Enzymes

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Editor-In-Chief: C. Michael Gibson, M.S., M.D. [1] ; Associate Editor: Cafer Zorkun, M.D., Ph.D. [2]

v • d • e Proteins: enzymes
Topics	Active site - Allosteric regulation - Binding site - Catalytically perfect enzyme - Coenzyme - Cofactor - Cooperativity - EC number Enzyme catalysis - Enzyme inhibitor - Enzyme kinetics - Lineweaver-Burk plot - Michaelis-Menten kinetics - List of enzymes
Types	EC1 Oxidoreductases/list - EC2 Transferases/list - EC3 Hydrolases/list - EC4 Lyases/list - EC5 Isomerases/list - EC6 Ligases/list

v • d • e

Oxidoreductases: alcohol oxidoreductases (EC 1.1)

1.1.1 NAD/NADP acceptor

Carbohydrate dehydrogenases - Alcohol dehydrogenase - Glycerol-3-phosphate dehydrogenase - L-xylulose reductase - Aldose reductase - Lactate dehydrogenase - 3-Hydroxyacyl CoA dehydrogenase - Malate dehydrogenase - Isocitrate dehydrogenase - Phosphogluconate dehydrogenase - Glucose-6-phosphate dehydrogenase - HMG-CoA reductase - Β-Ketoacyl ACP reductase - Hydroxysteroid dehydrogenase: 11 Beta (HSD11B1, HSD11B2) - 3 Beta (3-beta-HSD) - 17 Beta - Carnitine dehydrogenase - Beta-hydroxybutyryl-CoA dehydrogenase - IMP dehydrogenase - DXP reductoisomerase

1.1.3 oxygen acceptor

Glucose oxidase - L-gulonolactone oxidase - Xanthine oxidase

v • d • e Aldehyde/oxo oxidoreductases (EC 1.2)
1.2.1 - NAD or NADP acceptor	Aldehyde dehydrogenase - Acetaldehyde dehydrogenase (ALDH2) - Glyceraldehyde 3-phosphate dehydrogenase - Long-chain-aldehyde dehydrogenase
1.2.4 - disulfide acceptor	Oxoglutarate dehydrogenase - Pyruvate dehydrogenase - Branched-chain alpha-keto acid dehydrogenase complex

v • d • e Oxidoreductases: CH-CH oxidoreductases (EC 1.3)
1.3.1. NAD/NADP acceptor	Enoyl-acyl carrier protein reductase/Enoyl ACP reductase - 7-Dehydrocholesterol reductase - Biliverdin reductase - 2,4 Dienoyl-CoA reductase
1.3.3. Oxygen acceptor	Dihydroorotate dehydrogenase - Coproporphyrinogen III oxidase - Protoporphyrinogen oxidase -
1.3.5. Quinone	Succinate - coenzyme Q reductase
1.3.99. Other acceptors	Fumarate reductase - Butyryl CoA dehydrogenase - Acyl CoA dehydrogenase - 5-alpha reductase - Glutaryl-CoA dehydrogenase - Isovaleryl coenzyme A dehydrogenase

v • d • e CH-NH₂ oxidoreductases (EC 1.4) - primarily amino acid oxidoreductases
1.4.1 - NAD/NADP acceptor	Glutamate dehydrogenase (GLUD1)
1.4.3 - oxygen acceptor	D-amino acid oxidase - Amine oxidase - Lysyl oxidase - Monoamine oxidase
1.4.4 - disulfide acceptor	Glycine decarboxylase complex
1.4.99 - other acceptors	D-amino acid dehydrogenase - Amine dehydrogenase

v • d • e CH-NH oxidoreductases (EC 1.5)
1.5.1 - NAD or NADP acceptor	Dihydrofolate reductase - Saccharopine dehydrogenase - Methylenetetrahydrofolate reductase
1.5.3 - oxygen acceptor	Dihydrobenzophenanthridine oxidase - Sarcosine oxidase
1.5.5 - quinone acceptor	Electron-transferring-flavoprotein dehydrogenase

v • d • e NADH or NADPH oxidoreductases (EC 1.6)
Methemoglobin reductase - NADPH oxidase (P91-PHOX) - NADH dehydrogenase

v • d • e Oxidoreductases: nitrogenous donor (EC 1.7)
GMP reductase - Urate oxidase

