PDBsum entry 3tf4

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protein ligands metals Protein-protein interface(s) links
Glycosyl hydrolase PDB id
Protein chains
605 a.a. *
BGC ×2
_CA ×4
Waters ×1375
* Residue conservation analysis
PDB id:
Name: Glycosyl hydrolase
Title: Endo/exocellulase:cellotriose from thermomonospora
Structure: T. Fusca endo/exo-cellulase e4 catalytic domain and cellulose-binding domain. Chain: a, b. Fragment: catalytic domain and cellulose-binding domain. Synonym: endo-1,4-beta-d-glucanase. Engineered: yes. Other_details: crystal soaked in 0.3m cellotriose
Source: Thermobifida fusca. Organism_taxid: 2021. Gene: bamh1-pst1 fragment of t. Fusc. Expressed in: streptomyces lividans tk24. Expression_system_taxid: 457428. Genomic DNA carrying native e4 gene
2.20Å     R-factor:   0.214    
Authors: J.Sakon,D.B.Wilson,P.A.Karplus
Key ref: J.Sakon et al. (1997). Structure and mechanism of endo/exocellulase E4 from Thermomonospora fusca. Nat Struct Biol, 4, 810-818. PubMed id: 9334746
30-May-97     Release date:   04-Sep-97    
Go to PROCHECK summary

Protein chains
Pfam   ArchSchema ?
P26221  (GUN4_THEFU) -  Endoglucanase E-4
880 a.a.
605 a.a.
Key:    PfamA domain  Secondary structure  CATH domain

 Enzyme reactions 
   Enzyme class: E.C.  - Cellulase.
[IntEnz]   [ExPASy]   [KEGG]   [BRENDA]
      Reaction: Endohydrolysis of 1,4-beta-D-glucosidic linkages in cellulose, lichenin and cereal beta-D-glucans.
 Gene Ontology (GO) functional annotation 
  GO annot!
  Biological process     carbohydrate metabolic process   1 term 
  Biochemical function     catalytic activity     4 terms  


Nat Struct Biol 4:810-818 (1997)
PubMed id: 9334746  
Structure and mechanism of endo/exocellulase E4 from Thermomonospora fusca.
J.Sakon, D.Irwin, D.B.Wilson, P.A.Karplus.
Cellulase E4 from Thermomonospora fusca is unusual in that it has characteristics of both exo- and endo-cellulases. Here we report the crystal structure of a 68K M(r) fragment of E4 (E4-68) at 1.9 A resolution. E4-68 contains both a family 9 catalytic domain, exhibiting an (alpha/alpha)6 barrel fold, and a family III cellulose binding domain, having an antiparallel beta-sandwich fold. While neither of these folds is novel, E4-68 provides the first cellulase structure having interacting catalytic and cellulose binding domains. The complexes of E4-68 with cellopentaose, cellotriose and cellobiose reveal conformational changes associated with ligand binding and allow us to propose a catalytic mechanism for family 9 enzymes. We also provide evidence that E4 has two novel characteristics: first it combines exo- and endo-activities and second, when it functions as an exo-cellulase, it cleaves off cellotetraose units.

Literature references that cite this PDB file's key reference

  PubMed id Reference
21188618 J.L.Chir, C.F.Wan, C.H.Chou, and A.T.Wu (2011).
Hydrolysis of cellulose in synergistic mixtures of β-glucosidase and endo/exocellulase Cel9A from Thermobifida fusca.
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GPR39: a Zn(2+)-activated G protein-coupled receptor that regulates pancreatic, gastrointestinal and neuronal functions.
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21307602 Q.Yin, Y.Teng, Y.Li, M.Ding, and F.Zhao (2011).
Expression and characterization of full-length Ampullaria crossean endoglucanase EG65s and their two functional modules.
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21098515 Y.Honda, N.Shimaya, K.Ishisaki, M.Ebihara, and H.Taniguchi (2011).
Elucidation of exo-{beta}-D-glucosaminidase activity of a family 9 glycoside hydrolase (PBPRA0520) from Photobacterium profundum SS9.
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19957081 A.I.Chiriac, E.M.Cadena, T.Vidal, A.L.Torres, P.Diaz, and F.I.Pastor (2010).
Engineering a family 9 processive endoglucanase from Paenibacillus barcinonensis displaying a novel architecture.
