Activities in Family	β-galactosidase (EC 3.2.1.23) ; β-mannosidase (EC 3.2.1.25); β-glucuronidase (EC 3.2.1.31); α-L-arabinofuranosidase (EC 3.2.1.55); mannosylglycoprotein endo-β-mannosidase (EC 3.2.1.152); exo-β-glucosaminidase (EC 3.2.1.165); α-L-arabinopyranosidase (EC 3.2.1.-); β-galacturonidase (EC 3.2.1.-); β-xylosidase (EC 3.2.1.37); β-D-galactofuranosidase (EC 3.2.1.146); β-glucosidase (EC 3.2.1.21); β-galacturonidase RGII specific (EC 3.2.1.-); glycyrrhizin β-glucuronidase (EC 3.2.1.128)
Mechanism	Retaining
Clan	GH-A
3D Structure Status	( β / α ) 8 barrel
Catalytic Nucleophile/Base	Glu (experimental)
Catalytic Proton Donor	Glu (experimental)
External resources	CAZypedia; HOMSTRAD; PRINTS; PROSITE;
Commercial Enzyme Provider(s)	MEGAZYME; PROZOMIX;
Statistics	GenBank accession (37668); Uniprot accession (238); PDB accession (227); 3D entries (59); cryst (2)

Bacteria
Protein Name	EC#	Organism	GenBank	Uniprot	PDB/3D Carbohydrate Ligands Resolution (Å)
exo-β-glucosaminidase / exochitosanase (CsxA)	3.2.1.165	Amycolatopsis orientalis	AAX62629.2 AAX62629.1	Q56F26	2VZO[A,B] 2.24 2VZS[A,B] β-D-GlcpN 1.85 2VZT[A,B] β-D-GlcpN-(1-1)-<non_carb> 2.20 2VZU[A,B] β-D-GlcpN-(1-1)-<non_carb> 2.10 2VZV[A,B] β-D-GlcpN-(1-4)-β-D-GlcpN 2.70 2X05[A,B] <carb_like_X05> 2.30 2X09[A,B] <carb_like_X09> 2.40
β-glucuronidase/β-galacturonidase (AqGalA)	3.2.1.31 3.2.1.-	Aquimarina sp. SCSIO21287	UER37076.1		7VQM[A,B,C,D] β-D-GalpA 2.40
β-galactosidase (Arth?DG)	3.2.1.23	Arthrobacter sp. 32cB	AHY00656.1	A0A023UGN9	6ETZ[A] 1.80 6H1P[A] 3.01 6SE8[A] 1.84 6SE9[A] β-D-Fruf β-D-Galp β-D-Galp-(1-4)-β-D-Glcp 1.97 6SEA[A] β-D-Galp β-D-Galp-(1-4)-β-D-Glcp 1.87 6SEB[A] β-D-Galp1S-(1-1)-<non_carb> 2.27 6SEC[A] β-D-Galp-(1-1)-<non_carb> 2.77 6SED[A] β-D-Galp 2.23 6ZJP[A] 1.85 6ZJQ[A] β-D-Galp 1.70 6ZJR[A] 2.00 6ZJS[A] β-D-Galp 1.50 6ZJT[A] β-D-Galp β-D-Galp-(1-4)-β-D-Fruf 1.97 6ZJU[A] α-D-Glcp-(1-2)-β-D-Fruf 1.75 6ZJV[A] 2.25 6ZJW[A] β-D-Galp 2.12 6ZJX[A] 2.21
β-galactosidase (LacZ)	3.2.1.23	Arthrobacter sp. C2-2	CAD29775.1	Q8KRF6	1YQ2[A,B,C,D,E,F] 1.90
BACCELL_00063		Bacteroides cellulosilyticus DSM 14838	EEF92278.1		6B6L[A,B] 2.00 7RSK[A,B] 2.40
