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Glycoside Hydrolase Family 9

Activities in FamilyEndo-β-1,4-glucanase (xanthanase) (EC 3.2.1.-);
Xyloglucan-specific endo-β-1,4-glucanase / endo-xyloglucanase (EC 3.2.1.151);
Exo-β-1,4-glucosaminidase (EC 3.2.1.165);
Endo-β-1,4-glucanase (EC 3.2.1.4);
Endo-β-1,3(4)-glucanase / lichenase-laminarinase (EC 3.2.1.6);
Lichenase / endo-β-1,3-1,4-glucanase (EC 3.2.1.73);
Exo-β-1,4-glucanase / cellodextrinase (EC 3.2.1.74);
Endo-β-1,4-xylanase (EC 3.2.1.8);
Cellobiohydrolase (EC 3.2.1.91);
Mechanism Inverting e
3D Structure Status( α / α ) 6 barrel
Catalytic Nucleophile/BaseAsp (experimental)
Catalytic Proton DonorGlu (experimental)
Noteformerly known as cellulase family E.
External resourcesCAZypedia; HOMSTRAD; PROSITE;
Statistics GenBank accession (7645); Uniprot accession (204); PDB accession (57); 3D entries (22); cryst (0)
Summary
Download GH9 (7430) Taxonomic display Structure (22) Characterized (163)
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Bacteria
Protein Name EC#ReferenceOrganismGenBank UniprotPDB/3D
 endo-β-1,4-glucanase (Cel9B) 3.2.1.4 pubmed
 Acetivibrio cellulolyticus CD2 ATCC 33288 CAI94607.1 Q50HR0  
 endoglucanase (Clocl_2225) 3.2.1.4 pubmed
 Acetivibrio clariflavus DSM 19732 AEV68817.1 G8LXR4  
 endo-β-1,4-glucanase I (CelI;Cthe_0040) (Cel9I) 3.2.1.4 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN51281.1
WP_003511958.1		 Q02934  
 cellobiohydrolase / cellulase K (CelK; Cthe_0412) (Cel9K) 3.2.1.91   Acetivibrio thermocellus ATCC 27405 ABN51650.1 A3DCH1  
 cellobiohydrolase (CbhA;Cthe_0413) (Cbh9A) 3.2.1.91 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN51651.1
AAR87745.1		 A3DCH2
Q59325
Q6RSN8		 1RQ5[A]
1UT9[A]
 endo-β-1,4-glucanase F (CelF;Cthe_0543) (Cel9B) 3.2.1.4 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN51779.1 P26224  
 endo-β-1,4-glucanase R / cellotetraohydrolase (CelR;Cthe_0578) (Cel9R) 3.2.1.4 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN51814.1 A3DCY5 7UNP[A]
7V0I[A,B,C]
7V0J[A,B,C]
 broad substrate-specificity xyloglucanase J / processive endoglucanase (CelJ;Cthe_0624) (Cel9D/Cel9J+Cel44A) 3.2.1.151 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN51859.1 A3DD30  
 endo-β-1,4-glucanase Q (CelQ;Cthe_0625) (Cel9I) 3.2.1.4 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN51860.1 A0A1Y0K4H1
A3DD31		  
 endo-β-1,4-glucanase W (CelW; Cel9W; Cthe_0745) 3.2.1.4 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN51980.1 A3DDF1  
 endo-β-1,4-glucanase (CelD;Cthe_0825) (Cel9A) 3.2.1.4 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN52060.1 A3DDN1  
 endo-β-1,4-glucanase T (CelT;Cthe_2812) (Cel9T) 3.2.1.4 pubmed
 Acetivibrio thermocellus ATCC 27405 ABN54011.1 A3DJ82 2YIK[A]