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19787349 A.Sunna (2010).
Modular organisation and functional analysis of dissected modular beta-mannanase CsMan26 from Caldicellulosiruptor Rt8B.4.
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20373916 C.M.Fontes, and H.J.Gilbert (2010).
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SusG: a unique cell-membrane-associated alpha-amylase from a prominent human gut symbiont targets complex starch molecules.
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PDB codes: 3k8k 3k8l 3k8m
20057047 S.Petkun, S.Jindou, L.J.Shimon, S.Rosenheck, E.A.Bayer, R.Lamed, and F.Frolow (2010).
Structure of a family 3b' carbohydrate-binding module from the Cel9V glycoside hydrolase from Clostridium thermocellum: structural diversity and implications for carbohydrate binding.
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PDB codes: 2wnx 2wo4 2wob
20552664 T.V.Vuong, and D.B.Wilson (2010).
Glycoside hydrolases: catalytic base/nucleophile diversity.
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20173060 Y.Li, D.C.Irwin, and D.B.Wilson (2010).
Increased crystalline cellulose activity via combinations of amino acid changes in the family 9 catalytic domain and family 3c cellulose binding module of Thermobifida fusca Cel9A.
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20348303 Y.Vazana, S.Moraïs, Y.Barak, R.Lamed, and E.A.Bayer (2010).
Interplay between Clostridium thermocellum family 48 and family 9 cellulases in cellulosomal versus noncellulosomal states.
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19775243 A.C.Tolonen, A.C.Chilaka, and G.M.Church (2009).
Targeted gene inactivation in Clostridium phytofermentans shows that cellulose degradation requires the family 9 hydrolase Cphy3367.
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19617364 B.J.Watson, H.Zhang, A.G.Longmire, Y.H.Moon, and S.W.Hutcheson (2009).
Processive endoglucanases mediate degradation of cellulose by Saccharophagus degradans.
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The prospects of cellulase-producing bacteria for the bioconversion of lignocellulosic biomass.
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18853469 O.V.Oliveira, L.C.Freitas, T.P.Straatsma, and R.D.Lins (2009).
Interaction between the CBM of Cel9A from Thermobifida fusca and cellulose fibers.
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19116695 R.E.Nordon, S.J.Craig, and F.C.Foong (2009).
Molecular engineering of the cellulosome complex for affinity and bioenergy applications.
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18378599 D.B.Wilson (2008).
Three microbial strategies for plant cell wall degradation.
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18832174 E.J.Dimise, P.F.Widboom, and S.D.Bruner (2008).
Structure elucidation and biosynthesis of fuscachelins, peptide siderophores from the moderate thermophile Thermobifida fusca.
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18553392 F.Moser, D.Irwin, S.Chen, and D.B.Wilson (2008).
Regulation and characterization of Thermobifida fusca carbohydrate-binding module proteins E7 and E8.
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18256495 K.Murata, S.Kawai, B.Mikami, and W.Hashimoto (2008).
Superchannel of bacteria: biological significance and new horizons.
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18663584 L.C.Tsai, Y.N.Chen, and L.F.Shyur (2008).
Structural modeling of glucanase-substrate complexes suggests a conserved tyrosine is involved in carbohydrate recognition in plant 1,3-1,4-beta-D-glucanases.
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18203823 M.Qi, H.S.Jun, and C.W.Forsberg (2008).
Cel9D, an atypical 1,4-beta-D-glucan glucohydrolase from Fibrobacter succinogenes: characteristics, catalytic residues, and synergistic interactions with other cellulases.
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17985078 T.Mejia-Castillo, M.E.Hidalgo-Lara, L.G.Brieba, and J.Ortega-Lopez (2008).
Purification, characterization and modular organization of a cellulose-binding protein, CBP105, a processive beta-1,4-endoglucanase from Cellulomonas flavigena.
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18367275 V.G.Eijsink, G.Vaaje-Kolstad, K.M.Vårum, and S.J.Horn (2008).
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18306418 Y.Li, and D.B.Wilson (2008).
Chitin binding by Thermobifida fusca cellulase catalytic domains.
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17187076 A.L.van Bueren, M.Higgins, D.Wang, R.D.Burke, and A.B.Boraston (2007).
Identification and structural basis of binding to host lung glycogen by streptococcal virulence factors.