BcellWH2_02662 (Cbga_1)		Bacteroides cellulosilyticus WH2	ALJ59901.1		7SF2[A,B,C,D,E,F] 2.75
BF0029		Bacteroides fragilis NCTC 9343	CAH05808.1		3FN9[A,B,C,D] 2.70
β-glucuronidase (BF0328)	3.2.1.31	Bacteroides fragilis NCTC 9343	CAH06099.1	Q5LIC7	3CMG[A] 1.90 5Z1A[A] 1.86
β-glucuronidase (BMSP;HMPREF2532_04191)	3.2.1.31	Bacteroides ovatus KLE1656	KXT42440.1	A0A139KTC9	6D8K[B,C,D] 2.65 6D8K[A] 2.65
β-mannosidase (BT0458;BT_0458) (Man2A)	3.2.1.25	Bacteroides thetaiotaomicron VPI-5482	AAO75565.1 NP_809371.1	Q8AAK6	2JE8[A,B] 1.7 2VJX[A,B] <carb_like_IFL> 1.85 2VL4[A,B] <carb_like_MNM> 1.90 2VMF[A,B] <carb_like_MVL> 2.1 2VO5[A,B] 2.3 2VOT[A,B] 1.95 2VQT[A,B] <carb_like_15A> 2.1 2VQU[A,B] <carb_like_NOY> 1.9 2VR4[A,B] <carb_like_17B> 1.8 2WBK[A,B] β-D-Manp2F-(1-1)-<non_carb> 2.10 7OP6[A,B] 2.05 7OP7[A,B] 1.85
BT3179		Bacteroides thetaiotaomicron VPI-5482	AAO78285.1 NP_812091.1		3DEC[A] 2.80
BT1626		Bacteroides thetaiotaomicron VPI-5482	AAO76733.1 NP_810539.1		3BGA[A,B] 2.10
exo-β-galactosidase porphyran-specific (BuGH2A)	3.2.1.23	Bacteroides uniformis NP1	WP_234130811.1	A0A2D0TCC8	5T98[A,B] 2.60 5T99[A,B] 2.40
agarolytic β-galactosidase (BuGH2C)	3.2.1.23	Bacteroides uniformis NP1		A0A2D0TCC9	5T9A[B,C,D] 2.50 5T9G[A,B,C,D] 2.40 5T9A[A] 2.50
β-glucuronidase (BuGUS-2; M094_3283)	3.2.1.31	Bacteroides uniformis str. 3978 T3 i	KDS62590.1	A0A078SUX9 A0A414F142	5UJ6[A,B] 1.90 6D50[A,B] β-D-GlcpA 2.50 6D8G[A,B] 3.00 6NZG[A,B] 2.43
β-glucuronidase (BuGUS-1; ERS417307_01040; Uida_4)	3.2.1.31	Bacteroides uniformis str. 3978 T3 i	CUO13985.1	A0A174CQK8	6D1N[A,B] 2.20 6D41[A,B] α-D-GlcpA β-D-GlcpA 1.90 6D6W[A,B,C,D] α-D-GlcpA 1.80 6D7F[A,B,C,D,E,F] β-D-GlcpA-(1-1)-<non_carb> 2.40 6D89[A,B,C,D] 2.00
β-glucuronidase (BuGUS-3; M094_3281)		Bacteroides uniformis str. 3978 T3 i	KDS64778.1		6D1P[A,B] 2.35
β-galactosidase (BIF3)	3.2.1.23	Bifidobacterium bifidum DSM 20215	CAC14566.2 AAP64961.1 ABI07182.1 CAD19366.1	Q9F4D5	6QUB[A,B] β-D-Galp 1.95 6QUC[A,B] 2.30 6QUD[A] α-D-2-deoxy-Galp-(1-4)-GLU 2.10 7NIT[A,B,C,D,E,F] 2.89
lacto-N-neotetraose β-galactosidase (BbgIII)	3.2.1.23	Bifidobacterium bifidum JCM 1254	BAI94821.1	D4QAP3	5DMY[A,B,C] 1.95
