DB code: T00005

CATH domain 2.60.120.260 : Jelly Rolls
2.130.10.80 : Methylamine Dehydrogenase; Chain H Catalytic domain
2.60.40.10 : Immunoglobulin-like Catalytic domain
E.C. 1.1.3.9
CSA 1gog
M-CSA 1gog
MACiE M0322

CATH domain Related DB codes (homologues)
2.60.120.260 : Jelly Rolls M00124 T00065 T00066
2.60.40.10 : Immunoglobulin-like M00131 T00257 M00113 M00127 M00132 M00323 M00325 M00327 M00329 M00330 M00331 M00332 T00307 D00166 D00500 M00112 M00193 T00063 T00065 T00067 T00245

Uniprot Enzyme Name
UniprotKB Protein name Synonyms
Q01745
None

KEGG enzyme name
galactose oxidase
D-galactose oxidase
beta-galactose oxidase

UniprotKB: Accession Number Entry name Activity Subunit Subcellular location Cofactor
Q01745 GAOA_GIBZE D-galactose + O(2) = D-galacto-hexodialdose + H(2)O(2). Monomer. Secreted. Binds 1 Cu(2+) ion per subunit.

KEGG Pathways
Map code Pathways E.C.
MAP00052 Galactose metabolism

Compound table
Cofactors Substrates Products Intermediates
KEGG-id C00070 C00124 C00007 C03269 C00027
E.C.
Compound Copper D-Galactose O2 D-Galacto-hexodialdose H2O2
Type heavy metal carbohydrate others carbohydrate others
ChEBI 28694
30052
4139
15379
26689
27140
16240
PubChem 23978
6036
977
439962
22326046
784
1gofA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1gogA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1gohA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1k3iA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1t2xA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1gofA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Bound:_CU Unbound Unbound Analogue:ACY_703 Unbound
1gogA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Bound:_CU Unbound Unbound Unbound Unbound
1gohA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1k3iA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1t2xA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Bound:_CU Unbound Unbound Unbound Unbound
1gofA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1gogA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1gohA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1k3iA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound
1t2xA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain Unbound Unbound Unbound Unbound Unbound

Reference for Active-site residues
resource references E.C.
Swiss-prot;Q01745 & literature [9], [14] & [32]

Active-site residues
PDB Catalytic residues Cofactor-binding residues Modified residues Main-chain involved in catalysis Comment
1gofA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain
1gogA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain
1gohA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain
1k3iA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain
1t2xA01 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain
1gofA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain CYS 228;TYR 272;TRP 290;TYR 495 TYR 272;TYR 495;HIS 496(Copper binding)
1gogA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain CYS 228;TYR 272;TRP 290;TYR 495 TYR 272;TYR 495;HIS 496(Copper binding)
1gohA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain CYS 228;TYR 272;TRP 290;TYR 495 TYR 272;TYR 495;HIS 496(Copper binding)
1k3iA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain ;TYR 272;TRP 290;TYR 495 TYR 272;TYR 495;HIS 496(Copper binding) CSO 228
1t2xA02 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain CYS 228;TYR 272;TRP 290;TYR 495 TYR 272;TYR 495;HIS 496(Copper binding)
1gofA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain HIS 581(Copper binding)
1gogA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain HIS 581(Copper binding)
1gohA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain HIS 581(Copper binding)
1k3iA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain HIS 581(Copper binding)
1t2xA03 Pdbj logo s Rasmollogo id Rasmollogo chain Mmcif id Mmcif chain HIS 581(Copper binding)

References for Catalytic Mechanism
References Sections No. of steps in catalysis
[6]
Fig.6
[9]
p.88
[10]
Fig.7, p.82-83
[12]
Fig.9, p.770-771
[14]
p.808-810
[15]
Fig.1, p.510S
[18]
Scheme 1, Scheme 4, Scheme 5
[19]
Fig.8, Fig.9, p.8430-8435
[23]
Fig.2, p.248-249
[26]
Fig.4
[27]
Scheme 1
[32]
Fig.4, Fig.15, Fig.17, p.2358-2361

