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)

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Archaea
Protein Name	EC#	Reference	Organism	GenBank	Uniprot	PDB/3D
β-glucuronidase (GusB;Ssol98_010100007956)	3.2.1.31	pubmed	Saccharolobus solfataricus 98/2	AET42956.1	G8GCV1
Bacteria
Protein Name	EC#	Reference	Organism	GenBank	Uniprot	PDB/3D
β-galactosidase (LacZ)	3.2.1.23	pubmed	Actinobacillus pleuropneumoniae CM5	AAB17954.1	P70753
β-galactosidase (AgWH2A)	3.2.1.23	pubmed	Agarivorans gilvus WH0801	AXA12974.1	A0A2Z4WWH8
β-mannosidase (ManA; AfMan2; Atu1523)	3.2.1.25	pubmed	Agrobacterium fabrum str. C58	AAL42528.1 AAK87314.1 AAK87314.2 NP_354529.1 NP_532212.1	Q7CZ23
β-galactosidase	3.2.1.23	pubmed	Alkalilactibacillus ikkensis 517	ACO52514.1	C1KC07
β-galactosidase (GalA)	3.2.1.23	pubmed	Alteromonas sp. ANT48	QGZ98840.1	A0A6B9KBH1
β-galactosidase (Bgal)	3.2.1.23	doi	Alteromonas sp. ML117	AZK90254.1	A0A3S8S8V3
β-galactosidase	3.2.1.23	pubmed	Alteromonas sp. ML52	AZK90255.1	A0A3Q8SL57
β-galactosidase (Gal2A)	3.2.1.23	pubmed	Alteromonas sp. QD01	QPF69526.1	A0A7U3PJ41
exo-β-glucosaminidase / exochitosanase (CsxA)	3.2.1.165	pubmed	Amycolatopsis orientalis	AAX62629.2 AAX62629.1	Q56F26	2VZO[A,B] 2VZS[A,B] 2VZT[A,B] 2VZU[A,B] 2VZV[A,B] 2X05[A,B] 2X09[A,B]
β-glucuronidase/β-galacturonidase (AqGalA)	3.2.1.31 3.2.1.-	pubmed pubmed	Aquimarina sp. SCSIO21287	UER37076.1		7VQM[A,B,C,D]
β-galactosidase (BglA)	3.2.1.23	pubmed	Arthrobacter psychrolactophilus F2	BAF33372.1	Q08KN3
β-galactosidase (LacZ)	3.2.1.23	pubmed	Arthrobacter sp.	AAA69907.1	Q59140
β-galactosidase (BgaS)	3.2.1.23	pubmed	Arthrobacter sp. 20B	ACI41243.1	B6EAX7
β-galactosidase (Arth?DG)	3.2.1.23	pubmed	Arthrobacter sp. 32cB	AHY00656.1	A0A023UGN9	6ETZ[A] 6H1P[A] 6SE8[A] 6SE9[A] 6SEA[A] 6SEB[A] 6SEC[A] 6SED[A] 6ZJP[A] 6ZJQ[A] 6ZJR[A] 6ZJS[A] 6ZJT[A] 6ZJU[A] 6ZJV[A] 6ZJW[A] 6ZJX[A]
β-galactosidase (LacZ)	3.2.1.23	doi	Arthrobacter sp. C2-2	CAD29775.1	Q8KRF6	1YQ2[A,B,C,D,E,F]
β-galactosidase (GalA)	3.2.1.23	pubmed	Arthrobacter sp. ON14	ADJ18283.1	D9IU58
β-glucuronidase (Gus)	3.2.1.31	pubmed	Arthrobacter sp. RP10	AAV91790.1	Q5MJX7
β-glucuronidase (Gus)	3.2.1.31	pubmed	Arthrobacter sp. RP40.7	AAV91792.1	Q5MJX5
β-galactosidase (LacZ)	3.2.1.23	pubmed	Arthrobacter sp. SB	AAQ19029.1	Q7WTU5
β-xylosidase (BcellWH2_03638; BACWH2_3409; Lacz_21)	3.2.1.37	pubmed	Bacteroides cellulosilyticus WH2	ALJ60861.1 WP_029427454.1	A0A0P0GUA9
β-glucuronidase (BcellWH2_04226; BACWH2_3980; Lacz_22)	3.2.1.31	pubmed	Bacteroides cellulosilyticus WH2	ALJ61443.1 WP_029428765.1	A0A0N7IFX8
