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Title: The Dissociation Mechanism of Processive Cellulases

Abstract

Cellulase enzymes deconstruct recalcitrant cellulose into soluble sugars, making them a biocatalyst of biotechnological interest for use in the nascent lignocellulosic bioeconomy. Cellobiohydrolases (CBHs) are cellulases capable of liberating many sugar molecules in a processive manner without dissociating from the substrate. Within the complete processive cycle of CBHs, dissociation from the cellulose substrate is rate limiting, but the molecular mechanism of this step is unknown. Here, we present a direct comparison of potential molecular mechanisms for dissociation via Hamiltonian replica exchange molecular dynamics of the model fungal CBH, Trichoderma reesei Cel7A. Computational rate estimates indicate that stepwise cellulose dethreading from the binding tunnel is 4 orders of magnitude faster than a clamshell mechanism, in which the substrate-enclosing loops open and release the substrate without reversing. We also present the crystal structure of a disulfide variant that covalently links substrate-enclosing loops on either side of the substrate-binding tunnel, which constitutes a CBH that can only dissociate via stepwise dethreading. Biochemical measurements indicate that this variant has a dissociation rate constant essentially equivalent to the wild type, implying that dethreading is likely the predominant mechanism for dissociation.

Authors:
ORCiD logo [1];  [2]; ORCiD logo [1];  [1];  [3];  [3];  [3];  [2];  [1]
+ Show Author Affiliations
  1. National Renewable Energy Laboratory (NREL), Golden, CO (United States)
  2. Univ. of Tartu, Tartu (Estonia)
  3. Swedish Univ. of Agricultural Sciences, Uppsala (Sweden)
Publication Date:
Research Org.:
National Renewable Energy Laboratory (NREL), Golden, CO (United States)
Sponsoring Org.:
USDOE Office of Energy Efficiency and Renewable Energy (EERE), Sustainable Transportation Office. Bioenergy Technologies Office
OSTI Identifier:
1574196
Report Number(s):
NREL/JA-2700-74624
Journal ID: ISSN 0027-8424
Grant/Contract Number:  
AC36-08GO28308
Resource Type:
Accepted Manuscript
Journal Name:
Proceedings of the National Academy of Sciences of the United States of America
Additional Journal Information:
Journal Volume: 116; Journal Issue: 46; Journal ID: ISSN 0027-8424
Publisher:
National Academy of Sciences
Country of Publication:
United States
Language:
English
Subject:
09 BIOMASS FUELS; biofuels; cellulases; molecular mechanism; molecular simulation; crystal structure; BCPL

Citation Formats

Vermaas, Joshua, Kont, Riin, Beckham, Gregg T, Crowley, Michael F, Gudmundsson, Mikael, Sandgren, Mats, Stahlberg, Jerry, Valjamae, Priit, and Knott, Brandon C. The Dissociation Mechanism of Processive Cellulases. United States: N. p., 2019. Web. doi:10.1073/pnas.1913398116.
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Vermaas, Joshua, Kont, Riin, Beckham, Gregg T, Crowley, Michael F, Gudmundsson, Mikael, Sandgren, Mats, Stahlberg, Jerry, Valjamae, Priit, & Knott, Brandon C. The Dissociation Mechanism of Processive Cellulases. United States. https://doi.org/10.1073/pnas.1913398116
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Vermaas, Joshua, Kont, Riin, Beckham, Gregg T, Crowley, Michael F, Gudmundsson, Mikael, Sandgren, Mats, Stahlberg, Jerry, Valjamae, Priit, and Knott, Brandon C. Tue . "The Dissociation Mechanism of Processive Cellulases". United States. https://doi.org/10.1073/pnas.1913398116. https://www.osti.gov/servlets/purl/1574196.
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@article{osti_1574196,
title = {The Dissociation Mechanism of Processive Cellulases},
author = {Vermaas, Joshua and Kont, Riin and Beckham, Gregg T and Crowley, Michael F and Gudmundsson, Mikael and Sandgren, Mats and Stahlberg, Jerry and Valjamae, Priit and Knott, Brandon C},
abstractNote = {Cellulase enzymes deconstruct recalcitrant cellulose into soluble sugars, making them a biocatalyst of biotechnological interest for use in the nascent lignocellulosic bioeconomy. Cellobiohydrolases (CBHs) are cellulases capable of liberating many sugar molecules in a processive manner without dissociating from the substrate. Within the complete processive cycle of CBHs, dissociation from the cellulose substrate is rate limiting, but the molecular mechanism of this step is unknown. Here, we present a direct comparison of potential molecular mechanisms for dissociation via Hamiltonian replica exchange molecular dynamics of the model fungal CBH, Trichoderma reesei Cel7A. Computational rate estimates indicate that stepwise cellulose dethreading from the binding tunnel is 4 orders of magnitude faster than a clamshell mechanism, in which the substrate-enclosing loops open and release the substrate without reversing. We also present the crystal structure of a disulfide variant that covalently links substrate-enclosing loops on either side of the substrate-binding tunnel, which constitutes a CBH that can only dissociate via stepwise dethreading. Biochemical measurements indicate that this variant has a dissociation rate constant essentially equivalent to the wild type, implying that dethreading is likely the predominant mechanism for dissociation.},
doi = {10.1073/pnas.1913398116},
journal = {Proceedings of the National Academy of Sciences of the United States of America},
number = 46,
volume = 116,
place = {United States},
year = {Tue Nov 12 00:00:00 EST 2019},
month = {Tue Nov 12 00:00:00 EST 2019}
}
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https://doi.org/10.1073/pnas.1913398116
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Works referenced in this record:

Trichoderma reesei has no true exo-cellulase: all intact and truncated cellulases produce new reducing end groups on cellulose
journal, May 1993

  • Ståhlberg, Jerry; Johansson, Gunnar; Pettersson, Göran
  • Biochimica et Biophysica Acta (BBA) - General Subjects, Vol. 1157, Issue 1
  • DOI: 10.1016/0304-4165(93)90085-M

Mechanism of lignin inhibition of enzymatic biomass deconstruction
journal, December 2015

  • Vermaas, Josh V.; Petridis, Loukas; Qi, Xianghong
  • Biotechnology for Biofuels, Vol. 8, Issue 1
  • DOI: 10.1186/s13068-015-0379-8

Free Energy Diagram for the Heterogeneous Enzymatic Hydrolysis of Glycosidic Bonds in Cellulose
journal, July 2015

  • Sørensen, Trine Holst; Cruys-Bagger, Nicolaj; Borch, Kim
  • Journal of Biological Chemistry, Vol. 290, Issue 36
  • DOI: 10.1074/jbc.M115.659656

Fungal Cellulases
journal, January 2015

  • Payne, Christina M.; Knott, Brandon C.; Mayes, Heather B.
  • Chemical Reviews, Vol. 115, Issue 3
  • DOI: 10.1021/cr500351c

Cellulases and biofuels
journal, June 2009

Kinetics of Cellobiohydrolase (Cel7A) Variants with Lowered Substrate Affinity
journal, September 2014

  • Kari, Jeppe; Olsen, Johan; Borch, Kim
  • Journal of Biological Chemistry, Vol. 289, Issue 47
  • DOI: 10.1074/jbc.M114.604264

Slow Off-rates and Strong Product Binding Are Required for Processivity and Efficient Degradation of Recalcitrant Chitin by Family 18 Chitinases
journal, October 2015

  • Kurašin, Mihhail; Kuusk, Silja; Kuusk, Piret
  • Journal of Biological Chemistry, Vol. 290, Issue 48
  • DOI: 10.1074/jbc.M115.684977

Spontaneous Hydrolysis of Glycosides
journal, July 1998

  • Wolfenden, Richard; Lu, Xiangdong; Young, Gregory
  • Journal of the American Chemical Society, Vol. 120, Issue 27
  • DOI: 10.1021/ja9813055

Catalytic routes for the conversion of biomass into liquid hydrocarbon transportation fuels
journal, January 2011

  • Serrano-Ruiz, Juan Carlos; Dumesic, James A.
  • Energy Environ. Sci., Vol. 4, Issue 1
  • DOI: 10.1039/C0EE00436G

Scalable molecular dynamics with NAMD
journal, January 2005

  • Phillips, James C.; Braun, Rosemary; Wang, Wei
  • Journal of Computational Chemistry, Vol. 26, Issue 16, p. 1781-1802
  • DOI: 10.1002/jcc.20289

Molecular-Level Origins of Biomass Recalcitrance: Decrystallization Free Energies for Four Common Cellulose Polymorphs
journal, April 2011

