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J. Biol. Chem., Vol. 281, Issue 34, 24922-24933, August 25, 2006

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Crystal Structures of Clostridium thermocellum Xyloglucanase, XGH74A, Reveal the Structural Basis for Xyloglucan Recognition and Degradation^*

Carlos Martinez-Fleites¹, Catarina I. P. D. Guerreiro¹, Martin J. Baumann^¶1, Edward J. Taylor, José A. M. Prates, Luís M. A. Ferreira, Carlos M. G. A. Fontes, Harry Brumer^¶2, and Gideon J. Davies³

From the York Structural Biology Laboratory, Department of Chemitry, University of York, York YO10 5YW, United Kingdom, the CIISA, Faculdade de Medicina Veterinária, Universidade Técnica de Lisboa, Avenida da Universidade Técnica, 1300-477 Lisboa, Portugal, and the ^¶School of Biotechnology, Royal Institute of Technology (KTH), AlbaNova University Centre, 106 91 Stockholm, Sweden

The enzymatic degradation of the plant cell wall is central both to the natural carbon cycle and, increasingly, to environmentally friendly routes to biomass conversion, including the production of biofuels. The plant cell wall is a complex composite of cellulose microfibrils embedded in diverse polysaccharides collectively termed hemicelluloses. Xyloglucan is one such polysaccharide whose hydrolysis is catalyzed by diverse xyloglucanases. Here we present the structure of the Clostridium thermocellum xyloglucanase Xgh74A in both apo and ligand-complexed forms. The structures, in combination with mutagenesis data on the catalytic residues and the kinetics and specificity of xyloglucan hydrolysis reveal a complex subsite specificity accommodating seventeen monosaccharide moieties of the multibranched substrate in an open substrate binding terrain.

Received for publication, April 13, 2006 , and in revised form, June 12, 2006.

The atomic coordinates and structure factors (codes 2CN2 and 2CN3) have been deposited in the Protein Data Bank, Research Collaboratory for Structural Bioinformatics, Rutgers University, New Brunswick, NJ (http://www.rcsb.org/).

^* This work was funded by the Biotechnology and Biological Sciences Research Council (BBSRC), the Swedish Foundation for Strategic Research, the KTH Biofibre Materials Centre, and the Fundação para a Ciência e Tecnologia. The costs of publication of this article were defrayed in part by the payment of page charges. This article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.

¹ These authors contributed equally to this work.

² Fellow (Rådsforskare) of the Swedish Research Council.

³ To whom correspondence should be addressed. E-mail: davies{at}ysbl.york.ac.uk.

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