glycoside hydrolase cleaving tamarind xyloglucan (BcellWH2_04335; Lacz_23)	3.2.1.21	pubmed	Bacteroides cellulosilyticus WH2	ALJ61552.1 WP_029428707.1	A0A0P0GBJ1
β-glucuronidase (BF0328)	3.2.1.31	pubmed	Bacteroides fragilis NCTC 9343	CAH06099.1	Q5LIC7	3CMG[A] 5Z1A[A]
β-galactosidase (BgaA; BF1738)	3.2.1.23	pubmed	Bacteroides fragilis YCH46	BAD48485.1 BAC56904.1 BAC56904.2	G8K035 Q84ID4
BF1733 (BmnA)	3.2.1.25	pubmed	Bacteroides fragilis YCH46	BAD48480.1 BAC56899.1 BAC56899.2	G8K032 Q84ID9
β-galactosidase (Lacz_13;BoGH2A;BACOVA_02645;Bovatus_03065)	3.2.1.23	pubmed	Bacteroides ovatus ATCC 8483	ALJ47673.1
β-glucuronidase (Bovatus_02309;BACOVA_03100;Uida_1)	3.2.1.31	pubmed	Bacteroides ovatus ATCC 8483	ALJ46942.1
β-glucuronidase (BMSP;HMPREF2532_04191)	3.2.1.31	pubmed	Bacteroides ovatus KLE1656	KXT42440.1	A0A139KTC9	6D8K[B,C,D] 6D8K[A]
β-galacturonidase (BT0992; BT_0992)	3.2.1.-	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO76099.1 NP_809905.1	Q8A925
β-mannosidase (BT0458;BT_0458) (Man2A)	3.2.1.25	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO75565.1 NP_809371.1	Q8AAK6	2JE8[A,B] 2VJX[A,B] 2VL4[A,B] 2VMF[A,B] 2VO5[A,B] 2VOT[A,B] 2VQT[A,B] 2VQU[A,B] 2VR4[A,B] 2WBK[A,B] 7OP6[A,B] 7OP7[A,B]
bimodular β-L-arabinofuranosidase / β-glucuronidase (BT0996; BT_0996)	3.2.1.31	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO76103.1 NP_809909.1	Q8A921
β-galactosidase (BT4151;BT_4151)	3.2.1.23	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO79256.1 NP_813062.1	Q8A070
α-L-arabinopyranosidase (cleaves L-Arap-α-1,4-D-Gal; BT0983; BT_0983)	3.2.1.-	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO76090.1 NP_809896.1	Q8A934
β-galactosidase (BT4156;BT_4156)	3.2.1.23	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO79261.1 NP_813067.1	Q8A065
β-glucuronidase (BT4181;BT_4181)	3.2.1.31	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO79286.1 NP_813092.1	Q8A040
β-galactosidase (BT0461)	3.2.1.23	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO75568.1 NP_809374.1	Q8AAK3
β-galactosidase (cleaves D-Galp-β-1,2-L-Acef; BT0993)	3.2.1.23	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO76100.1 NP_809906.1	Q8A924
β-galactosidase (BT4667;BT_4667)	3.2.1.23	pubmed	Bacteroides thetaiotaomicron VPI-5482	AAO79772.1 NP_813578.1	Q89YR4
exo-β-galactosidase porphyran-specific (BuGH2A)	3.2.1.23	doi	Bacteroides uniformis NP1	WP_234130811.1	A0A2D0TCC8	5T98[A,B] 5T99[A,B]
agarolytic β-galactosidase (BuGH2C)	3.2.1.23	pubmed	Bacteroides uniformis NP1		A0A2D0TCC9	5T9A[B,C,D] 5T9G[A,B,C,D] 5T9A[A]
β-glucuronidase (BuGUS-2; M094_3283)	3.2.1.31	pubmed	Bacteroides uniformis str. 3978 T3 i	KDS62590.1	A0A078SUX9 A0A414F142	5UJ6[A,B] 6D50[A,B] 6D8G[A,B] 6NZG[A,B]
