, 2005
Last sequence update:	March 1, 2001
Last modified:	July 7, 2009
This is version 77 of the entry and version 2 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation project

HPI (Human Proteome Initiative)