v • d • e Oxidoreductases: sulfur oxidoreductases (EC 1.8)
Dihydrolipoamide dehydrogenase - Glutathione reductase - Thioredoxin reductase - Sulfite oxidase

v • d • e Diphenol family oxidoreductases (EC 1.10)
Coenzyme Q - cytochrome c reductase - Catechol oxidase - Laccase - Alternative oxidase

v • d • e Oxidoreductases: peroxidases (EC 1.11)
Catalase - Cytochrome c peroxidase - Eosinophil peroxidase - Glutathione peroxidase - Horseradish peroxidase - Lactoperoxidase - Myeloperoxidase - Thyroid peroxidase - Deiodinase (Tetraiodothyronine 5' deiodinase)

v • d • e Oxidoreductases: monooxygenases (EC 1.13)
1.13.11 - two atoms of oxygen	Catechol dioxygenase - Homogentisate 1,2-dioxygenase - Cysteine dioxygenase - 4-Hydroxyphenylpyruvate dioxygenase - Indoleamine 2,3-dioxygenase - Lipoxygenase (5)
1.13.12 - one atom of oxygen	Firefly luciferase
1.13.99 - other	Inositol oxygenase

v • d • e Oxidoreductases: dioxygenases, including steroid hydroxylases (EC 1.14)
1.14.11 - 2-oxoglutarate	Prolyl hydroxylase - Lysyl hydroxylase
1.14.13 - NADH or NADPH	Flavin-containing monooxygenase - Nitric oxide synthase - Cholesterol 7 alpha-hydroxylase - Methane monooxygenase - 3A4 -51A1
1.14.14 - reduced flavin or flavoprotein	19A1 - 2D6 - 2E1
1.14.15 - reduced iron-sulfur protein	11B1 - 11B2 - 11A1
1.14.16 - reduced pteridine	Phenylalanine hydroxylase - Tyrosine hydroxylase - Tryptophan hydroxylase
1.14.17 - reduced ascorbate	Dopamine beta hydroxylase
1.14.18-19 - other	Tyrosinase - Stearoyl-CoA desaturase-1
1.14.99 - miscellaneous	Cyclooxygenase - Heme oxygenase (HMOX1) - Squalene monooxygenase - 17A1 - 21A2

v • d • e Other oxidoreductases (EC 1.15-1.18)
1.15 - Acting on superoxide as acceptor	Superoxide dismutase
1.16 - Oxidizing metal ions	Ceruloplasmin
1.17 - Acting on CH or CH2 groups	Xanthine oxidase - Ribonucleotide reductase
1.18 - Acting on iron-sulfur proteins as donors	Nitrogenase

v • d • e Transferase: one carbon transferases (EC 2.1)
2.1.1 - Methyltransferases	5-hydroxyindole-O-methyltransferase/Acetylserotonin O-methyltransferase - Betaine-homocysteine methyltransferase - Catechol-O-methyl transferase - Histamine N-methyltransferase - Homocysteine methyltransferase - 5-Methyltetrahydrofolate-homocysteine methyltransferase - Phosphatidyl ethanolamine methyltransferase - Phenylethanolamine N-methyltransferase - DNMT3B - Histone methyltransferase - Thymidylate synthase - Tryptamine N-methyltransferase - DNA methyltransferase
2.1.2 - Hydroxy methyl-, Formyl- and Related Transferases	Serine hydroxymethyltransferase - Phosphoribosylglycinamide formyltransferase - Inosine monophosphate synthase - Glutamate formimidoyltransferase - Aminomethyltransferase
2.1.3 - Carboxy- and Carbamoyl transferases	Ornithine transcarbamylase
2.1.4 - Amidino transferases	Arginine:glycine amidinotransferase

v • d • e Aldehyde-ketone transferases (EC 2.2)
Acetolactate synthase - Transaldolase - Transketolase

v • d • e Transferases: acyltransferases (EC 2.3)
2.3.1: other than amino-acyl groups (mostly acetyltransferases)	N-Acetylglutamate synthase - Choline acetyltransferase - Acetyl-Coenzyme A acetyltransferase - Dihydrolipoyl transacetylase - Acetyl-CoA C-acyltransferase - Beta-galactoside transacetylase - Carnitine O-palmitoyltransferase (CPT1, CPT2) - Acyltransferase like 2 - Chloramphenicol acetyltransferase - Aminolevulinic acid synthase - Beta-ketoacyl-ACP synthase - Glyceronephosphate O-acyltransferase - Lecithin-cholesterol acyltransferase - Histone acetyltransferase (P300/CBP) - Serotonin N-acetyl transferase
2.3.2 - Aminoacyltransferases	Gamma glutamyl transpeptidase - Peptidyl transferase - Transglutaminase (Tissue transglutaminase, Keratinocyte transglutaminase, Factor XIII)
2.3.3 - converted into alkyl on transfer	Citrate synthase - ATP citrate lyase - HMG-CoA synthase