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PDB codes: 2j43 2j44
17121820 D.K.Poon, S.G.Withers, and L.P.McIntosh (2007).
Direct demonstration of the flexibility of the glycosylated proline-threonine linker in the Cellulomonas fimi Xylanase Cex through NMR spectroscopic analysis.
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17905885 F.Mingardon, A.Chanal, C.Tardif, E.A.Bayer, and H.P.Fierobe (2007).
Exploration of new geometries in cellulosome-like chimeras.
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Crystallization and preliminary diffraction studies of CBM3b of cellobiohydrolase 9A from Clostridium thermocellum.
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17453195 S.Zhang, Q.Y.Yin, Y.H.Li, M.Ding, G.J.Xu, and F.K.Zhao (2007).
Molecular and biochemical characterization of Ba-EGA, a cellulase secreted by Bacillus sp. AC-1 from Ampullaria crosseans.
  Appl Microbiol Biotechnol, 75, 1327-1334.  
17369336 Y.Li, D.C.Irwin, and D.B.Wilson (2007).
Processivity, substrate binding, and mechanism of cellulose hydrolysis by Thermobifida fusca Cel9A.
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16527981 B.Li, J.P.Yu, J.S.Brunzelle, G.N.Moll, W.A.van der Donk, and S.K.Nair (2006).
Structure and mechanism of the lantibiotic cyclase involved in nisin biosynthesis.
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PDB codes: 2g02 2g0d
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15755956 A.L.Demain, M.Newcomb, and J.H.Wu (2005).
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15995203 S.O.Han, H.Yukawa, M.Inui, and R.H.Doi (2005).
Molecular cloning and transcriptional and expression analysis of engO, encoding a new noncellulosomal family 9 enzyme, from Clostridium cellulovorans.
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15849405 W.Hashimoto, K.Momma, Y.Maruyama, M.Yamasaki, B.Mikami, and K.Murata (2005).
Structure and function of bacterial super-biosystem responsible for import and depolymerization of macromolecules.
  Biosci Biotechnol Biochem, 69, 673-692.  
15073875 D.B.Wilson (2004).
Studies of Thermobifida fusca plant cell wall degrading enzymes.
  Chem Rec, 4, 72-82.  
15487947 E.A.Bayer, J.P.Belaich, Y.Shoham, and R.Lamed (2004).
The cellulosomes: multienzyme machines for degradation of plant cell wall polysaccharides.
  Annu Rev Microbiol, 58, 521-554.  
15604820 L.Hildén, and G.Johansson (2004).
Recent developments on cellulases and carbohydrate-binding modules with cellulose affinity.
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15211513 L.L.Videau, W.B.Arendall, and J.S.Richardson (2004).
The cis-Pro touch-turn: a rare motif preferred at functional sites.
  Proteins, 56, 298-309.  
14761991 Q.Xu, E.A.Bayer, M.Goldman, R.Kenig, Y.Shoham, and R.Lamed (2004).
Architecture of the Bacteroides cellulosolvens cellulosome: description of a cell surface-anchoring scaffoldin and a family 48 cellulase.
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15148314 T.Itoh, S.Akao, W.Hashimoto, B.Mikami, and K.Murata (2004).
Crystal structure of unsaturated glucuronyl hydrolase, responsible for the degradation of glycosaminoglycan, from Bacillus sp. GL1 at 1.8 A resolution.
  J Biol Chem, 279, 31804-31812.
PDB code: 1vd5
16233728 W.Hashimoto, M.Yamasaki, T.Itoh, K.Momma, B.Mikami, and K.Murata (2004).
Super-channel in bacteria: structural and functional aspects of a novel biosystem for the import and depolymerization of macromolecules.
  J Biosci Bioeng, 98, 399-413.  
12837787 D.Mandelman, A.Belaich, J.P.Belaich, N.Aghajari, H.Driguez, and R.Haser (2003).
X-Ray crystal structure of the multidomain endoglucanase Cel9G from Clostridium cellulolyticum complexed with natural and synthetic cello-oligosaccharides.
  J Bacteriol, 185, 4127-4135.
PDB codes: 1g87 1ga2 1k72 1kfg
12581217 K.Suzuki, T.Ojima, and K.Nishita (2003).
Purification and cDNA cloning of a cellulase from abalone Haliotis discus hannai.
  Eur J Biochem, 270, 771-778.  
