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The structure, determined for the 40-kDa precursor form of SpeB at 1.6-\u212b resolution, reveals that the protein is a distant homologue of the papain superfamily that includes the mammalian cathepsins B, K, L, and S. Despite negligible sequence identity, the protease portion has the canonical papain fold, albeit with major loop insertions and deletions. The catalytic site differs from most other cysteine proteases in that it lacks the Asn residue of the Cys-His-Asn triad. The prosegment has a unique fold and inactivation mechanism that involves displacement of the catalytically essential His residue by a loop inserted into the active site. The structure also reveals the surface location of an integrin-binding Arg-Gly-Asp (RGD) motif that is a feature unique to SpeB among cysteine proteases and is linked to the pathogenesis of the most invasive strains of\n                    <jats:italic>S. pyogenes<\/jats:italic>\n                    .\n                  <\/jats:p>","DOI":"10.1073\/pnas.040549997","type":"journal-article","created":{"date-parts":[[2002,7,26]],"date-time":"2002-07-26T10:41:48Z","timestamp":1027680108000},"page":"2235-2240","update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":93,"title":["Crystal structure of the zymogen form of the group A\n                    <i>Streptococcus<\/i>\n                    virulence factor SpeB: An integrin-binding cysteine protease"],"prefix":"10.1073","volume":"97","author":[{"given":"Todd F.","family":"Kagawa","sequence":"first","affiliation":[{"name":"School of Biological Sciences and Department of Chemistry, University of Auckland, Private Bag 92\u2013019, Auckland, New Zealand; Institute of Molecular Biosciences, Massey University, Palmerston North, New Zealand; European Molecular Biology Laboratory Grenoble Outstation, 6 Av. J. Horowitz, Grenoble 38042, France; Institute of Human Bacterial Pathogenesis, Department of Pathology, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030; and Laboratory of Human Bacterial Pathogenesis, Rocky..."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jakki C.","family":"Cooney","sequence":"additional","affiliation":[{"name":"School of Biological Sciences and Department of Chemistry, University of Auckland, Private Bag 92\u2013019, Auckland, New Zealand; Institute of Molecular Biosciences, Massey University, Palmerston North, New Zealand; European Molecular Biology Laboratory Grenoble Outstation, 6 Av. J. Horowitz, Grenoble 38042, France; Institute of Human Bacterial Pathogenesis, Department of Pathology, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030; and Laboratory of Human Bacterial Pathogenesis, Rocky..."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heather M.","family":"Baker","sequence":"additional","affiliation":[{"name":"School of Biological Sciences and Department of Chemistry, University of Auckland, Private Bag 92\u2013019, Auckland, New Zealand; Institute of Molecular Biosciences, Massey University, Palmerston North, New Zealand; European Molecular Biology Laboratory Grenoble Outstation, 6 Av. J. Horowitz, Grenoble 38042, France; Institute of Human Bacterial Pathogenesis, Department of Pathology, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030; and Laboratory of Human Bacterial Pathogenesis, Rocky..."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sean","family":"McSweeney","sequence":"additional","affiliation":[{"name":"School of Biological Sciences and Department of Chemistry, University of Auckland, Private Bag 92\u2013019, Auckland, New Zealand; Institute of Molecular Biosciences, Massey University, Palmerston North, New Zealand; European Molecular Biology Laboratory Grenoble Outstation, 6 Av. J. Horowitz, Grenoble 38042, France; Institute of Human Bacterial Pathogenesis, Department of Pathology, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030; and Laboratory of Human Bacterial Pathogenesis, Rocky..."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mengyao","family":"Liu","sequence":"additional","affiliation":[{"name":"School of Biological Sciences and Department of Chemistry, University of Auckland, Private Bag 92\u2013019, Auckland, New Zealand; Institute of Molecular Biosciences, Massey University, Palmerston North, New Zealand; European Molecular Biology Laboratory Grenoble Outstation, 6 Av. J. 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Horowitz, Grenoble 38042, France; Institute of Human Bacterial Pathogenesis, Department of Pathology, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030; and Laboratory of Human Bacterial Pathogenesis, Rocky..."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James M.","family":"Musser","sequence":"additional","affiliation":[{"name":"School of Biological Sciences and Department of Chemistry, University of Auckland, Private Bag 92\u2013019, Auckland, New Zealand; Institute of Molecular Biosciences, Massey University, Palmerston North, New Zealand; European Molecular Biology Laboratory Grenoble Outstation, 6 Av. J. 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