HMPREF0168_2111 (UidA)	3.2.1.31	Bifidobacterium dentium ATCC 27679	WP_003843201.1	E0QAF3	5Z1B[A,B,C,D] β-D-GlcpA-(1-1)-<non_carb> β-D-GlcpA-(1-2)-unknown 1.45
BDP_2112 (Lacz1)		Bifidobacterium dentium Bd1	ADB10673.1		6LD0[A,B,C,D] 1.90 6LD6[A,B,C,D] 2.20 6LDB[A,B,C,D] 1.65 6LDC[A,B,C,D] 2.18 6LDD[A,B,C,D] 2.21
β-glucuronidase (BglR;CPE0147)	3.2.1.31	Clostridium perfringens str. 13	BAB79853.1 CAD12654.1 NP_561063.1	Q8VNV4 Q8XP19	4JKM[A,B] 2.26 6CXS[A,B] 2.80
β-glucuronidase (DtGlcA; DICTH_1429)	3.2.1.31	Dictyoglomus thermophilum H-6-12	ACI18585.1	B5YFE0	6XXW[A] 1.85
β-mannosidase (DtMan;DICTH_1692)	3.2.1.25	Dictyoglomus thermophilum H-6-12	ACI19676.1	B5YAN4	5N6U[A,B,C,D] D-1-deoxy-Manp 3.08
β-galacturonidase (EtGalAse)	3.2.1.-	Eisenbergiella tayi	WP_069152983.1	A0A1E3AEY6	6NCW[A,B,C,D] 2.10 6NCX[A,B,C,D] α-D-GalpA 2.25
RG69_19155 (LacZ)		Escherichia coli S40	APL67431.1		6TSH[A,B,C,D] <carb_like_DGJ> 2.30 6TSK[A,B,C,D] β-L-Ribp 2.30 6TTE[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.20
HPE47_22190 (UidA)		Escherichia coli SCU-488	QKP93781.1		6LEG[A,B,C,D] 2.60 6LEJ[A,B] 2.62 6LEL[A,B] 2.50 6LEM[A,B] 3.19
β-galactosidase Z (LacZ;Lac-Z;b0344;EO53_16565)	3.2.1.23	Escherichia coli str. K-12 substr. MG1655	AAA24053.1 AAB18068.1 AAC73447.1 AAW06767.1 AAY73171.1 ABN72582.1 AEL16982.1 AIZ92504.1 ALI41840.1 AMC97588.1 AUG15134.1 AVI54877.1 CAA23573.1 CAC87491.1 NP_414878.1	B8LFD6 G0ZKW2 P00722 Q8VNN2	1BGL[A] 2.50 1BGM[I] 2.50 1DP0[A] 1.70 1F49[A] 2.50 1F4A[A] 2.80 1F4H[A] 2.80 1GHO[I] 2.50 1HN1[A,B,C,D] 3.00 1JYN[A] β-D-Galp-(1-4)-β-D-Glcp 1.80 1JYV[A] β-D-Galp-(1-1)-<non_carb> 1.75 1JYW[A] β-D-Galp-(1-1)-<non_carb> 1.55 1JYX[A] β-D-Galp1S-(1-1)-<non_carb> 1.75 1JYY[A] β-D-Galp2F-(1-4)-β-D-Glcp 2.70 1JYZ[I] β-D-Galp2F-(1-4)-β-D-Glcp 2.70 1JZ0[A] α-D-Galp2F-(1-4)-GLU 2.60 1JZ1[I] α-D-Galp2F-(1-4)-GLU 2.60 1JZ2[A] α-D-Galp2F-(1-4)-GLU 2.10 1JZ3[A] α-D-2-deoxy-Galp-(1-4)-GLU 1.75 1JZ4[A] α-D-2-deoxy-Galp-(1-4)-GLU 2.10 1JZ5[A] α-D-Galp β-D-Galp 1.80 1JZ6[A] <carb_like_GTZ> 2.10 1JZ7[A] β-D-Galp 1.50 1JZ8[A] β-D-Galp-(1-6)-β-D-Glcp β-D-Galp-(1-6)-β-D-Manp 1.50 1PX3[A,B,C,D] 1.60 1PX4[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 1.60 