β-glucuronidase (BuGUS-1; ERS417307_01040; Uida_4)	3.2.1.31	pubmed	Bacteroides uniformis str. 3978 T3 i	CUO13985.1	A0A174CQK8	6D1N[A,B] 6D41[A,B] 6D6W[A,B,C,D] 6D7F[A,B,C,D,E,F] 6D89[A,B,C,D]
β-galactosidase (BgaC;BAD_1582)	3.2.1.23	pubmed	Bifidobacterium adolescentis ATCC 15703	BAF40363.1	A1A3T0
β-galactosidase (BIF1;OLGA1)	3.2.1.23	pubmed	Bifidobacterium bifidum DSM 20215	CAB75342.1	Q9L3L4
β-galactosidase (BIF3)	3.2.1.23	pubmed	Bifidobacterium bifidum DSM 20215	CAC14566.2 AAP64961.1 ABI07182.1 CAD19366.1	Q9F4D5	6QUB[A,B] 6QUC[A,B] 6QUD[A] 7NIT[A,B,C,D,E,F]
β-galactosidase (BIF2)	3.2.1.23	pubmed	Bifidobacterium bifidum DSM 20215	CAC14565.1	Q9F4D6
lacto-N-neotetraose β-galactosidase (BbgIII)	3.2.1.23	pubmed	Bifidobacterium bifidum JCM 1254	BAI94821.1	D4QAP3	5DMY[A,B,C]
β-galactosidase P1 (BbgI;Bbg1)	3.2.1.23	pubmed	Bifidobacterium bifidum NCIMB 41171	ABE27151.1 ABE27151.2	Q0ZI53
β-galactosidase S1 specific for lactose (BbgIII)	3.2.1.108	pubmed	Bifidobacterium bifidum NCIMB 41171	ABE27152.1	A4K5H9
β-galactosidase K2 (BbgIV)	3.2.1.23	pubmed	Bifidobacterium bifidum NCIMB 41171	ABE00939.1	Q0ZII7
β-galactosidase (LacZ;Lacz6;Bbr_1552_lacZ;Bbr_1552)	3.2.1.23	pubmed	Bifidobacterium breve UCC2003	ABE96223.1
β-galactosidase (Bbr_0010;Lacz1)	3.2.1.23	pubmed	Bifidobacterium breve UCC2003	ABE94727.1
HMPREF0168_2111 (UidA)	3.2.1.31	pubmed	Bifidobacterium dentium ATCC 27679	WP_003843201.1	E0QAF3	5Z1B[A,B,C,D]
β-galactosidase 2A (Blon_2334) (Bga2A)	3.2.1.23	pubmed	Bifidobacterium longum subsp. infantis ATCC 15697 = JCM 1222 = DSM 20088	ACJ53392.1	B7GNN6
β-galactosidase (Blon_0268)	3.2.1.23	pubmed	Bifidobacterium longum subsp. infantis ATCC 15697 = JCM 1222 = DSM 20088	ACJ51396.1	B7GTV5
β-galactosidase I (β-GalI)	3.2.1.23	pubmed	Bifidobacterium longum subsp. infantis HL96	AAL02052.1	Q93GI6
β-galactosidase (LacZ)	3.2.1.23	pubmed	Bifidobacterium longum subsp. longum MB219	CAB44428.1	Q9X6Y5
β-galactosidase (B8809_0611)	3.2.1.23	pubmed	Bifidobacterium longum subsp. longum NCIMB8809	ALO72284.1
galactomannan-binding β-1,4-mannosidase (CbMan2A;Athe_0227) (Man2A)	3.2.1.25	pubmed	Caldicellulosiruptor bescii DSM 6725	ACM59377.1	B9MME5
β-mannosidase 2A (CfMan2A) (Man2A)	3.2.1.25	pubmed	Cellulomonas fimi ATCC 484	AAD42775.1 AEE46894.1	F4H7D3 Q9XCV4
β-galactosidase	3.2.1.23	pubmed	Clostridium acetobutylicum	AAA23216.1	P24131
β-glucuronidase (BglR;CPE0147)	3.2.1.31	pubmed	Clostridium perfringens str. 13	BAB79853.1 CAD12654.1 NP_561063.1	Q8VNV4 Q8XP19	4JKM[A,B] 6CXS[A,B]
β-glucuronidase (UidA)	3.2.1.31	pubmed	Clostridium sp. Marseille-P299	WP_066714276.1