v • d • e Transferases: glycosyltransferases (EC 2.4)
2.4.1 - Hexosyltransferases: glucosyltransferase	Phosphorylase (Starch, Glycogen, Myo-) - Glycogen synthase - Debranching enzyme - Branching enzyme - 1,3-beta-glucan synthase - Ceramide glucosyltransferase
2.4.1 - Hexosyltransferases: other	Galactosyltransferase (Lactose synthase, B-N-acetylglucosaminyl-glycopeptide b-1,4-galactosyltransferase) - Glucuronosyltransferase (UGT1A1, UGT2B7, Hyaluronan synthase) - Fucosyltransferase (POFUT1, POFUT2)
2.4.2 - Pentosyltransferases: ADP ribose	Cholera toxin - Diphtheria toxin - Pertussis toxin - Poly ADP ribose polymerase
2.4.2 - Pentosyltransferases: other	Purine nucleoside phosphorylase - Adenine phosphoribosyltransferase - Hypoxanthine-guanine phosphoribosyltransferase - Uracil phosphoribosyltransferase - Amidophosphoribosyltransferase

v • d • e Alkyl and aryl transferases (EC 2.5)
Dimethylallyltranstransferase - Thiaminase I - Methionine adenosyltransferase - Riboflavin synthase - Dihydropteroate synthetase - Spermidine synthase - Glutathione S-transferase - Farnesyl-diphosphate farnesyltransferase - Spermine synthase - Alkylglycerone phosphate synthase - Farnesyltransferase - Geranylgeranyltransferase type 1 - Porphobilinogen deaminase

v • d • e Transferase: transaminases (EC 2.6)
Aspartate transaminase - Alanine transaminase - GABA transaminase - Tyrosine aminotransferase - Ornithine aminotransferase - Branched chain aminotransferase

v • d • e Phosphotransferases/kinases (EC 2.7)
2.7.1 - OH acceptor	Hexo- - Gluco- - Fructo- (Hepatic fructo-) - Galacto- - Phosphofructo- (1, 2) - Thymidine - NAD+ - Glycerol - Pantothenate - Mevalonate - Pyruvate - Deoxycytidine - PFP - Diacylglycerol - Bruton's tyrosine - Phosphoinositide 3 (Class I PI 3, Class II PI 3) - Sphingosine
2.7.2 - COOH acceptor	Phosphoglycerate - Aspartate
2.7.3 - N acceptor	Creatine
2.7.4 - PO4 acceptor	Phosphomevalonate - Adenylate - Nucleoside-diphosphate
2.7.6 - P2O7	Ribose-phosphate diphosphokinase - Thiamine pyrophosphokinase
2.7.7 - nucleotidyl-	Integrase - PNPase - Polymerase - RNase PH - UDP-glucose pyrophosphorylase - Galactose-1-phosphate uridylyltransferase -Terminal deoxynucleotidyl transferase - RNA replicase - Reverse transcriptase (Telomerase) - Transposase
2.7.8 - other phos.	N-acetylglucosamine-1-phosphate transferase
2.7.10-11 - protein	Tyrosine - Serine/threonine-specific

v • d • e Sulfur-containing group transferases (EC 2.8)
Alcohol sulfotransferase - Tyrosylprotein sulfotransferase