12511483 R.Gilad, L.Rabinovich, S.Yaron, E.A.Bayer, R.Lamed, H.J.Gilbert, and Y.Shoham (2003).
CelI, a noncellulosomal family 9 enzyme from Clostridium thermocellum, is a processive endoglucanase that degrades crystalline cellulose.
  J Bacteriol, 185, 391-398.  
12511497 T.Arai, R.Araki, A.Tanaka, S.Karita, T.Kimura, K.Sakka, and K.Ohmiya (2003).
Characterization of a cellulase containing a family 30 carbohydrate-binding module (CBM) derived from Clostridium thermocellum CelJ: importance of the CBM to cellulose hydrolysis.
  J Bacteriol, 185, 504-512.  
12475987 W.Hashimoto, H.Nankai, B.Mikami, and K.Murata (2003).
Crystal structure of Bacillus sp. GL1 xanthan lyase, which acts on the side chains of xanthan.
  J Biol Chem, 278, 7663-7673.
PDB codes: 1j0m 1j0n
11844767 A.Belaich, G.Parsiegla, L.Gal, C.Villard, R.Haser, and J.P.Belaich (2002).
Cel9M, a new family 9 cellulase of the Clostridium cellulolyticum cellulosome.
  J Bacteriol, 184, 1378-1384.  
11958931 A.Toyoda, and H.Minato (2002).
Cloning, nucleotide sequence and expression of the gene encoding the cellulose-binding protein A (CBPA) of Eubacterium cellulosolvens 5.
  FEMS Microbiol Lett, 207, 141-146.  
12204381 A.Toyoda, and H.Minato (2002).
Identification of the cellulose-binding and the cell wall-binding domains of Eubacterium cellulosolvens 5 cellulose-binding protein A (CBPA).
  FEMS Microbiol Lett, 214, 113-118.  
12406778 A.V.Wymelenberg, S.Denman, D.Dietrich, J.Bassett, X.Yu, R.Atalla, P.Predki, U.Rudsander, T.T.Teeri, and D.Cullen (2002).
Transcript analysis of genes encoding a family 61 endoglucanase and a putative membrane-anchored family 9 glycosyl hydrolase from Phanerochaete chrysosporium.
  Appl Environ Microbiol, 68, 5765-5768.  
12200278 I.A.Kataeva, R.D.Seidel, A.Shah, L.T.West, X.L.Li, and L.G.Ljungdahl (2002).
The fibronectin type 3-like repeat from the Clostridium thermocellum cellobiohydrolase CbhA promotes hydrolysis of cellulose by modifying its surface.
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16233198 I.Maru, J.Ohnishi, Y.Ohta, and Y.Tsukada (2002).
Why is sialic acid attracting interest now? Complete enzymatic synthesis of sialic acid with N-acylglucosamine 2-epimerase.
  J Biosci Bioeng, 93, 258-265.  
11980476 M.Abou-Hachem, E.N.Karlsson, P.J.Simpson, S.Linse, P.Sellers, M.P.Williamson, S.J.Jamieson, H.J.Gilbert, D.N.Bolam, and O.Holst (2002).
Calcium binding and thermostability of carbohydrate binding module CBM4-2 of Xyn10A from Rhodothermus marinus.
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12186865 V.Receveur, M.Czjzek, M.Schülein, P.Panine, and B.Henrissat (2002).
Dimension, shape, and conformational flexibility of a two domain fungal cellulase in solution probed by small angle X-ray scattering.
  J Biol Chem, 277, 40887-40892.  
11481469 D.M.van Aalten, D.Komander, B.Synstad, S.Gåseidnes, M.G.Peter, and V.G.Eijsink (2001).
Structural insights into the catalytic mechanism of a family 18 exo-chitinase.
  Proc Natl Acad Sci U S A, 98, 8979-8984.
PDB codes: 1e6n 1e6p 1e6r 1e6z
11327868 D.N.Bolam, H.Xie, P.White, P.J.Simpson, S.M.Hancock, M.P.Williamson, and H.J.Gilbert (2001).
Evidence for synergy between family 2b carbohydrate binding modules in Cellulomonas fimi xylanase 11A.
  Biochemistry, 40, 2468-2477.
PDB codes: 1heh 1hej
11478884 H.Xie, H.J.Gilbert, S.J.Charnock, G.J.Davies, M.P.Williamson, P.J.Simpson, S.Raghothama, C.M.Fontes, F.M.Dias, L.M.Ferreira, and D.N.Bolam (2001).