3CZJ[A,B,C,D] β-D-Galp 2.05 3DYM[A,B,C,D] 2.05 3DYO[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 1.80 3DYP[A,B,C,D] 1.75 3E1F[1,2,3,4] 3.00 3I3B[A,B,C,D] α-D-Galp β-D-Galp 2.20 3I3D[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.20 3I3E[A,B,C,D] 2.10 3IAP[A,B,C,D] 2.00 3IAQ[A,B,C,D] 2.70 3J7H[A,B,C,D] 3.20 3MUY[1,2,3,4] 2.50 3MUZ[1,2,3,4] 1.90 3MV0[1,2,3,4] 2.20 3MV1[1,2,3,4] 2.20 3SEP[A,B,C,D] 2.05 3T08[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.00 3T09[A,B,C,D] α-D-Galp β-D-Galp 1.75 3T0A[A,B,C,D] 1.90 3T0B[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.40 3T0D[A,B,C,D] α-D-Galp β-D-Galp 1.93 3T2O[A,B,C,D] 1.85 3T2P[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.60 3T2Q[A,B,C,D] α-D-Galp β-D-Galp 2.40 3VD3[A,B,C,D] 2.80 3VD4[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.00 3VD5[A,B,C,D] 2.70 3VD7[A,B,C,D] <carb_like_GTZ> 2.87 3VD9[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.05 3VDA[A,B,C,D] 2.50 3VDB[A,B,C,D] α-D-Galp β-D-Galp 2.05 3VDC[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.55 4CKD[A,B,C,D] 13.00 4DUV[A,B,C,D] α-D-2-deoxy-Galp-(1-4)-GLU 2.10 4DUW[A,B,C,D] β-D-Galp-(1-6)-β-D-Glcp 2.20 4DUX[A,B,C,D] β-L-Ribp 2.30 4TTG[A,B,C,D] 1.60 4V40[A,B,C,D,E,F,G,H,I,J, 4V41[A,B,C,D,E,F,G,H,I,J, 4V44[A,B,C,D,E,F,G,H,I,J, 4V45[A,B,C,D,E,F,G,H,I,J, 5A1A[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 2.20 6CVM[A,B,C,D] β-D-Galp1S-(1-1)-<non_carb> 1.90 6DRV[A,B,C,D] 2.20 6KUZ[A,B,C,D] β-D-Galp-(1-1)-<non_carb> 2.83 6X1Q[A,B,C,D] 1.80 7BRS[A,B,C,D] β-D-Galp-(1-1)-<non_carb> 2.67 7BTK[A,B,C,D] β-D-Galp-(1-1)-<non_carb> 2.70
β-glucuronidase (UidA;GusA;GurA;b1617)	3.2.1.31	Escherichia coli str. K-12 substr. MG1655	AAC74689.1 AAA68923.1 AAB30197.1 AAR75623.1 ABE20937.1 ACR88323.1 AIZ91278.1 ALI40610.1 AMC98822.1 AUG16387.1 AVI56132.1 AYG19328.1 CAJ55495.1 CAJ55496.1 CAJ55497.1 CAJ55498.1 CAJ55499.1 CAJ55500.1 NP_416134.1 WP_001551153.1	A0A0D8WFE1 A0A1X3I6P5 A0A7Z1D535 A9DEX9 A9DEY7 A9DEZ1 A9DEZ6 A9DF00 A9DF05 P05804	3K46[A,B] 2.50 3K4A[A,B] 2.90 3K4D[A,B] <carb_like_EVA> 2.39 3LPF[A,B] 2.26 3LPG[A,B] 2.42 4JHZ[A,B] 2.83 5CZK[A,B] 2.39 7PR6[A,B]