v • d • e Hydrolase: esterases (EC 3.1)
3.1.1: Carboxylic ester hydrolases	Cholinesterase - Pectinesterase - 6-phosphogluconolactonase - PAF acetylhydrolase Lipase (Gastric/Lingual, Pancreatic, Lysosomal, Hormone-sensitive, Endothelial, Hepatic, Lipoprotein, Monoacylglycerol, Diacylglycerol) Phospholipase (A1, A2, B)
3.1.2: Thioesterase	Palmitoyl thioesterase - Ubiquitin carboxy-terminal hydrolase L1
3.1.3: Phosphatase	Alkaline phosphatase - Acid phosphatase (Prostatic)/Tartrate resistant acid phosphatase/Purple acid phosphatases - Nucleotidase - Glucose 6-phosphatase - Fructose 1,6-bisphosphatase - Calcineurin - Phosphoprotein phosphatase (PP2A) - OCRL - Pyruvate dehydrogenase phosphatase - Fructose 2,6-bisphosphatase - Protein tyrosine phosphatase - PTEN
3.1.4: Phosphodiesterase	Autotaxin - Phospholipase (C, D) - Sphingomyelin phosphodiesterase - PDE1 - PDE2 - PDE3 - PDE5
3.1.6: Sulfatase	Arylsulfatase B - Steroid sulfatase - Galactosamine-6 sulfatase - Arylsulfatase A - Iduronate-2-sulfatase - N-acetylglucosamine-6-sulfatase
other	Nuclease

v • d • e

Hydrolase: sugar hydrolases (EC 3.2)

3.2.1: Glycoside hydrolases

Amylase (Alpha-Amylase) - Chitinase - Lysozyme - Neuraminidase - Galactosidases (Alpha, Beta) - alpha-Mannosidase - Glucuronidase - Hyaluronidase - Pullulanase - Glucocerebrosidase - Galactosylceramidase - Alpha-N-acetylglucosaminidase - Fucosidase - Hexosaminidase - Iduronidase - Disaccharidase (Sucrase/Sucrase-isomaltase/Invertase, Maltase, Trehalase, Lactase) - Glucosidases (Cellulase, Alpha-glucosidase, Beta-glucosidase, Debranching enzyme)

3.2.2: Hydrolysing N-Glycosyl compounds

DNA glycosylases: Oxoguanine glycosylase

v • d • e Ether bond hydrolases (EC 3.3)
Adenosylhomocysteinase - Epoxide hydrolase - Leukotriene A4 hydrolase

v • d • e Hydrolase: proteases (EC 3.4)
Exopeptidase 3.4.11-19	Angiotensin-converting enzyme - Dipeptidase - Dipeptidyl peptidase-4 - DD-transpeptidase Metalloexopeptidases: Aminopeptidase (Alanine, Cystinyl, Leucyl, Glutamyl) - Carboxypeptidase (A, B, C, E, Glutamate II)
Endopeptidase 3.4.21-24	Serine proteases - Cysteine protease - Aspartic acid protease - Metalloendopeptidases
Cathepsin 3.4.18,21,22,23	A - B - C - K

v • d • e Carbon-nitrogen non-peptide hydrolases (EC 3.5)
3.5.1 - Amidohydrolases	Asparaginase - Glutaminase - Urease - Biotinidase - Aspartoacylase - Ceramidase -Aspartylglucosaminidase - Fatty acid amide hydrolase - Histone deacetylase (Sirtuin)
3.5.2	Barbiturase - Beta-lactamase
3.5.3	Arginase
3.5.4 - Aminohydrolases	Guanine deaminase - Adenosine deaminase - AMP deaminase - Inosine monophosphate synthase - DCMP deaminase - GTP cyclohydrolase I
Other	Nitrilase - Thiaminase II

v • d • e Hydrolases: acid anhydride hydrolases (EC 3.6)
3.6.1	Pyrophosphatase (Inorganic, Thiamine) - Apyrase - Thiamine triphosphatase
3.6.3-4	ATPase
3.6.5	GTPase

v • d • e Carbon-carbon hydrolases (EC 3.7)
Fumarylacetoacetate hydrolase - Kynureninase

v • d • e Carbon-carbon lyases (EC 4.1)
4.1.1: Carboxy-lyases	Pyruvate decarboxylase -Oxaloacetate decarboxylase - Acetoacetate decarboxylase - Malonyl-CoA decarboxylase - Glutamate decarboxylase - Ornithine decarboxylase - Lysine decarboxylase - Phosphoribosylaminoimidazole carboxylase - Histidine decarboxylase - Uridine monophosphate synthetase/Orotidine 5'-phosphate decarboxylase - Aromatic L-amino acid decarboxylase - Phosphoenolpyruvate carboxylase - Pyrophosphomevalonate decarboxylase - Uroporphyrinogen III decarboxylase - RuBisCO - Phosphoenolpyruvate carboxykinase - Adenosylmethionine decarboxylase
4.1.2: Aldehyde-lyases	Aldolase A/Aldolase B - 2-hydroxyphytanoyl-CoA lyase
4.1.3: Oxo-acid-lyases	3-hydroxy-3-methylglutaryl-CoA lyase