References
[1]
Resource
Comments
Medline ID
PubMed ID 20059
Journal Arch Biochem Biophys
Year 1977
Volume 182
Pages 712-22
Authors Kwiatkowski LD, Siconolfi L, Weiner RE, Giordano RS, Bereman RD, Ettinger MJ, Kosman DJ
Title Histidine as an essential residue in the active site of the copper enzyme galactose oxidase.
Related PDB
Related UniProtKB
[2]
Resource
Comments
Medline ID
PubMed ID 902277
Journal Carbohydr Res
Year 1977
Volume 57
Pages 273-80
Authors Whyte JN, Englar JR
Title An enzyme-P.M.R.-spectroscopic determination of the enantiomers of galactose.
Related PDB
Related UniProtKB
[3]
Resource
Comments
Medline ID
PubMed ID 6248103
Journal Biochemistry
Year 1980
Volume 19
Pages 1304-8
Authors Kosman DJ, Peisach J, Mims WB
Title Pulsed electron paramagnetic resonance studies of the copper(II) site in galactose oxidase.
Related PDB
Related UniProtKB
[4]
Resource
Comments
Medline ID
PubMed ID 7213759
Journal Biochim Biophys Acta
Year 1981
Volume 657
Pages 495-506
Authors Marwedel BJ, Kurland RJ
Title Fluoride ion as an NMR relaxation probe of galactose oxidase-substrate binding.
Related PDB
Related UniProtKB
[5]
Resource
Comments
Medline ID
PubMed ID 6267193
Journal J Inorg Biochem
Year 1981
Volume 14
Pages 223-35
Authors Winkler ME, Bereman RD, Kurland RJ
Title Kinetic and magnetic resonance studies of substrate binding to galactose oxidase copper(II).
Related PDB
Related UniProtKB
[6]
Resource
Comments
Medline ID
PubMed ID 2834363
Journal J Biol Chem
Year 1988
Volume 263
Pages 6074-80
Authors Whittaker MM, Whittaker JW
Title The active site of galactose oxidase.
Related PDB
Related UniProtKB
[7]
Resource
Comments
Medline ID
PubMed ID 2542244
Journal J Biol Chem
Year 1989
Volume 264
Pages 7792-4
Authors van der Meer RA, Jongejan JA, Duine JA
Title Pyrroloquinoline quinone as cofactor in galactose oxidase (EC 1.1.3.9).
Related PDB
Related UniProtKB
[8]
Resource
Comments
Medline ID
PubMed ID 2161837
Journal J Biol Chem
Year 1990
Volume 265
Pages 9610-3
Authors Whittaker MM, Whittaker JW
Title A tyrosine-derived free radical in apogalactose oxidase.
Related PDB
Related UniProtKB
[9]
Resource
Comments X-RAY CRYSTALLOGRAPHY (1.7 ANGSTROMS).
Medline ID 91163641
PubMed ID 2002850
Journal Nature
Year 1991
Volume 350
Pages 87-90
Authors Ito N, Phillips SE, Stevens C, Ogel ZB, McPherson MJ, Keen JN, Yadav KD, Knowles PF
Title Novel thioether bond revealed by a 1.7 A crystal structure of galactose oxidase.
Related PDB 1gof 1gog 1goh
Related UniProtKB Q01745
[10]
Resource
Comments
Medline ID
PubMed ID 1290941
Journal Faraday Discuss
Year 1992
Volume -93
Pages 75-84
Authors Ito N, Phillips SE, Stevens C, Ogel ZB, McPherson MJ, Keen JN, Yadav KD, Knowles PF
Title Three-dimensional structure of galactose oxidase: an enzyme with a built-in secondary cofactor.
Related PDB
Related UniProtKB
[11]
Resource
Comments
Medline ID
PubMed ID 8224464
Journal Biochem Soc Trans
Year 1993
Volume 21
Pages 319S
Authors Baron AJ, Stevens C, Wilmot CM, Knowles PF, Phillips SE, McPherson MJ
Title Preliminary studies of two active site mutants of galactose oxidase.
Related PDB
Related UniProtKB
[12]
Resource
Comments
Medline ID
PubMed ID 8386015
Journal Biophys J
Year 1993
Volume 64
Pages 762-72
Authors Whittaker MM, Whittaker JW
Title Ligand interactions with galactose oxidase: mechanistic insights.