β-glucuronidase (EeGUS; EUBELI_20590)	3.2.1.31	[Eubacterium] eligens ATCC 27750	ACR73734.1	A0A174ZZA3 C4Z6Z2	6BJQ[A] β-D-GlcpA 2.70 6BJW[A] 3.00 6BO6[A] 2.80 6D4O[A] 2.90
ORF		Faecalibacterium prausnitzii			7KGY[A,B,C,D] 2.20
β-glucuronidase (CG447_12395)	3.2.1.31	Faecalibacterium prausnitzii A2165	ATP00638.1 EEU97254.1	C7H4D2	6ED2[A] 2.30
β-glucuronidase (C4Q21_10165)	3.2.1.31	Faecalibacterium prausnitzii APC918/95b	AXB29257.1	A0A3F3JX71	6U7I[A,B,C,D] 2.70
β-glucuronidase (Fp2GUS; FP2_04120)	3.2.1.31	Faecalibacterium prausnitzii L2-6 L2/6	CBK98066.1	D4K3H3	6MVF[A,B,C,D,E,F] 2.55
[ulvan] β-1,2-glucuronidase (BN863_22060)	3.2.1.31	Formosa agariphila KMM 3901	CDF79918.1	T2KN75	6HPD[A] 2.43
FsGUS (β-galacturonidase/β-glucuronidase)	3.2.1.31 3.2.1.-	Fusicatenibacter saccharivorans		A0A174EHD1	6NCY[B] 2.05 6NCZ[A,B,C,D,E,F] β-D-GlcpA-(1-1)-<non_carb> 2.20 6NCY[A] 2.05
β-glucuronidase (HMPREF0539_0313; UidA)	3.2.1.31	Lacticaseibacillus rhamnosus LMS2-1	EEN81625.1	A0A2A5L2J1 C2JTS9	6ECA[A,B] 2.85
β-galactosidase A / B / C / D / 91 (Bga;BgaD;β-Gal-A;β-Gal-B;β-Gal-C;β-Gal-D;β-Glα-91)	3.2.1.23	Niallia circulans ATCC 31382	BAJ61032.1	E5RWQ2	4YPJ[A,B] 2.50
β-galactosidase	3.2.1.23	Niallia circulans ATCC 31382	QJU69416.1	A0A6M5K904	7CWD[A] α-D-Galp-(1-4)-GLU α-D-Glcp 2.00 7CWI[A] 1.95
β-glucuronidase (ZP_02032394.1;PARMER_02407) (26901)	3.2.1.31	Parabacteroides merdae ATCC 43184	ZP_02032394.1	A7AG62 K5ZWV5	6D7J[A,B,C,D] 2.24 6DXU[A,B,C,D] 1.90
β-galactosidase (BgaL; ParbDG)	3.2.1.23	Paracoccus sp. 32d	ACY69080.1	D1LZK0	5EUV[A,B] 2.40 5LDR[A,B] α-D-Glcp β-D-Galp 3.15
β-glucuronidase (BSEG_01974)	3.2.1.31	Phocaeicola dorei 5_1_36/D4	EEO45833.1	C3R9X4	6ED1[A,B,C,D] 2.90
BVU_0391		Phocaeicola vulgatus ATCC 8482	ABR38109.1		3GM8[A] 2.40
β-glucuronidase (Rh2GUS)	3.2.1.31	Roseburia hominis	WP_118096903.1		6MVH[A,B,C,D] 2.40
FMN-binding betα-glucuronidase		Roseburia hominis			7KGZ[A,B] 2.40
β-glucuronidase (Rg3GUS)	3.2.1.31	Ruminococcus gnavus	WP_118581144.1		6MVG[A,B,C] 2.80
β-glucuronidase (UidA)	3.2.1.31	Ruminococcus gnavus E1	AAQ76046.1	A0A2N5PVZ8 Q6W7J7	5Z18[A,B,C,D,E,F] 2.50 5Z19[A,B,C,D,E,F] 2.50 6EC6[A,B] 2.85 6JZ1[A,B] 1.73 6JZ2[A,B] 1.29 6JZ3[A,B] 1.50 6JZ4[A,B] <carb_like_EVA> 1.71 6JZ5[A,B] β-D-GlcpA 1.58 6JZ6[A,B] 1.61 6JZ7[A,B] 1.45 6JZ8[A,B] β-D-GlcpA 1.58