v • d • e Carbon-oxygen lyases (EC 4.2) (primarily dehydratases)
Carbonic anhydrase - Fumarase - Aconitase - Enolase (Alpha) - Enoyl-CoA hydratase/3-Hydroxyacyl ACP dehydrase - Methylglutaconyl-CoA hydratase - Tryptophan synthase - Cystathionine beta synthase - Porphobilinogen synthase - 3-isopropylmalate dehydratase - Urocanate hydratase - Uroporphyrinogen III synthase - Nitrile hydratase

v • d • e Carbon-nitrogen lyases (EC 4.3)
4.3.1 - ammonia-lyases	Histidine ammonia-lyase - Formiminotransferase cyclodeaminase - Serine dehydratase
4.3.2 - amidine-lyases	Argininosuccinate lyase - Adenylosuccinate lyase

v • d • e Carbon-sulfur lyases (EC 4.4)
Cystathionine gamma-lyase - Cystathionine-beta-lyase - Leukotriene C4 synthase

v • d • e Carbon-halide lyases (EC 4.5)
Dichloromethane dehalogenase - Halohydrin dehalogenase

v • d • e Phosphorus-oxygen lyases (EC 4.6)
Adenylate cyclase - Guanylate cyclase (2D, c)

v • d • e Isomerases: Epimerase and racemases (EC 5.1)
Phenylalanine racemase (ATP-hydrolysing) - Serine racemase - Mandelate racemase - UDP galactose epimerase - Methylmalonyl CoA epimerase

v • d • e Geometric isomerases (EC 5.2)
Cyclophilin - FKBP - Parvulin - Prolyl isomerase

v • d • e Intramolecular oxidoreductases isomerases (EC 5.3)
5.3.1 - Aldoses/Ketoses	Triosephosphate isomerase - Phosphopentose isomerase - Mannose phosphate isomerase - Glucose isomerase
5.3.3 - C=C	Isopentenyl-diphosphate delta isomerase - Enoyl CoA isomerase
5.3.4 - S-S	Protein disulfide isomerase (PDIA3)
5.3.99 - other	Prostaglandin D2 synthase - Prostaglandin E synthase - Prostacyclin synthase - Thromboxane-A synthase

v • d • e Isomerase: mutases (EC 5.4)
5.4.2 Phosphomutases	Phosphoglycerate mutase - Bisphosphoglycerate mutase - Phosphoglucomutase
5.4.99 Other groups	Methylmalonyl-CoA mutase - Lanosterol synthase

v • d • e Intramolecular lyases (EC 5.5)
3-carboxy-cis,cis-muconate_cycloisomerase - Alpha-pinene-oxide_decyclase - Bornyl_diphosphate_synthase
Carboxy-cis,cis-muconate_cyclase - Chalcone_isomerase - Chloromuconate_cycloisomerase
Copalyl_diphosphate_synthase - Cycloeucalenol_cycloisomerase - Dichloromuconate_cycloisomerase
Ent-copalyl_diphosphate_synthase - Inositol-3-phosphate_synthase - Tetrahydroxypteridine_cycloisomerase

v • d • e Isomerase: topoisomerases (EC 5.99)
Type I topoisomerase - Type II topoisomerase (gyrase, topoisomerase IV)

v • d • e Enzymes: ligases (EC 6)
6.1 - Carbon-Oxygen	Aminoacyl tRNA synthetase
6.2 - Carbon-Sulfur	Succinyl coenzyme A synthetase - Acetyl Co-A synthetase - Long fatty acyl CoA synthetase
6.3 - Carbon-Nitrogen	Glutamine synthetase - Ubiquitin ligase (Von Hippel-Lindau tumor suppressor, UBE3A, Mdm2, Anaphase-promoting complex) - Glutathione synthetase - CTP synthase - Adenylosuccinate synthase - Argininosuccinate synthetase - Holocarboxylase synthetase - GMP synthase - Asparagine synthetase - Carbamoyl phosphate synthase (I , II )
6.4 - Carbon-Carbon	Carbon-carbon ligases
6.5 - Phosphoric Ester	DNA ligase

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