Clostridium thermocellum Xyn10B carbohydrate-binding module 22-2: the role of conserved amino acids in ligand binding.
  Biochemistry, 40, 9167-9176.
PDB codes: 1h6x 1h6y
11160085 I.A.Kataeva, R.D.Seidel, X.L.Li, and L.G.Ljungdahl (2001).
Properties and mutation analysis of the CelK cellulose-binding domain from the Clostridium thermocellum cellulosome.
  J Bacteriol, 183, 1552-1559.  
11679332 M.L.Wu, Y.C.Chuang, J.P.Chen, C.S.Chen, and M.C.Chang (2001).
Identification and characterization of the three chitin-binding domains within the multidomain chitinase Chi92 from Aeromonas hydrophila JP101.
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11514661 W.A.Breyer, and B.W.Matthews (2001).
A structural basis for processivity.
  Protein Sci, 10, 1699-1711.  
10714996 C.Gaudin, A.Belaich, S.Champ, and J.P.Belaich (2000).
CelE, a multidomain cellulase from Clostridium cellulolyticum: a key enzyme in the cellulosome?
  J Bacteriol, 182, 1910-1915.  
11106416 C.I.Avitia, F.X.Castellanos-Juárez, E.Sánchez, A.Téllez-Valencia, P.Fajardo-Cavazos, W.L.Nicholson, and M.Pedraza-Reyes (2000).
Temporal secretion of a multicellulolytic system in Myxobacter sp. AL-1. Molecular cloning and heterologous expression of cel9 encoding a modular endocellulase clustered in an operon with cel48, an exocellobiohydrolase gene.
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10823940 D.M.van Aalten, B.Synstad, M.B.Brurberg, E.Hough, B.W.Riise, V.G.Eijsink, and R.K.Wierenga (2000).
Structure of a two-domain chitotriosidase from Serratia marcescens at 1.9-A resolution.
  Proc Natl Acad Sci U S A, 97, 5842-5847.
PDB code: 1e15
10985769 G.Parsiegla, C.Reverbel-Leroy, C.Tardif, J.P.Belaich, H.Driguez, and R.Haser (2000).
Crystal structures of the cellulase Cel48F in complex with inhibitors and substrates give insights into its processive action.
  Biochemistry, 39, 11238-11246.
PDB codes: 1f9d 1f9o 1fae 1fbo 1fbw
10913296 J.Kormos, P.E.Johnson, E.Brun, P.Tomme, L.P.McIntosh, C.A.Haynes, and D.G.Kilburn (2000).
Binding site analysis of cellulose binding domain CBD(N1) from endoglucanse C of Cellulomonas fimi by site-directed mutagenesis.
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10819965 S.J.Charnock, D.N.Bolam, J.P.Turkenburg, H.J.Gilbert, L.M.Ferreira, G.J.Davies, and C.M.Fontes (2000).
The X6 "thermostabilizing" domains of xylanases are carbohydrate-binding modules: structure and biochemistry of the Clostridium thermocellum X6b domain.
  Biochemistry, 39, 5013-5021.
PDB code: 1dyo
10824094 S.Zhang, D.C.Irwin, and D.B.Wilson (2000).
Site-directed mutation of noncatalytic residues of Thermobifida fusca exocellulase Cel6B.
  Eur J Biochem, 267, 3101-3115.  
10788483 T.Ikegami, T.Okada, M.Hashimoto, S.Seino, T.Watanabe, and M.Shirakawa (2000).
Solution structure of the chitin-binding domain of Bacillus circulans WL-12 chitinase A1.
  J Biol Chem, 275, 13654-13661.
PDB code: 1ed7
11018131 T.Y.Wong, L.A.Preston, and N.L.Schiller (2000).
ALGINATE LYASE: review of major sources and enzyme characteristics, structure-function analysis, biological roles, and applications.
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  10210191 D.H.Juers, R.E.Huber, and B.W.Matthews (1999).
Structural comparisons of TIM barrel proteins suggest functional and evolutionary relationships between beta-galactosidase and other glycohydrolases.
  Protein Sci, 8, 122-136.  
10542312 G.Tokuda, N.Lo, H.Watanabe, M.Slaytor, T.Matsumoto, and H.Noda (1999).
Metazoan cellulase genes from termites: intron/exon structures and sites of expression.