β-glucuronidase (SAG0698)	3.2.1.31	Streptococcus agalactiae 2603V/R	AAM99585.1 NP_687713.1 Q8E0N2		4JKK[A] 2.59 4JKL[A,B] 2.29
β-1,4-galactosidase A (BgaA;SP0648;SP_0648)	3.2.1.23	Streptococcus pneumoniae TIGR4	AAK74795.1 ABA35731.1 NP_345155.1	A0A0H2UP19 Q9KGU0	4CU6[A] 2.70 4CU7[A] 2.20 4CU8[A] <carb_like_DGJ> 2.50 4CUC[A] β-D-Galp-(1-4)-β-D-GlcpNAc 2.20
β-galactosidase Z (LacZ;TmLac;TM1193)	3.2.1.23	Thermotoga maritima MSB8	AAD36268.1 AAA50597.1 AGL50124.1 AHD18900.1 AKE28970.1 CAA04513.1 NP_228998.1	G4FEF3 Q56307	6S6Z[A,B,C,D,E,F,G,H] 2.10 6SD0[A,B,C,D] 3.70
Betα-glucuronidase (uidA_2)	3.2.1.31	uncultured Clostridium sp.		A0A1C5YG41	6U7J[A,B,C,D] 2.20
exo-β-mannanase (XacMan2A; XAC3075) (XacMan2A)	3.2.1.25	Xanthomonas citri pv. citri str. 306	AAM37920.1 NP_643384.1	Q8PI23	6BYC[A] 1.90 6BYE[A,B] β-D-Manp 2.13 6BYG[A,B] β-D-Manp 2.00 6BYI[A,B] β-D-Manp 2.20
Eukaryota
Protein Name	EC#	Organism	GenBank	Uniprot	PDB/3D Carbohydrate Ligands Resolution (Å)
β-mannosidase (MndA;AnigerBM2;An11g06540)	3.2.1.25	Aspergillus niger CBS 513.88	CAK96951.1 XP_001394632.2	A2QWU9	cryst
β-glucuronidase (PGUS;GUS)	3.2.1.31	Aspergillus oryzae Li-3	ABU68712.1	A7XS03	5C70[A,B] 3.10 5C71[A,B,C,D] α-D-GlcpA-(1-1)-<non_carb> 2.62
β-glucuronidase b (GusB;GUSB)	3.2.1.31	Homo sapiens	AAA52561.1 AAH14142.1 AAQ96851.1 AAR75622.1 ABH92596.1 ACH09936.1 ACM86199.1 BAD97126.1 BAG35619.1 BAG61991.1 BAG64768.1 CAC41170.1 NP_000172.1	B2R6X2 B4DTH9 P08236	1BHG[A,B] 2.53 3HN3[A,B,D,E] β-D-Manp 1.70
β-galactosidase (Lac4;BgaL)	3.2.1.23	Kluyveromyces lactis CBS2359	AAA35265.1	P00723	3OB8[A,B,C,D] β-D-Galp 2.80 3OBA[A,B,C,D] 2.75
β-mannosidase	3.2.1.25	Mus musculus	AAK18177.1 AAH31409.1 BAE24954.1 BAE38715.1 BAE41690.1 NP_081564.1	A0A0R4J092 Q8K2I4 Q99MS1	6DDT[A] 2.10 6DDU[A] β-D-Manp 2.67
β-mannosidase (ThMan2A) (Man2A)	3.2.1.25	Trichoderma harzianum IOC-3844	AID21733.1	A0A068CMC5 A0A075B5H6	4CVU[A] 1.90 4UOJ[A,B] 2.50
exo-β-glucosaminidase (Gls93)	3.2.1.165	Trichoderma reesei PC-3-7	BAD99604.1	Q4R1C4	cryst
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