CAZy - GH

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 IUBMB 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,

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

(iv) illustrates the difficulty of deriving relationships between family membership and substrate specificity

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 below.

Catalytic Mechanism

In most cases, the hydrolysis of the glycosidic bond is catalyzed by two amino acid 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 two families of glycosidases utilizing NAD+ as a cofactor [6] [7].

For a detailed description of the catalytic mechanism (and variations thereof) of glycoside hydrolases, please visit CAZypedia.

Tables for Direct Access

GH 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	116	117	118	119	120
121	122	123	124	125	126	127	128	129	130	131	132	133	134	135	136	137	138	139	140	141	142	143	144	145	146	147	148	149	150	151	152	153	154	155	156	157	158	159	160
161	162	163	164	165	166	167	168

Non-Classified Sequences

GH SubFamily Number

GH5	1 2 4 5 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

GH Classification Statistics

Modules in present families	874712
Non-Classified modules	17797

GH Clans of Related Families

GH-A	(β/α)₈	1 2 5 10 17 26 30 35 39 42 50 51 53 59 72 79 86 113 128 147 148 157 158 167

GH-B	β-jelly roll	7 16

GH-C	β-jelly roll	11 12

GH-D	(β/α)₈	27 31 36

GH-E	6-fold β-propeller	33 34 83 93

GH-F	5-fold β-propeller	43 62 117

GH-G	(α/α)₆	37 63 100 125

GH-H	(β/α)₈	13 70 77

GH-I	α+β	24 80

GH-J	5-fold β-propeller	32 68

GH-K	(β/α)₈	18 20 85

GH-L	(α/α)₆	15 65

GH-M	(α/α)₆	8 48

GH-N	β-helix	28 49

GH-O	(α/α)₆	52 116

GH-P	(α/α)₆	127 146

GH-Q	(α/α)₆	94 149 161

GH-R	(β/α)₈	29 107

EC Activities found in GH families

Caution : Because of the modular nature of CAZymes, these activities may not be directly associated with the family but simply borne by adjacent modules.

1.14.99.53	18

2.4.1.-	1 3 5 13 16 17 20 27 31 32 33 35 39 65 70 72 94 130 NC

2.4.1.2	15

2.4.1.4	13

2.4.1.5	70

2.4.1.7	13

2.4.1.8	65

2.4.1.9	68

2.4.1.10	32 68

2.4.1.12	6

2.4.1.18	13 57

2.4.1.19	13

2.4.1.20	94

2.4.1.25	13 57 77 133

2.4.1.30	149

2.4.1.31	94

2.4.1.49	94

2.4.1.64	65

2.4.1.67	36

2.4.1.82	36

2.4.1.97	149 161

2.4.1.99	32

2.4.1.100	32

2.4.1.140	70

2.4.1.161	13 31

2.4.1.183	13

2.4.1.207	16

2.4.1.211	112

2.4.1.216	65

2.4.1.230	65

2.4.1.243	32

2.4.1.247	112

2.4.1.248	66

2.4.1.279	65

2.4.1.281	130

2.4.1.282	65

2.4.1.319	130

2.4.1.320	130

2.4.1.321	94

2.4.1.329	13

2.4.1.333	94 NC

2.4.1.334	65

2.4.1.339	130

2.4.1.340	130

2.4.1.352	13

2.4.1.359	13

2.4.2.-	10

2.4.99.16	13

3.1.1.-	5 106

3.1.1.72	5 11 62

3.1.1.73	10 62 78

3.2.1.-	1 2 3 5 6 8 9 10 11 13 16 17 18 19 20 26 28 30 32 33 35 38 39 42 43 44 48