Related PDB
Related UniProtKB
[13]
Resource
Comments
Medline ID
PubMed ID 7929198
Journal J Biol Chem
Year 1994
Volume 269
Pages 25095-105
Authors Baron AJ, Stevens C, Wilmot C, Seneviratne KD, Blakeley V, Dooley DM, Phillips SE, Knowles PF, McPherson MJ
Title Structure and mechanism of galactose oxidase. The free radical site.
Related PDB
Related UniProtKB
[14]
Resource
Comments
Medline ID
PubMed ID 8182749
Journal J Mol Biol
Year 1994
Volume 238
Pages 794-814
Authors Ito N, Phillips SE, Yadav KD, Knowles PF
Title Crystal structure of a free radical enzyme, galactose oxidase.
Related PDB
Related UniProtKB
[15]
Resource
Comments
Medline ID
PubMed ID 8654695
Journal Biochem Soc Trans
Year 1995
Volume 23
Pages 510S
Authors Reynolds MP, Baron AJ, Wilmot CM, Phillips SE, Knowles PF, McPherson MJ
Title Tyrosine 495 is a key residue in the active site of galactose oxidase.
Related PDB
Related UniProtKB
[16]
Resource
Comments
Medline ID
PubMed ID 8524154
Journal Methods Enzymol
Year 1995
Volume 258
Pages 235-62
Authors Ito N, Knowles PF, Phillips SE
Title X-ray crystallographic studies of cofactors in galactose oxidase.
Related PDB
Related UniProtKB
[17]
Resource
Comments
Medline ID
PubMed ID 8524155
Journal Methods Enzymol
Year 1995
Volume 258
Pages 262-77
Authors Whittaker JW
Title Spectroscopic studies of galactose oxidase.
Related PDB
Related UniProtKB
[18]
Resource
Comments
Medline ID
PubMed ID 8916929
Journal Biochemistry
Year 1996
Volume 35
Pages 14425-35
Authors Wachter RM, Branchaud BP
Title Thiols as mechanistic probes for catalysis by the free radical enzyme galactose oxidase.
Related PDB
Related UniProtKB
[19]
Resource
Comments
Medline ID
PubMed ID 9622494
Journal Biochemistry
Year 1998
Volume 37
Pages 8426-36
Authors Whittaker MM, Ballou DP, Whittaker JW
Title Kinetic isotope effects as probes of the mechanism of galactose oxidase.
Related PDB
Related UniProtKB
[20]
Resource
Comments
Medline ID
PubMed ID 9684896
Journal Protein Sci
Year 1998
Volume 7
Pages 1626-31
Authors Baumgartner S, Hofmann K, Chiquet-Ehrismann R, Bucher P
Title The discoidin domain family revisited: new members from prokaryotes and a homology-based fold prediction.
Related PDB
Related UniProtKB
[21]
Resource
Comments
Medline ID
PubMed ID 9438841
Journal Science
Year 1998
Volume 279
Pages 537-40
Authors Wang Y, DuBois JL, Hedman B, Hodgson KO, Stack TD
Title Catalytic galactose oxidase models: biomimetic Cu(II)-phenoxyl-radical reactivity.
Related PDB
Related UniProtKB
[22]
Resource
Comments
Medline ID
PubMed ID 10805536
Journal Anal Biochem
Year 2000
Volume 280
Pages 173-7
Authors Basu SS, Dotson GD, Raetz CR
Title A facile enzymatic synthesis of uridine diphospho-[14C]galacturonic acid.
Related PDB
Related UniProtKB
[23]
Resource
Comments
Medline ID
PubMed ID 10819469
Journal J Biol Inorg Chem
Year 2000
Volume 5
Pages 236-50
Authors Rothlisberger U, Carloni P, Doclo K, Parrinello M
Title A comparative study of galactose oxidase and active site analogs based on QM/MM Car-Parrinello simulations.
Related PDB
Related UniProtKB
[24]
Resource
Comments
Medline ID
PubMed ID 11948877
Journal Chembiochem
Year 2001
Volume 2
Pages 884-94
Authors Bulter T, Schumacher T, Namdjou DJ, Gutierrez Gallego R, Clausen H, Elling L