  • Beckham, Gregg T.; Matthews, James F.; Peters, Baron
  • The Journal of Physical Chemistry B, Vol. 115, Issue 14
  • DOI: 10.1021/jp1106394

Mechanism of initial rapid rate retardation in cellobiohydrolase catalyzed cellulose hydrolysis
journal, April 2010

  • Jalak, Jürgen; Väljamäe, Priit
  • Biotechnology and Bioengineering, Vol. 106, Issue 6
  • DOI: 10.1002/bit.22779

Cello-Oligosaccharide Hydrolysis by Cellobiohydrolase II from Trichoderma Reesei. Association and Rate Constants Derived from an Analysis of Progress Curves
journal, September 1996

High Speed Atomic Force Microscopy Visualizes Processive Movement of Trichoderma reesei Cellobiohydrolase I on Crystalline Cellulose
journal, October 2009

  • Igarashi, Kiyohiko; Koivula, Anu; Wada, Masahisa
  • Journal of Biological Chemistry, Vol. 284, Issue 52
  • DOI: 10.1074/jbc.M109.034611

The Mechanism of Cellulose Hydrolysis by a Two-Step, Retaining Cellobiohydrolase Elucidated by Structural and Transition Path Sampling Studies
journal, December 2013

  • Knott, Brandon C.; Haddad Momeni, Majid; Crowley, Michael F.
  • Journal of the American Chemical Society, Vol. 136, Issue 1
  • DOI: 10.1021/ja410291u

Observing and modeling BMCC degradation by commercial cellulase cocktails with fluorescently labeled Trichoderma reseii Cel7A through confocal microscopy
journal, July 2012

  • Luterbacher, Jeremy S.; Walker, Larry P.; Moran-Mirabal, Jose M.
  • Biotechnology and Bioengineering, Vol. 110, Issue 1
  • DOI: 10.1002/bit.24597

Biomass Recalcitrance: Engineering Plants and Enzymes for Biofuels Production
journal, February 2007

  • Himmel, M. E.; Ding, S.-Y.; Johnson, D. K.
  • Science, Vol. 315, Issue 5813, p. 804-807
  • DOI: 10.1126/science.1137016

Site-directed mutation of noncatalytic residues of Thermobifida fusca exocellulase Cel6B: Mutagenesis of Thermobifida fusca exocellulase Cel6B
journal, June 2000

Time to Rethink Cellulosic Biofuels?
journal, January 2018

  • Dale, Bruce
  • Biofuels, Bioproducts and Biorefining, Vol. 12, Issue 1
  • DOI: 10.1002/bbb.1856

Activity Studies and Crystal Structures of Catalytically Deficient Mutants of Cellobiohydrolase I fromTrichoderma reesei
journal, November 1996

  • Ståhlberg, Jerry; Divne, Christina; Koivula, Anu
  • Journal of Molecular Biology, Vol. 264, Issue 2
  • DOI: 10.1006/jmbi.1996.0644

Degrees of Difficulty of Water-Consuming Reactions in the Absence of Enzymes
journal, August 2006

XSEDE: Accelerating Scientific Discovery
journal, September 2014

  • Towns, John; Cockerill, Timothy; Dahan, Maytal
  • Computing in Science & Engineering, Vol. 16, Issue 5
  • DOI: 10.1109/MCSE.2014.80

Microbial engineering for the production of advanced biofuels
journal, August 2012

  • Peralta-Yahya, Pamela P.; Zhang, Fuzhong; del Cardayre, Stephen B.
  • Nature, Vol. 488, Issue 7411
  • DOI: 10.1038/nature11478

Single-molecule Imaging Analysis of Elementary Reaction Steps of Trichoderma reesei Cellobiohydrolase I (Cel7A) Hydrolyzing Crystalline Cellulose I α and III I
journal, April 2014

  • Shibafuji, Yusuke; Nakamura, Akihiko; Uchihashi, Takayuki
  • Journal of Biological Chemistry, Vol. 289, Issue 20
  • DOI: 10.1074/jbc.M113.546085

Effects of Lytic Polysaccharide Monooxygenase Oxidation on Cellulose Structure and Binding of Oxidized Cellulose Oligomers to Cellulases
journal, April 2015