3.2.1.1	13 14 57 119

3.2.1.2	13 14

3.2.1.3	15 97

3.2.1.4	5 6 7 8 9 10 12 26 44 45 48 51 74 124 148 NC

3.2.1.6	9 16

3.2.1.7	32

3.2.1.8	3 5 6 8 9 10 11 16 26 30 43 44 51 62 98 141

3.2.1.10	13 31

3.2.1.11	31 49 66

3.2.1.14	18 19 23 48

3.2.1.15	28

3.2.1.17	18 19 22 23 24 25 73 108 NC

3.2.1.18	1 33 34 83 156 NC

3.2.1.20	4 13 31 63 76 97 122

3.2.1.21	1 2 3 5 9 16 30 39 116 NC

3.2.1.22	4 27 31 36 57 97 110

3.2.1.23	1 2 3 35 39 42 50 147 163 165 NC

3.2.1.24	31 38 92

3.2.1.25	1 2 5 113 164

3.2.1.26	32 68 100

3.2.1.28	13 15 37 65

3.2.1.31	1 2 30 79 137 154

3.2.1.32	8 26

3.2.1.33	13 133

3.2.1.35	16 56 84

3.2.1.36	79

3.2.1.37	1 2 3 5 10 30 39 43 51 52 54 116 120

3.2.1.38	1 3 30 35

3.2.1.39	5 16 17 55 64 81 128 152 157 158

3.2.1.40	33 78 106

3.2.1.41	13 57

3.2.1.45	3 5 30 116

3.2.1.46	5 59

3.2.1.48	13 31

3.2.1.49	27 31 36 109 129

3.2.1.50	89

3.2.1.51	29 141 151 NC

3.2.1.52	3 5 18 20 84 109 116

3.2.1.53	123

3.2.1.54	13 57

3.2.1.55	2 3 5 10 39 43 51 54 62

3.2.1.57	49

3.2.1.58	5 16 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	29 95

3.2.1.64	32

3.2.1.65	32

3.2.1.67	4 28

3.2.1.68	13

3.2.1.70	13 15

3.2.1.71	144 162

3.2.1.73	5 6 7 8 9 11 12 16 17 26 51

3.2.1.74	1 3 5 9

3.2.1.75	5 30

3.2.1.76	39

3.2.1.78	5 9 26 44 45 113 134

3.2.1.80	32

3.2.1.81	16 50 86 118

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

3.2.1.88	27 36 97

3.2.1.89	53 147

3.2.1.91	5 6 9

3.2.1.93	13

3.2.1.94	27

3.2.1.95	49

3.2.1.96	18 20 73 85

3.2.1.97	101

3.2.1.98	13

3.2.1.99	43

3.2.1.100	5 26

3.2.1.101	76

3.2.1.102	98

3.2.1.103	16

3.2.1.104	5

3.2.1.105	1

3.2.1.106	63

3.2.1.107	65

3.2.1.108	1

3.2.1.109	114

3.2.1.111	29

3.2.1.113	38 47 92

3.2.1.114	38 92

3.2.1.116	13

3.2.1.117	1

3.2.1.118	1

3.2.1.119	1

3.2.1.120	3

3.2.1.122	4

3.2.1.123	5

3.2.1.125	1

3.2.1.126	3

3.2.1.129	58

3.2.1.130	99

3.2.1.131	67 115

3.2.1.132	3 5 7 8 46 75 80

3.2.1.133	13

3.2.1.135	13

3.2.1.136	30

3.2.1.139	4 67

3.2.1.140	20 136

3.2.1.141	13

3.2.1.145	43

3.2.1.146	2 5 43 117 159

3.2.1.147	1

3.2.1.149	1 5

3.2.1.150	74

3.2.1.151	5 9 12 16 26 44 45 48 74

3.2.1.152	2

3.2.1.153	32

3.2.1.154	32

3.2.1.155	3

3.2.1.156	8

3.2.1.157	82

3.2.1.158	96

3.2.1.159	117

3.2.1.161	1

3.2.1.162	150

3.2.1.164	5 30

3.2.1.165	2 9 35

3.2.1.166	79

3.2.1.167	79

3.2.1.169	84

3.2.1.170	63

3.2.1.171	28

3.2.1.172	105

3.2.1.173	138

3.2.1.174	78 106

3.2.1.175	1 17

3.2.1.176	7 9 48

3.2.1.177	31

3.2.1.178	16 86

3.2.1.181	16

3.2.1.182	1

3.2.1.185	2 127 137 142 143 146

3.2.1.197	130

3.2.1.198	99

3.2.1.199	31

3.2.1.204	31

3.2.1.205	15

3.2.1.208	63

3.2.1.211	168

3.2.1.212	107

3.2.1.17_or_4.2.2.n1	23

3.5.1.-	153

3.5.2.17	1

4.2.2.n1	23 102 103 104 NC

4.2.2.13	31

4.2.2.15	33

4.2.2.16	32

4.2.2.17	91

4.2.2.18	91

5.4.99.11	13

5.4.99.15	13

5.4.99.16	13

Glycoside Hydrolase family classification

Introduction

Catalytic Mechanism

Tables for Direct Access

Note

Notes

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	116	117	118	119	120
121	122	123	124	125	126	127	128	129	130	131	132	133	134	135	136	137	138	139	140	141	142	143	144	145	146	147	148	149	150	151	152	153	154	155	156	157	158	159	160
161	162	163	164	165	166	167	168

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	116	117	118	119	120
121	122	123	124	125	126	127	128	129	130	131	132	133	134	135	136	137	138	139	140	141	142	143	144	145	146	147	148	149	150	151	152	153	154	155	156	157	158	159	160
161	162	163	164	165	166	167	168

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	116	117	118	119	120
121	122	123	124	125	126	127	128	129	130	131	132	133	134	135	136	137	138	139	140	141	142	143	144	145	146	147	148	149	150	151	152	153	154	155	156	157	158	159	160
161	162	163	164	165	166	167	168