Title Chemoenzymatic synthesis of biotinylated nucleotide sugars as substrates for glycosyltransferases.
Related PDB
Related UniProtKB
[25]
Resource
Comments
Medline ID
PubMed ID 11170534
Journal Inorg Chem
Year 2001
Volume 40
Pages 294-300
Authors Wright C, Im SC, Twitchett MB, Saysell CG, Sokolowski A, Sykes AG
Title Thermodynamic, kinetic and pH studies on the reactions of NCS-, N3-, and CH3CO2- with Fusarium galactose oxidase.
Related PDB
Related UniProtKB
[26]
Resource
Comments
Medline ID
PubMed ID 11698678
Journal Proc Natl Acad Sci U S A
Year 2001
Volume 98
Pages 12932-7
Authors Firbank SJ, Rogers MS, Wilmot CM, Dooley DM, Halcrow MA, Knowles PF, McPherson MJ, Phillips SE
Title Crystal structure of the precursor of galactose oxidase: an unusual self-processing enzyme.
Related PDB 1k3i
Related UniProtKB
[27]
Resource
Comments
Medline ID
PubMed ID 12203454
Journal Angew Chem Int Ed Engl
Year 2002
Volume 41
Pages 3047-50
Authors Thomas F, Gellon G, Gautier-Luneau I, Saint-Aman E, Pierre JL
Title A structural and functional model of galactose oxidase: control of the one-electron oxidized active form through two differentiated phenolic arms in a tripodal ligand.
Related PDB
Related UniProtKB
[28]
Resource
Comments
Medline ID
PubMed ID 12009437
Journal Bioelectrochemistry
Year 2002
Volume 56
Pages 23-5
Authors Tkac J, Vostiar I, Gemeiner P, Sturdik E
Title Indirect evidence of direct electron communication between the active site of galactose oxidase and a graphite electrode.
Related PDB
Related UniProtKB
[29]
Resource
Comments
Medline ID
PubMed ID 12058898
Journal J Chromatogr A
Year 2002
Volume 954
Pages 137-50
Authors Mazitsos CF, Rigden DJ, Tsoungas PG, Clonis YD
Title Galactosyl-biomimetic dye-ligands for the purification of Dactylium dendroides galactose oxidase.
Related PDB
Related UniProtKB
[30]
Resource
Comments
Medline ID
PubMed ID 12027633
Journal Org Lett
Year 2002
Volume 4
Pages 1863-6
Authors Andreana PR, Xie W, Cheng HN, Qiao L, Murphy DJ, Gu QM, Wang PG
Title In situ preparation of beta-D-1-O-hydroxylamino carbohydrate polymers mediated by galactose oxidase.
Related PDB
Related UniProtKB
[31]
Resource
Comments
Medline ID
PubMed ID 12027632
Journal Org Lett
Year 2002
Volume 4
Pages 1859-62
Authors Chang D, Feiten HJ, Engesser KH, van Beilen JB, Witholt B, Li Z
Title Practical syntheses of N-substituted 3-hydroxyazetidines and 4-hydroxypiperidines by hydroxylation with Sphingomonas sp. HXN-200.
Related PDB
Related UniProtKB
[32]
Resource
Comments
Medline ID
PubMed ID 12797833
Journal Chem Rev
Year 2003
Volume 103
Pages 2347-63
Authors Whittaker JW
Title Free radical catalysis by galactose oxidase.
Related PDB
Related UniProtKB
[33]
Resource
Comments
Medline ID
PubMed ID 15047910
Journal Protein Eng Des Sel
Year 2004
Volume 17
Pages 141-8
Authors Wilkinson D, Akumanyi N, Hurtado-Guerrero R, Dawkes H, Knowles PF, Phillips SE, McPherson MJ
Title Structural and kinetic studies of a series of mutants of galactose oxidase identified by directed evolution.
Related PDB 1t2x
Related UniProtKB

Comments
This enzyme catalyze two distinct reactions:
(A) Reduction of alcohol substrate, to produce aldehyde. (Hydride transfer)
(B) Reduction of dioxygen (O2), to produce hydrogen peroxide (H2O2) (Hydride transfer)

Created Updated
2005-04-28 2009-02-26