  • Vermaas, Josh V.; Crowley, Michael F.; Beckham, Gregg T.
  • The Journal of Physical Chemistry B, Vol. 119, Issue 20
  • DOI: 10.1021/acs.jpcb.5b00778

Improving the thermal stability of cellobiohydrolase Cel7A from Hypocrea jecorina by directed evolution
journal, August 2017

  • Goedegebuur, Frits; Dankmeyer, Lydia; Gualfetti, Peter
  • Journal of Biological Chemistry, Vol. 292, Issue 42
  • DOI: 10.1074/jbc.M117.803270

CHARMM: The biomolecular simulation program
journal, July 2009

  • Brooks, B. R.; Brooks, C. L.; Mackerell, A. D.
  • Journal of Computational Chemistry, Vol. 30, Issue 10
  • DOI: 10.1002/jcc.21287

Structural insights into the inhibition of cellobiohydrolase Cel7A by xylo-oligosaccharides
journal, April 2015

  • Momeni, Majid Haddad; Ubhayasekera, Wimal; Sandgren, Mats
  • FEBS Journal, Vol. 282, Issue 11
  • DOI: 10.1111/febs.13265

Predominant Nonproductive Substrate Binding by Fungal Cellobiohydrolase I and Implications for Activity Improvement
journal, June 2018

  • Rabinovich, Mikhail L.; Melnik, Maria S.; Herner, Mikhail L.
  • Biotechnology Journal, Vol. 14, Issue 3
  • DOI: 10.1002/biot.201700712

Deconstruction of Lignocellulosic Biomass to Fuels and Chemicals
journal, July 2011

Expression, crystal structure and cellulase activity of the thermostable cellobiohydrolase Cel7A from the fungus Humicola grisea var. thermoidea
journal, August 2014

  • Momeni, Majid Haddad; Goedegebuur, Frits; Hansson, Henrik
  • Acta Crystallographica Section D Biological Crystallography, Vol. 70, Issue 9
  • DOI: 10.1107/S1399004714013844

Stereochemistry and the Mechanism of Enzymatic Reactions
journal, November 1953

Sabatier Principle for Interfacial (Heterogeneous) Enzyme Catalysis
journal, October 2018

Cellulose: Fascinating Biopolymer and Sustainable Raw Material
journal, May 2005

  • Klemm, Dieter; Heublein, Brigitte; Fink, Hans-Peter
  • Angewandte Chemie International Edition, Vol. 44, Issue 22
  • DOI: 10.1002/anie.200460587

Transient Kinetics and Rate-Limiting Steps for the Processive Cellobiohydrolase Cel7A: Effects of Substrate Structure and Carbohydrate Binding Domain
journal, November 2013

  • Cruys-Bagger, Nicolaj; Tatsumi, Hirosuke; Ren, Guilin Robin
  • Biochemistry, Vol. 52, Issue 49
  • DOI: 10.1021/bi401210n

Trade-off between Processivity and Hydrolytic Velocity of Cellobiohydrolases at the Surface of Crystalline Cellulose
journal, March 2014

  • Nakamura, Akihiko; Watanabe, Hiroki; Ishida, Takuya
  • Journal of the American Chemical Society, Vol. 136, Issue 12
  • DOI: 10.1021/ja4119994

Probing Carbohydrate Product Expulsion from a Processive Cellulase with Multiple Absolute Binding Free Energy Methods
journal, March 2011

  • Bu, Lintao; Beckham, Gregg T.; Shirts, Michael R.
  • Journal of Biological Chemistry, Vol. 286, Issue 20
  • DOI: 10.1074/jbc.M110.212076

Deconstruction of lignocellulosic biomass with ionic liquids
journal, January 2013

  • Brandt, Agnieszka; Gräsvik, John; Hallett, Jason P.
  • Green Chemistry, Vol. 15, Issue 3
  • DOI: 10.1039/c2gc36364j

Engineering the Exo-loop of Trichoderma reesei Cellobiohydrolase, Cel7A. A comparison with Phanerochaete chrysosporium Cel7D
journal, October 2003

  • von Ossowski, Ingemar; Ståhlberg, Jerry; Koivula, Anu
  • Journal of Molecular Biology, Vol. 333, Issue 4, p. 817-829
  • DOI: 10.1016/S0022-2836(03)00881-7