β-glucuronidase (DtGlcA; DICTH_1429)	3.2.1.31	pubmed	Dictyoglomus thermophilum H-6-12	ACI18585.1	B5YFE0	6XXW[A]
β-mannosidase (DtMan;DICTH_1692)	3.2.1.25	pubmed	Dictyoglomus thermophilum H-6-12	ACI19676.1	B5YAN4	5N6U[A,B,C,D]
β-galacturonidase (EtGalAse)	3.2.1.-	pubmed	Eisenbergiella tayi	WP_069152983.1	A0A1E3AEY6	6NCW[A,B,C,D] 6NCX[A,B,C,D]
β-galactosidase (Bga;BgaB5)	3.2.1.23	doi	Enterobacter cloacae B5	ABB73039.1	Q2XQU3
β-galactosidase (LacZ)	3.2.1.23	pubmed	Enterobacter cloacae GAO	BAA07673.1	Q47077
β-galactosidase (LacZ)	3.2.1.23	pubmed	Erwinia rhapontici E602	AYC35409.1	A0A385ZAF4
β-galactosidase (LacZ;EcDH1_3262)	3.2.1.23	pubmed	Escherichia coli DH1	ACX40886.1 BAJ42187.1	C9QQT4
β-galactosidase Z (LacZ;Lac-Z;b0344;EO53_16565)	3.2.1.23	pubmed	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] 1BGM[I] 1DP0[A] 1F49[A] 1F4A[A] 1F4H[A] 1GHO[I] 1HN1[A,B,C,D] 1JYN[A] 1JYV[A] 1JYW[A] 1JYX[A] 1JYY[A] 1JYZ[I] 1JZ0[A] 1JZ1[I] 1JZ2[A] 1JZ3[A] 1JZ4[A] 1JZ5[A] 1JZ6[A] 1JZ7[A] 1JZ8[A] 1PX3[A,B,C,D] 1PX4[A,B,C,D] 3CZJ[A,B,C,D] 3DYM[A,B,C,D] 3DYO[A,B,C,D] 3DYP[A,B,C,D] 3E1F[1,2,3,4] 3I3B[A,B,C,D] 3I3D[A,B,C,D] 3I3E[A,B,C,D] 3IAP[A,B,C,D] 3IAQ[A,B,C,D] 3J7H[A,B,C,D] 3MUY[1,2,3,4] 3MUZ[1,2,3,4] 3MV0[1,2,3,4] 3MV1[1,2,3,4] 3SEP[A,B,C,D] 3T08[A,B,C,D] 3T09[A,B,C,D] 3T0A[A,B,C,D] 3T0B[A,B,C,D] 3T0D[A,B,C,D] 3T2O[A,B,C,D] 3T2P[A,B,C,D] 3T2Q[A,B,C,D] 3VD3[A,B,C,D] 3VD4[A,B,C,D] 3VD5[A,B,C,D] 3VD7[A,B,C,D] 3VD9[A,B,C,D] 3VDA[A,B,C,D] 3VDB[A,B,C,D] 3VDC[A,B,C,D] 4CKD[A,B,C,D] 4DUV[A,B,C,D] 4DUW[A,B,C,D] 4DUX[A,B,C,D] 4TTG[A,B,C,D] 4V40[A,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P] 4V41[A,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P] 4V44[A,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P] 4V45[A,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P] 5A1A[A,B,C,D] 6CVM[A,B,C,D] 6DRV[A,B,C,D] 6KUZ[A,B,C,D] 6X1Q[A,B,C,D] 7BRS[A,B,C,D] 7BTK[A,B,C,D]
β-glucuronidase (UidA;GusA;GurA;b1617)	3.2.1.31	pubmed	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] 3K4A[A,B] 3K4D[A,B] 3LPF[A,B] 3LPG[A,B] 4JHZ[A,B] 5CZK[A,B] 7PR6[A,B]
evolved β-galactosidase (EbgA;b3076)	3.2.1.23	pubmed	Escherichia coli str. K-12 substr. MG1655	AAA61971.1 AAA57877.1 AAC76111.1 AAT48164.1 AIZ89904.1 ALI39224.1 AMC95983.1 AUG17794.1 AVI53249.1 AYG17960.1 CAA36274.1 NP_417547.2	P06864
β-glucuronidase (EeGUS; EUBELI_20590)	3.2.1.31	pubmed	[Eubacterium] eligens ATCC 27750	ACR73734.1	A0A174ZZA3 C4Z6Z2	6BJQ[A] 6BJW[A] 6BO6[A] 6D4O[A]
β-glucuronidase (CG447_12395)	3.2.1.31	pubmed	Faecalibacterium prausnitzii A2165	ATP00638.1 EEU97254.1	C7H4D2	6ED2[A]
β-glucuronidase (C4Q21_10165)	3.2.1.31	pubmed	Faecalibacterium prausnitzii APC918/95b	AXB29257.1	A0A3F3JX71	6U7I[A,B,C,D]