 endoglucanase T (CelT) (Cel9T) 3.2.1.4 pubmed
 Acetivibrio thermocellus F1 BAB79196.2 Q8VV73  
 endoglucanase J (CelJ) (Cel9D-Cel44A) 3.2.1.4 pubmed
 Acetivibrio thermocellus F1 BAA12070.1 A3DD30
P71140		  
 endoglucanase Q (CelQ) (Cel9I) 3.2.1.4 pubmed
 Acetivibrio thermocellus F1 BAB33148.1 Q9AJF8 5GXX[A,B]
5GXY[A,B]
5GXZ[A,B]
5GY0[A,B]
5GY1[A,B]
 cellobiohydrolase A (CbhA) (Cbh9A) 3.2.1.91 pubmed
 Acetivibrio thermocellus F7 CAA56918.2
CAA06693.1
CAA56918.1		 Q59325
Q7B3G3		  
 cellotetraose-producing endo-β-1,4-glucanase (CelR) (Cel9R) 3.2.1.4 pubmed
 Acetivibrio thermocellus F7 CAE51308.1 Q70DK3  
 endo-β-1,4-glucanase N (CelN) (Cel9N) 3.2.1.4 pubmed
 Acetivibrio thermocellus F7 CAB76935.1 Q9L3J5  
 cellobiohydrolase K (CelK) (Cel9K) 3.2.1.91 pubmed
 Acetivibrio thermocellus JW 20 / F7 AAC06139.1
CAA06692.1		 O68438
O69308
P0C2S1		  
 endo-β-1,4-glucanase D (CelD) (Cel9A) 3.2.1.4 pubmed
 Acetivibrio thermocellus NCIB 10682 CAA28255.1 A3DDN1
P04954
P0C2S4		 1CLC[A]
4CJ0[A]
4CJ1[A]
 endo-β-1,4-glucanase F (CelF) (Cel9B) 3.2.1.4 pubmed
 Acetivibrio thermocellus NCIB 10682 CAA43035.1 P26224  
 endo-β-1,4-glucanase I (CelI;CMC) (Cel9C=Cel9I) 3.2.1.4 pubmed
 Acetivibrio thermocellus NCIB10682 / F7 AAA20892.1
CAB76932.1		 Q02934
Q9L3J8		 2XFG[A]
 cellulase (CelA;Ac_Cel9A;AaCel9A;Aaci_2475) (Cel9A) 3.2.1.4 pubmed
 Alicyclobacillus acidocaldarius subsp. acidocaldarius DSM 446 ACV59481.1
CAC34051.1		 C8WSJ6
Q9AJS0		 3EZ8[A]
3GZK[A]
3H2W[A]
3H3K[A]
3RX5[A]
3RX7[A]
3RX8[A]
5E2J[A,B]
 endo-β-1,4-glucanase (AmCel9; IW967_08975) 3.2.1.4 pubmed
 Alicyclobacillus mali (ex Roth et al. 2021) FL 18 MBF8377998.1    
 β-1,3(4)-glucanase 3.2.1.6 pubmed
 Alicyclobacillus sp. A4 ADH59533.1 D7PC10  
 endoglucanase (BLi01880;BL01232;BlCel9) 3.2.1.4 pubmed
 Bacillus licheniformis DSM 13 = ATCC 14580 AAU40775.1
AAU23415.1		 Q65JI9 8U49[A]
8U4A[A]
8U4F[A]
 cellulase (Cel9A) (Cel9A) 3.2.1.4 pubmed
 Bacillus licheniformis GXN151 AAR29083.1 Q6SYB5  
 endo-β-1,4-glucanase (GH9A) 3.2.1.4 pubmed
 Bacillus licheniformis SVD1 BAL45507.1 H1AD14  
 β-1,4-glucanase, partial 3.2.1.4 pubmed
 Bacillus licheniformis TH15 UYB06747.1
MBY8346522.1		    
 endo-β-1,4-glucanase (EGX) 3.2.1.4 pubmed
 Bacillus pumilus C-9 ADP09350.1 E3UTE0  
 endoglucanase 3.2.1.4 doi
 Bacillus pumilus C18N2 AEQ94264.1 G8I0B7  
 endo-β-1,4-glucanase A (EglA) 3.2.1.4 pubmed
 Bacillus pumilus CL16 AAQ91573.1 Q5YLG1  
 endo-β-1,4-glucanase 3.2.1.4 pubmed
 Bacillus pumilus S-27 AAF15367.1 Q9R9H6  
 endo-β-1,4-glucanase IV 3.2.1.4 pubmed
 Bacillus sp. (in: firmicutes) KSM-522 BAA24918.1 P28622  
 endoglucanase A (Ega;Bα-EGA) 3.2.1.4 pubmed
 Bacillus sp. AC-1 ABM68635.1 A6Y862  
 cellulase (CelA20) 3.2.1.4 doi
 bacterium enrichment culture clone CelA20 ACR23658.1 C5HG55  