Inter-domain Synergism Is Required for Efficient Feeding of Cellulose Chain into Active Site of Cellobiohydrolase Cel7A
journal, October 2016

  • Kont, Riin; Kari, Jeppe; Borch, Kim
  • Journal of Biological Chemistry, Vol. 291, Issue 50
  • DOI: 10.1074/jbc.M116.756007

Processive action of cellobiohydrolase Cel7A from Trichoderma reesei is revealed as ‘burst’ kinetics on fluorescent polymeric model substrates
journal, January 2005

  • Kipper, Kalle; VÄLjamÄE, Priit; Johansson, Gunnar
  • Biochemical Journal, Vol. 385, Issue 2
  • DOI: 10.1042/BJ20041144

Hydrolysis of cellooligosaccharides by Trichoderma reesei cellobiohydrolases: Experimental data and kinetic modeling
journal, January 1994

Dynamics of reactions involving diffusive barrier crossing
journal, April 1981

  • Schulten, Klaus; Schulten, Zan; Szabo, Attila
  • The Journal of Chemical Physics, Vol. 74, Issue 8
  • DOI: 10.1063/1.441684

Systematic deletions in the cellobiohydrolase (CBH) Cel7A from the fungus Trichoderma reesei reveal flexible loops critical for CBH activity
journal, December 2018

  • Schiano-di-Cola, Corinna; Røjel, Nanna; Jensen, Kenneth
  • Journal of Biological Chemistry, Vol. 294, Issue 6
  • DOI: 10.1074/jbc.RA118.006699

Exo-exo synergy between Cel6A and Cel7A from Hypocrea jecorina : Role of carbohydrate binding module and the endo-lytic character of the enzymes : Synergy Between Cel6A and Cel7A
journal, March 2017

  • Badino, Silke F.; Christensen, Stefan J.; Kari, Jeppe
  • Biotechnology and Bioengineering, Vol. 114, Issue 8
  • DOI: 10.1002/bit.26276

Thermoactivation of a cellobiohydrolase
journal, January 2018

  • Westh, Peter; Borch, Kim; Sørensen, Trine
  • Biotechnology and Bioengineering, Vol. 115, Issue 4
  • DOI: 10.1002/bit.26513

Costs and benefits of processivity in enzymatic degradation of recalcitrant polysaccharides
journal, November 2006

  • Horn, S. J.; Sikorski, P.; Cederkvist, J. B.
  • Proceedings of the National Academy of Sciences, Vol. 103, Issue 48
  • DOI: 10.1073/pnas.0608909103

Glycoside Hydrolase Processivity Is Directly Related to Oligosaccharide Binding Free Energy
journal, December 2013

  • Payne, Christina M.; Jiang, Wei; Shirts, Michael R.
  • Journal of the American Chemical Society, Vol. 135, Issue 50
  • DOI: 10.1021/ja407287f

A distributed cellulosic biorefinery system in the US Midwest based on corn stover
journal, September 2016

  • Kim, Seungdo; Dale, Bruce E.
  • Biofuels, Bioproducts and Biorefining, Vol. 10, Issue 6
  • DOI: 10.1002/bbb.1712

Loop variants of the thermophile Rasamsonia emersonii Cel7A with improved activity against cellulose : Loop Variants of the Thermophile
journal, September 2016

  • Sørensen, Trine Holst; Windahl, Michael Skovbo; McBrayer, Brett
  • Biotechnology and Bioengineering, Vol. 114, Issue 1
  • DOI: 10.1002/bit.26050

Carbohydrate–Protein Interactions That Drive Processive Polysaccharide Translocation in Enzymes Revealed from a Computational Study of Cellobiohydrolase Processivity
journal, June 2014

  • Knott, Brandon C.; Crowley, Michael F.; Himmel, Michael E.
  • Journal of the American Chemical Society, Vol. 136, Issue 24
  • DOI: 10.1021/ja504074g

Biofuels 2020: Biorefineries based on lignocellulosic materials
journal, July 2016

  • Valdivia, Miguel; Galan, Jose Luis; Laffarga, Joaquina
  • Microbial Biotechnology, Vol. 9, Issue 5
  • DOI: 10.1111/1751-7915.12387

Towards a molecular-level theory of carbohydrate processivity in glycoside hydrolases
journal, June 2014

Concerted motions and large-scale structural fluctuations of Trichoderma reesei Cel7A cellobiohydrolase
journal, January 2018

