CAZy Family Glycoside Hydrolase

CAZy ~ Carbohydrate-Active enZymes

Families

Glycoside Hydrolases

GlycosylTransferases

Polysaccharide Lyases

Carbohydrate Esterases

Carbohydrate-Binding Modules

Welcome to the Glycoside Hydrolase family server

Introduction

Glycoside hydrolases (EC 3.2.1.-) are a widespread group of enzymes which hydrolyse the glycosidic bond between two or more carbohydrates or between a carbohydrate and a non-carbohydrate moiety. The IUB-MB Enzyme nomenclature of glycoside hydrolases is based on their substrate specificity and occasionally on their molecular mechanism; such a classification does not reflect (and was not intended to) the structural features of these enzymes. A classification of glycoside hydrolases in families based on amino acid sequence similarities has been proposed a few years ago. Because there is a direct relationship between sequence and folding similarities, such a classification:

(i) reflects the structural features of these enzymes better than their sole substrate specificity,

(ii) helps to reveal the evolutionary relationships between these enzymes, and

(iii) provides a convenient tool to derive mechanistic information[1-2].

The Carbohydrate-Active Enzymes database (CAZy) provides a continuously updated list of the glycoside hydrolase families. Because the fold of proteins is better conserved than their sequences, some of the families can be grouped in 'clans'

(i) when new sequences are found to be related to more than one family,

(ii) when the sensitivity of sequence comparison methods is increased or

(iii) when structural determinations demonstrate the resemblance between members of different families [3].

The established 'clans' are given in form of a table.

Catalytic Mechanism

In most cases, the hydrolysis of the glycosidic bond is performed by two catalytic residues of the enzyme: a general acid (proton donor) and a nucleophile/base [4]. Depending on the spatial position of these catalytic residues, hydrolysis occurs via overall retention or overall inversion of the anomeric configuration. For each family listed in the CAZy database, we indicate (when it is known) the stereochemical outcome of the reaction catalyzed as well as the type of amino-acid residues acting as a nucleophile/base and as a proton donor. In some cases, the catalytic nucleophile is not borne by the enzyme, and is replaced by the acetamido group at C-2 of the substrate [5]. A completely unrelated mechanism has been demonstrated recently for a family of glycosidases utilizing NAD+ as a cofactor [6].

Tables for Direct Access

Family Number

1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22	23	24	25	26	27	28	29	30

31	32	33	34	35	36	37	38	39	40	41	42	43	44	45	46	47	48	49	50	51	52	53	54	55	56	57	58	59	60

61	62	63	64	65	66	67	68	69	70	71	72	73	74	75	76	77	78	79	80	81	82	83	84	85	86	87	88	89	90

91	92	93	94	95	96	97	98	99	100	101	102	103	104	105	106	107	108	109	110	111	112	113	114	115