  Biochim Biophys Acta, 1447, 146-159.  
  10419961 H.D.Simpson, and F.Barras (1999).
Functional analysis of the carbohydrate-binding domains of Erwinia chrysanthemi Cel5 (Endoglucanase Z) and an Escherichia coli putative chitinase.
  J Bacteriol, 181, 4611-4616.  
10548733 K.A.Byrne, S.A.Lehnert, S.E.Johnson, and S.S.Moore (1999).
Isolation of a cDNA encoding a putative cellulase in the red claw crayfish Cherax quadricarinatus.
  Gene, 239, 317-324.  
10224066 N.A.Spiridonov, and D.B.Wilson (1999).
Characterization and cloning of celR, a transcriptional regulator of cellulase genes from Thermomonospora fusca.
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10425686 P.J.Simpson, D.N.Bolam, A.Cooper, A.Ciruela, G.P.Hazlewood, H.J.Gilbert, and M.P.Williamson (1999).
A family IIb xylan-binding domain has a similar secondary structure to a homologous family IIa cellulose-binding domain but different ligand specificity.
  Structure, 7, 853-864.
PDB codes: 1xbd 2xbd
10391926 S.Kawaminami, H.Takahashi, S.Ito, Y.Arata, and I.Shimada (1999).
A multinuclear NMR study of the active site of an endoglucanase from a strain of Bacillus. Use of Trp residues as structural probes.
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  10542174 S.Y.Ding, E.A.Bayer, D.Steiner, Y.Shoham, and R.Lamed (1999).
A novel cellulosomal scaffoldin from Acetivibrio cellulolyticus that contains a family 9 glycosyl hydrolase.
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10390637 Y.Shoham, R.Lamed, and E.A.Bayer (1999).
The cellulosome concept as an efficient microbial strategy for the degradation of insoluble polysaccharides.
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9649302 B.K.Barr, D.E.Wolfgang, K.Piens, M.Claeyssens, and D.B.Wilson (1998).
Active-site binding of glycosides by Thermomonospora fusca endocellulase E2.
  Biochemistry, 37, 9220-9229.  
  9537366 D.Irwin, D.H.Shin, S.Zhang, B.K.Barr, J.Sakon, P.A.Karplus, and D.B.Wilson (1998).
Roles of the catalytic domain and two cellulose binding domains of Thermomonospora fusca E4 in cellulose hydrolysis.
  J Bacteriol, 180, 1709-1714.  
9818257 E.A.Bayer, H.Chanzy, R.Lamed, and Y.Shoham (1998).
Cellulose, cellulases and cellulosomes.
  Curr Opin Struct Biol, 8, 548-557.  
9755156 G.Parsiegla, M.Juy, C.Reverbel-Leroy, C.Tardif, J.P.Belaïch, H.Driguez, and R.Haser (1998).
The crystal structure of the processive endocellulase CelF of Clostridium cellulolyticum in complex with a thiooligosaccharide inhibitor at 2.0 A resolution.
  EMBO J, 17, 5551-5562.
PDB code: 1fce
9682475 K.Riedel, J.Ritter, S.Bauer, and K.Bronnenmeier (1998).
The modular cellulase CelZ of the thermophilic bacterium Clostridium stercorarium contains a thermostabilizing domain.
  FEMS Microbiol Lett, 164, 261-267.  
9643544 K.Riedel, and K.Bronnenmeier (1998).
Intramolecular synergism in an engineered exo-endo-1,4-beta-glucanase fusion protein.
  Mol Microbiol, 28, 767-775.  
9519404 K.U.Wendt, and G.E.Schulz (1998).
Isoprenoid biosynthesis: manifold chemistry catalyzed by similar enzymes.
  Structure, 6, 127-133.  
  9657993 N.A.Spiridonov, and D.B.Wilson (1998).
Regulation of biosynthesis of individual cellulases in Thermomonospora fusca.
  J Bacteriol, 180, 3529-3532.  
9737854 P.E.Johnson, A.L.Creagh, E.Brun, K.Joe, P.Tomme, C.A.Haynes, and L.P.McIntosh (1998).
Calcium binding by the N-terminal cellulose-binding domain from Cellulomonas fimi beta-1,4-glucanase CenC.
  Biochemistry, 37, 12772-12781.  
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