β-glucuronidase (Fp2GUS; FP2_04120)	3.2.1.31	pubmed	Faecalibacterium prausnitzii L2-6 L2/6	CBK98066.1	D4K3H3	6MVF[A,B,C,D,E,F]
β-galacturonidase (FcGalAse; BHW29_02690)	3.2.1.-	pubmed	Faecalibacterium sp. CAG:74_58_120	OLA24209.1	A0A1Q6Q230
β-galactosidase (Gal4214-1)	3.2.1.23	pubmed	Flavobacterium sp. 4214	AAX48919.1	Q2VT50
[ulvan] β-1,2-glucuronidase (BN863_22060)	3.2.1.31	pubmed	Formosa agariphila KMM 3901	CDF79918.1	T2KN75	6HPD[A]
FsGUS (β-galacturonidase/β-glucuronidase)	3.2.1.31 3.2.1.-	pubmed pubmed	Fusicatenibacter saccharivorans		A0A174EHD1	6NCY[B] 6NCZ[A,B,C,D,E,F] 6NCY[A]
β-galactosidase (BglP)	3.2.1.23	pubmed	gamma proteobacterium NT2-2	AMK07416.1	A0A140D6D2
β-galactosidase (LacZ)	3.2.1.23	pubmed	Klebsiella pneumoniae 285	AEA30145.1	F2X2W7
β-galactosidase (GalB)	3.2.1.23	pubmed	Klebsiella sp. T17-2	AMK07522.1	A0A126QG41
β-glucuronidase (HMPREF0539_0313; UidA)	3.2.1.31	pubmed	Lacticaseibacillus rhamnosus LMS2-1	EEN81625.1	A0A2A5L2J1 C2JTS9	6ECA[A,B]
β-galactosidase (LacL+LacM)	3.2.1.23	pubmed	Lactiplantibacillus plantarum ATCC 8014	AAL09167.1 ATQ34817.1	Q938Q6
β-galactosidase (LacL+LacM)	3.2.1.23	pubmed	Lactiplantibacillus plantarum FUA3112	CAZ66936.1	C6H176
β-galactosidase (LacL+LacM,lp_3483+lp_3484)	3.2.1.23	pubmed	Lactiplantibacillus plantarum WCFS1	CAD65569.1 CCC80451.1 NP_786691.1	F9UUE9
β-galactosidase large subunit (LacL)	3.2.1.23	pubmed	Lactobacillus acidophilus ATCC 4356	ACC38286.1	B2LWG2
β-galactosidase Z	3.2.1.23	pubmed	Lactobacillus delbrueckii	AAA25240.1 AAA25244.1 AAY08349.1 ACW22137.1 CAC07488.1	P0C1Y0 Q1G9Z4
β-galactosidase Z (LacZ;Lbulagal;Ldb1201)	3.2.1.23	pubmed	Lactobacillus delbrueckii subsp. bulgaricus ATCC 11842 = JCM 1002	CAI98003.1 AEN14565.1	Q1G9Z4
β-galactosidase (BgaL3)	3.2.1.23	doi	Lactobacillus delbrueckii subsp. bulgaricus L3	ACE06986.1	B3GS90
β-glucuronidase (GusA)	3.2.1.31	pubmed	Lactobacillus gasseri ADH	AAK07836.1 AAS22718.1	Q9AHJ8
β-galactosidase large subunit (LacL;WANG_0293)	3.2.1.23	pubmed	Lactobacillus kefiranofaciens ZW3	AEG39988.1
β-galactosidase (LacZ)	3.2.1.23	doi	Lactococcus lactis ATCC 7962	AAD11504.1 AAC63020.1 CAA56341.1	O87523 Q48727
β-galactosidase (LacZ;L0025)	3.2.1.23	pubmed	Lactococcus lactis subsp. lactis Il1403	AAK06078.1 NP_268137.2	Q48727
β-galactosidase L+S (LacL+LacS) (heterodimer)	3.2.1.23	pubmed	Latilactobacillus sakei subsp. sakei DSM 20017	CAA57730.1	Q48846
β-galactosidase L+M (LacL+LacM)	3.2.1.23	pubmed	Leuconostoc lactis	AAA25267.1 CAL48858.1	Q02603
LVIS_2259 + LVIS_2258	3.2.1.23	pubmed	Levilactobacillus brevis ATCC 367	ABJ65308.1	Q03NB4
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