* Non-Classified Sequences


Clans of Related Families

GH-A	Fold ( / ) ₈	1	2	5	10	17	26	30	35	39	42	50	51	53	59	72	79	86	113

GH-B	Fold -jelly roll	7	16

GH-C	Fold -jelly roll	11	12

GH-D	Fold ( / ) ₈	27	31	36

GH-E	Fold 6-fold -propeller	33	34	83	93

GH-F	Fold 5-fold -propeller	43	62

GH-G	Fold ( / ) ₆	37	63

GH-H	Fold ( / ) ₈	13	70	77

GH-I	Fold +	24	46	80

GH-J	Fold 5-fold -propeller	32	68

GH-K	Fold ( / ) ₈	18	20	85

GH-L	Fold ( / ) ₆	15	65

GH-M	Fold ( / ) ₆	8	48

GH-N	Fold -helix	28	49

EC Number


2.4.1.4	13

2.4.1.5	70

2.4.1.7	13

2.4.1.8	13	65

2.4.1.9	68

2.4.1.10	68

2.4.1.18	13	57

2.4.1.19	13

2.4.1.20	94

2.4.1.25	13	57	77

2.4.1.49	94

2.4.1.64	65

2.4.1.67	36

2.4.1.82	36

2.4.1.99	32

2.4.1.100	32

2.4.1.140	70

2.4.1.183	13

2.4.1.207	12	16

2.4.1.211	112

2.4.1.230	65

2.4.1.243	32

2.4.1.-	16	17	31	32	33	66	70	72	94	112

3.2.1.1	13	57

3.2.1.2	14

3.2.1.3	15

3.2.1.4	5	6	7	8	9	12	44	45	48	51	61	74

3.2.1.6	16

3.2.1.7	32

3.2.1.8	5	8	10	11	43

3.2.1.10	13

3.2.1.11	49	66

3.2.1.14	18	19	48

3.2.1.15	28

3.2.1.17	22	23	24	25	73	108

3.2.1.18	33	34	83

3.2.1.20	4	13	31	63	97

3.2.1.21	1	3	9

3.2.1.22	4	27	36	57	110

3.2.1.23	1	2	35	42

3.2.1.24	38

3.2.1.25	1	2	5

3.2.1.26	32	68	100

3.2.1.28	15	37	65

3.2.1.31	1	2	79

3.2.1.32	10	26

3.2.1.33	13

3.2.1.35	56	69	84

3.2.1.37	3	30	39	43	51	52	54

3.2.1.38	1

3.2.1.39	16	17	55	64	81

3.2.1.40	28	78	106

3.2.1.41	13	57

3.2.1.45	30

3.2.1.46	59

3.2.1.48	31

3.2.1.49	27	36	109

3.2.1.50	89

3.2.1.51	29	95

3.2.1.52	3	20	84

3.2.1.53	95

3.2.1.54	13

3.2.1.55	3	43	51	54	62

3.2.1.57	49

3.2.1.58	3	5	17	55

3.2.1.59	71	87

3.2.1.60	13

3.2.1.61	87

3.2.1.62	1

3.2.1.63	95

3.2.1.64	32

3.2.1.65	32

3.2.1.67	28

3.2.1.68	13

3.2.1.70	13	15

3.2.1.73	5	8	12	16	17

3.2.1.74	3

3.2.1.75	5	30

3.2.1.76	39

3.2.1.78	5	26	113

3.2.1.80	32

3.2.1.81	16	50	86

3.2.1.82	28

3.2.1.83	16

3.2.1.84	31	63

3.2.1.85	1

3.2.1.86	1	4

3.2.1.89	53

3.2.1.91	5	6	9	48

3.2.1.93	13

3.2.1.94	27

3.2.1.95	49

3.2.1.96	18	73	85

3.2.1.97	101

3.2.1.98	13

3.2.1.99	43

3.2.1.101	76

3.2.1.103	16

3.2.1.105	1

3.2.1.106	63

3.2.1.108	1

3.2.1.109	114

3.2.1.113	47

3.2.1.114	38

3.2.1.116	13

3.2.1.117	1

3.2.1.118	1

3.2.1.122	4

3.2.1.123	5

3.2.1.125	1

3.2.1.129	58

3.2.1.130	99

3.2.1.131	67

3.2.1.132	5	7	8	18	46	75	80

3.2.1.133	13

3.2.1.135	13

3.2.1.136	5

3.2.1.139	4	67

3.2.1.140	20

3.2.1.141	13

3.2.1.145	43

3.2.1.147	1

3.2.1.149	1

3.2.1.150	74

3.2.1.151	5	12	16	44	74

3.2.1.152	2

3.2.1.153	32

3.2.1.154	32

3.2.1.156	8

3.2.1.157	82

3.2.1.158	96

3.2.1.161	1

3.2.1.165	2	35

3.2.1.-	1	2	3	4	5	6	7	8	9	10	12	13	16	17	18	19	20	23	26	27	28	30	31	35	36	39	43	48	51
	56	61	63	66	73	79	88	90	92	93	97	98	102	103	104	105	107	110

3.5.2.17	1

4.2.2.13	31

4.2.2.16	32

4.2.2.-	23

5.4.99.11	13

5.4.99.15	13

5.4.99.16	13

5.4.99.16	16

5.4.99.16	18

Note

Some glycoside hydrolases are multifunctional enzymes that contain catalytic domains that belong to different GH families. The GenBank, SWISS-PROT, and/or PDB entries for these proteins are therefore mentioned in each of the relevant list of family members.

References


[1]	Henrissat B (1991) A classification of glycosyl hydrolases based on amino-acid sequence similarities. Biochem. J. 280:309-316 [PMID: 1747104].

[2]	Henrissat B, Bairoch A (1993) New families in the classification of glycosyl hydrolases based on amino- acid sequence similarities. Biochem. J. 293:781-788 [PMID: 8352747].

[3]	Henrissat B, Bairoch A (1996) Updating the sequence-based classification of glycosyl hydrolases. Biochem. J. 316:695-696 [PMID: 8687420].

[4]	Davies G, Henrissat B (1995) Structures and mechanisms of glycosyl hydrolases. Structure 3:853-859 [PMID: 8535779].

[5]	Terwisscha van Scheltinga A, Armand S, Kalk KH, Isogai A, Henrissat B, Dijkstra BW (1995) Stereochemistry of chitin hydrolysis by a plant chitinase/lysozyme and x-ray structure of a complex with allosamidin. Evidence for substrate assisted catalysis. Biochemistry 34:15619-15623 [PMID: 7495789].

[6]	Rajan SS, Yang X, Collart F, Yip VL, Withers SG, Varrot A, Thompson J, Davies GJ, Anderson WF (2004) Novel catalytic mechanism of glycoside hydrolysis based on the structure of an NAD+/Mn2+ -dependent phospho-alpha-glucosidase from Bacillus subtilis. Structure. 12:1619-1629 [PMID: 15341727].