  • Silveira, Rodrigo L.; Skaf, Munir S.
  • Physical Chemistry Chemical Physics, Vol. 20, Issue 11
  • DOI: 10.1039/C8CP00101D

A steady-state theory for processive cellulases
journal, July 2013

  • Cruys-Bagger, Nicolaj; Elmerdahl, Jens; Praestgaard, Eigil
  • FEBS Journal, Vol. 280, Issue 16
  • DOI: 10.1111/febs.12397

Endo-exo Synergism in Cellulose Hydrolysis Revisited
journal, June 2012

  • Jalak, Jürgen; Kurašin, Mihhail; Teugjas, Hele
  • Journal of Biological Chemistry, Vol. 287, Issue 34
  • DOI: 10.1074/jbc.M112.381624

Cellobiohydrolase 1 from Trichoderma reesei degrades cellulose in single cellobiose steps
journal, December 2015

  • Brady, Sonia K.; Sreelatha, Sarangapani; Feng, Yinnian
  • Nature Communications, Vol. 6, Issue 1
  • DOI: 10.1038/ncomms10149

Lignocellulose degradation mechanisms across the Tree of Life
journal, December 2015

Pre-steady-state Kinetics for Hydrolysis of Insoluble Cellulose by Cellobiohydrolase Cel7A
journal, April 2012

  • Cruys-Bagger, Nicolaj; Elmerdahl, Jens; Praestgaard, Eigil
  • Journal of Biological Chemistry, Vol. 287, Issue 22
  • DOI: 10.1074/jbc.M111.334946

Anomeric Selectivity and Product Profile of a Processive Cellulase
journal, December 2016

Molecular Dynamics Simulations of Family 7 Cellobiohydrolase Mutants Aimed at Reducing Product Inhibition
journal, September 2014

  • Silveira, Rodrigo L.; Skaf, Munir S.
  • The Journal of Physical Chemistry B, Vol. 119, Issue 29
  • DOI: 10.1021/jp509911m

Processivity of Cellobiohydrolases Is Limited by the Substrate
journal, November 2010

  • Kurašin, Mihhail; Väljamäe, Priit
  • Journal of Biological Chemistry, Vol. 286, Issue 1
  • DOI: 10.1074/jbc.M110.161059

Traffic Jams Reduce Hydrolytic Efficiency of Cellulase on Cellulose Surface
journal, September 2011

Structural, Biochemical, and Computational Characterization of the Glycoside Hydrolase Family 7 Cellobiohydrolase of the Tree-killing Fungus Heterobasidion irregulare
journal, January 2013

  • Momeni, Majid Haddad; Payne, Christina M.; Hansson, Henrik
  • Journal of Biological Chemistry, Vol. 288, Issue 8
  • DOI: 10.1074/jbc.M112.440891

Distinct roles of N- and O-glycans in cellulase activity and stability
journal, December 2017

  • Amore, Antonella; Knott, Brandon C.; Supekar, Nitin T.
  • Proceedings of the National Academy of Sciences, Vol. 114, Issue 52
  • DOI: 10.1073/pnas.1714249114

Degrees of Difficulty of Water-Consuming Reactions in the Absence of Enzymes
journal, October 2006

Effects of Lytic Polysaccharide Monooxygenase Oxidation on Cellulose Structure and Binding of Oxidized Cellulose Oligomers to Cellulases
journal, January 2013

  • Vermaas, Joshua V.; Payne, Christina M.; Crowley, Michael F.
  • Biophysical Journal, Vol. 104, Issue 2
  • DOI: 10.1016/j.bpj.2012.11.658

Fungal cellulases
journal, February 1992

  • Wood, Thomas M.
  • Biochemical Society Transactions, Vol. 20, Issue 1
  • DOI: 10.1042/bst0200046

Cellulose: Fascinating Biopolymer and Sustainable Raw Material
journal, September 2005

  • Klemm, Dieter; Heublein, Brigitte; Fink, Hans-Peter
  • ChemInform, Vol. 36, Issue 36
  • DOI: 10.1002/chin.200536238

CHARMM: the biomolecular simulation program
text, January 2009

  • Brooks, B. R.; Brooks, C. L.; Mackerell, A. D.
  • Wiley-Blackwell
  • DOI: 10.5167/uzh-19245
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