 endo-xyloglucanase (CelD; BoGH9A; BACOVA_02649; Bovatus_03068) 3.2.1.151 pubmed
 Bacteroides ovatus ATCC 8483 ALJ47676.1 A7LXT3 6DHT[A]
 endo-1,4-glucanase C (Ced1) 3.2.1.4 pubmed
 Butyrivibrio fibrisolvens H17C CAA39264.1 P23658  
 cellulase (CelA;CbCelA;Athe_1867;Cbes1867;Cbes_1867;CbCBH48A) (Cel9A+Cbh48A) 3.2.1.4 pubmed
 Caldicellulosiruptor bescii DSM 6725 ACM60955.1
CAB06786.1		 B9MKU7
P96311		 4DOD[A]
4DOE[A]
 bifunctional cellulase / β-mannanase (CelC-ManB;CbCel9B/Man5A;Athe_1865;Cbes_1865;Orf1952) (Cel9B-Man5A) 3.2.1.4 pubmed
 Caldicellulosiruptor bescii DSM 6725 ACM60953.1 B9MKU5  
 endoglucanase (J30; BLM47_05245) 3.2.1.4 pubmed
 Candidatus Reconcilbacillus cellulovorans PDO10807.1 A0A2A6E235 5U0H[A]
5U2O[A]
 endo-β-1,4-glucanase B (CenB;Celf_0019) (Cel9A) 3.2.1.4 pubmed
 Cellulomonas fimi ATCC 484 AAA23086.1
AEE44171.1		 F4H3U7
P26225		  
 endo-β-1,4-glucanase C (CenC) (Cel9B) 3.2.1.4 pubmed
 Cellulomonas fimi ATCC 484 CAA40993.1
AAA23088.1
AEE45671.1		 P14090  
 endoglucanase (CBP105) (processive) 3.2.1.4 pubmed
 Cellulomonas flavigena CDBB-531 AAW62376.2 Q3ZMA8  
 cellulose endo-β-1,4-glucosidase (Cfla_3563) 3.2.1.4 pubmed
 Cellulomonas flavigena DSM 20109 ADG76435.1
ARU79882.1		 A0A1Y0K4G5
D5UDH9		  
 endo-β-1,4-glucanase A (CelA;CJA_2472) (Cel9A) 3.2.1.4 pubmed
 Cellvibrio japonicus Ueda107 ACE85757.1
ARU79885.1
CAA31082.1		 B3PKK4
P10476		  
 endoglucanase (CelG;CAC0916) 3.2.1.4 pubmed
 Clostridium acetobutylicum ATCC 824 AAK78892.1
NP_347552.1		 Q97KK3  
 endoglucanase O (EngO) 3.2.1.4 pubmed
 Clostridium cellulovorans AAT66046.1 Q6DTY2  
 endo-β-1,4-glucanase (EngZ;NC1;Clocel_2741) 3.2.1.4 pubmed
 Clostridium cellulovorans 743B ADL52438.1
YP_003844202.1		 D9SRK9  
 endo-β-1,4-glucanase K (EngK) 3.2.1.4 pubmed
 Clostridium cellulovorans 743B AAF06107.1 Q9RGE8  
 endo-β-1,4-glucanase L (EngL; Clocel_2819) 3.2.1.4 pubmed
 Clostridium cellulovorans 743B AAF06109.1
ADL52515.1		 C5RQ37
D9SS68
Q9RGE6		  
 endo-β-1,4-glucanase M (EngM) 3.2.1.4 pubmed
 Clostridium cellulovorans 743B AAF06111.2 Q9LAJ2  
 endo-β-1,4-glucanase Y (EngY) 3.2.1.4 pubmed
 Clostridium cellulovorans ATCC 35269 AAG59608.1 Q9AQF4  
 endo-β-1,4-glucanase H (EngH) 3.2.1.4 pubmed
 Clostridium cellulovorans ATCC 35296 AAC38572.2 O65987  
 xanthan endo-β-1,4-glucosidase (CspXan9; VQ071_26415) 3.2.1.- doi
 Cohnella sp. 56 VKM B-36720 MEK0317390.1    
 endoglucanase 9A (CHU_1336) (Cel9A) 3.2.1.4 pubmed
 Cytophaga hutchinsonii ATCC 33406 ABG58608.1 A0A6N4SQK1
Q11VF8		  
 endoglucanase / cellulose-binding protein A / E1 (Cbpe1;CbpA) 3.2.1.4 pubmed
 Eubacterium cellulosolvens 5 BAB86305.1 Q8RR29  
 endoglucanase / cellulose-binding protein B (CbpB) 3.2.1.4 pubmed
 Eubacterium cellulosolvens 5 BAE20171.1 Q402U6  
 endo-1,4-glucanase A (EndA) 3.2.1.4 pubmed
 Fibrobacter succinogenes AR1 AAA24894.1
AAA24895.1		 P23665  
 endo-β-1,4-glucanase C (EgC;CMCase) 3.2.1.4 pubmed
 Fibrobacter succinogenes BL2 AAC41523.1 Q59442  
 endo-β-1,4-glucanase B (EndB;EgB;Fisuc_0057;FSU_0451) (Cel9G) 3.2.1.4 pubmed
 Fibrobacter succinogenes subsp. succinogenes S85 AAA68129.1
ACX73672.1
ADL25229.1		 C9RJ28
Q59444		  
 endo-1,4-glucanase D (CelD; Fisuc_1860; FSU_2362) (Cel9C) 3.2.1.4 pubmed
 Fibrobacter succinogenes subsp. succinogenes S85 AAC44386.1
ACX75452.1
ADL24960.1		 D9S4N9
P77864		  
 endoglucanase 1 (CelE;End1;Fisuc_1859;FSU_2361) (Cel9B) 3.2.1.4 pubmed
 Fibrobacter succinogenes subsp. succinogenes S85 ABU45498.1
AAC44385.1
ACX75451.1
ADL26362.1		 A7UG67
P71326		  
 1,4-β-glucan glucohydrolase (Fisuc_2033;FSU_2558) (Cel9D) 3.2.1.74 pubmed
 Fibrobacter succinogenes subsp. succinogenes S85 ABY60376.1
ACX75620.1
ADL26040.1		 B0FWI3
C9RJA3		  
 endo-β-1,4-glucanase 9A (CpCel9; cphy3367; Cphy_3367) (Cel9A) 3.2.1.4 pubmed
 Lachnoclostridium phytofermentans ISDg ABX43720.1 A9KT90  
 endo-xanthanase (MiXen; AB663_001095) 3.2.1.- pubmed
 Microbacterium sp. XT11 ALX66163.1    
 endocellulase (EgltA) (Cel9) 3.2.1.4 pubmed
 Myxobacter sp. AL-1 AAF19168.1 Q9RAI8  
 endo-β-1,4-glucanase B (CelB) (Cel9B) 3.2.1.4 pubmed
 Paenibacillus barcinonensis BP-23 CAB38941.1 Q9Z4I1  
 endo-β-1,4-glucanase (Cel9A; PcMulGH9) 3.2.1.4 pubmed
 Paenibacillus curdlanolyticus B-6 AND99925.1 A0A172QQ23  
 endo-xanthanase (Xan9; PspXan9) 3.2.1.- doi
 Paenibacillus nanensis 62047 / Paenibacillus sp. 62047 AXR85426.1 A0A346PYN7
A0A3F2YM16
A0A3F2YM17		 6FHJ[A]
6FHN[A]
 endo-β-1,4-glucanase (Cel9P) 3.2.1.4 pubmed
 Paenibacillus sp. BME-14 ACJ68032.1 B7U9C9  
 exo-β-glucosaminidase (GlcNase;PpGlcNase;PBPRA0520) 3.2.1.165 pubmed
 Photobacterium profundum SS9 CAG18943.1 Q6LUT2 5DGQ[A,B]
5DGR[A,B]
 endoglucanase (fragment) (Cel9A) 3.2.1.4 pubmed
 Pseudomonas sp. SK38 AAG49558.1 Q9APG3  
 endo-β-1,4-glucanase (CelA) 3.2.1.4 pubmed
 Pseudomonas sp. YD-15 AAD01959.1 Q9Z3X7  
 cellulase 9R (Ccel_1648) (Cel9R) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75999.1 B8I2K6  
 endo-xyloglucanase 9X (Ccel_2621) (Cel9X) 3.2.1.151 pubmed
 Ruminiclostridium cellulolyticum H10 ACL76949.1 B8I6V7  
 cellulase 9W (Ccel_2226) (Cel9W) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL76568.1 B8I4R4  
 cellulase 9U / cellulase 9S (Ccel_0755) (Cel9S=Cel9U) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75133.1 B8I7X6  
 β-glucanase 9T (Ccel_1249) (Cel9T) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75605.1 B8I0N0  
 cellulase 9E (CelE; Ccel_0732) (Cel9E) 3.2.1.91 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75111.1
AAA73869.2		 B8I7V4
Q46002		  
 endo-β-1,4-glucanase 9G (CelG; Ccel_0731) (Cel9G) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75110.1
AAA73868.1		 P37700 1G87[A,B]
1GA2[A,B]
1K72[A,B]
1KFG[A,B]
 cellulase 9Q (Ccel_0231) (Cel9Q) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL74618.1
ABG76967.1		 Q0PRN4  
 cellulase 9P (P90; Ccel_0753) (Cel9P) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75131.1
ABG76972.1		 Q0PRM9  
 cellulase M / 9M (CelM; Ccel_0737) (Cel9M) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75116.1
AAG45160.1		 B8I7V9
Q9EYQ2		 1IA6[A]
1IA7[A]
 cellulase 9J (CelJ; Ccel_0735) (Cel9J) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75114.1
AAG45158.1		 B8I7V7
Q9EYQ4		  
 cellulase 9H (CelH; Ccel_0734) (Cel9H) 3.2.1.4 pubmed
 Ruminiclostridium cellulolyticum H10 ACL75113.1
AAG45157.1		 B8I7V6
Q9EYQ5		  
 endo-β-1,4-glucanase (GH9A;RUM_01230) (Cel9A) 3.2.1.4 pubmed
 Ruminococcus champanellensis 18P13 = JCM 17042 CBL16391.1 D4L9U6  
 endo-β-1,4-glucanase (GH9C;RUM_08690) (Cel9C) 3.2.1.4 pubmed
 Ruminococcus champanellensis 18P13 = JCM 17042 CBL17047.1 D4LBQ2  
 endo-β-1,4-glucanase (GH9D;RUM_14490) (Cel9D) 3.2.1.4 pubmed
 Ruminococcus champanellensis 18P13 = JCM 17042 CBL17554.1 D4LD59  
 endo-β-1,4-glucanase (GH9E;RUM_15950) (Cel9E) 3.2.1.4 pubmed
 Ruminococcus champanellensis 18P13 = JCM 17042 CBL17683.1 D4LDI8  
 endo-β-1,4-glucanase (GH9F;RUM_15960) (Cel9F) 3.2.1.4 pubmed
 Ruminococcus champanellensis 18P13 = JCM 17042 CBL17684.1 D4LDI9  
 endo-β-1,4-glucanase (GH9G;RUM_24010) (Cel9G) 3.2.1.4 pubmed
 Ruminococcus champanellensis 18P13 = JCM 17042 CBL18389.1 D4LFJ4  
 endo-β-1,4-glucanase I (Cel1) 3.2.1.4 pubmed
 Streptomyces reticuli CAA46570.1 Q05156  
 endo-β-1,3-1,4-glucanase (slr0897;Glc) 3.2.1.73 pubmed
 Synechocystis sp. PCC 6803 BAA10447.1
AGF52739.1
ALJ68659.1
BAF51695.1
BAK51232.1
NP_442377.1
QWO79905.1		 A4PCG1
Q55365		  
 endo-β-1,4-glucanase (E1; EGtf1; Tfu_1627) (Cel9B) 3.2.1.4 pubmed
 Thermobifida fusca YX AAC06387.1
AAZ55662.1		 Q08166
Q47PF7		  
 endo/exo-β-1,4-glucanase 9A (E4; CelD; TfCel9A; Tfu_2176) (processive) (Cel9A) 3.2.1.4 pubmed
 Thermobifida fusca YX AAB42155.1
AAA27397.1
AAZ56209.1		 P26221
Q47MW0		 1JS4[A,B]
1TF4[A,B]
3TF4[A,B]
4TF4[A,B]
 endo-β-1,4-glucanase (ThCel9A) (Cel9A) 3.2.1.4 pubmed
 Thermobifida halotolerans YIM 90462 AEH04391.1 G4XKY7  
 endo-β-1,4-glucanase Z / avicelase I (CelZ) 3.2.1.4 pubmed
 Thermoclostridium stercorarium CAA39010.1 P23659  
 endo-glucanase 9Z (Cel9Z; CelZ; Cst_c18960) 3.2.1.4 pubmed
 Thermoclostridium stercorarium subsp. stercorarium DSM 8532 AGI39870.1
AGC68873.1		 L7VTG9  
 endoglucanase (Cel9A; BP_Cel9A) 3.2.1.6
3.2.1.151		 pubmed
pubmed
 uncultured bacterium AXR85444.1 A0A346PYQ5  
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