MICROBIOLOGY AND MOLECULAR BIOLOGY REVIEWS, Dec. 2004, p. 692–744 Vol. 68, No. 4 1092-2172/04/$08.00ϩ0 DOI: 10.1128/MMBR.68.4.692–744.2004 Copyright © 2004, American Society for Microbiology. All Rights Reserved. Type V Protein Secretion Pathway: the Autotransporter Story Ian R. Henderson,1* Fernando Navarro-Garcia,2 Mickae¨l Desvaux,1 Rachel C. Fernandez,3 and Dlawer Ala’Aldeen4 Division of Immunity and Infection, University of Birmingham, Birmingham,1 and Division of Microbiology & Infectious Diseases, University Hospital, Nottingham,4 United Kingom; Department of Cell Biology, CINVESTAV, Mexico City, Mexico2; and Department of Microbiology & Immunology, University of British Columbia, Vancouver, British Columbia, Canada3 INTRODUCTION .......................................................................................................................................................693 Downloaded from SECRETION BY NUMBERS: PROTEIN SECRETION MECHANISMS IN GRAM-NEGATIVE BACTERIA...........................................................................................................................................................693 Type I Protein Secretion........................................................................................................................................694 Two-Step Type II Protein Secretion .....................................................................................................................695 The Injectisome: Type III Protein Secretion.......................................................................................................696 Type IV Protein and Nucleoprotein Secretion....................................................................................................697 TYPE V PROTEIN SECRETION.............................................................................................................................697 Autotransporter Secretion Pathway (Type Va)...................................................................................................698 Inner membrane transport................................................................................................................................699 Periplasmic transit .............................................................................................................................................701 http://mmbr.asm.org/ ␤-Domain .............................................................................................................................................................702 Translocation of the passenger domain...........................................................................................................702 Processing of autotransporters at the surface................................................................................................704 Energy of transport ............................................................................................................................................705 Evolution and distribution of autotransporter proteins ...............................................................................705 Two-Partner Secretion Pathway (Type Vb).........................................................................................................705 Type Vc.....................................................................................................................................................................708 CLUSTER 1: SUBTILASE FAMILY OF AUTOTRANSPORTERS ....................................................................709 Ssp.............................................................................................................................................................................709 PspA and PspB........................................................................................................................................................709 Ssa1...........................................................................................................................................................................709 on December 9, 2018 by guest SphB1 .......................................................................................................................................................................709 AspA/NalP................................................................................................................................................................710 CLUSTERS 2 AND 11: HELICOBACTER PYLORI AUTOTRANSPORTERS...................................................710 VacA, the Vacuolating Cytotoxin ..........................................................................................................................710 Molecular and structural features ...................................................................................................................710 Vacuolating activity ............................................................................................................................................711 Other functions ...................................................................................................................................................711 Epidemiology of H. pylori infection ..................................................................................................................711 BabA, SabA, and AlpA ...........................................................................................................................................712 CLUSTERS 3 AND 4: AIDA AUTOTRANSPORTER FAMILY...........................................................................713 AIDA-I ..........................................................................................................................................................................713 Antigen 43 ................................................................................................................................................................714 IcsA ...........................................................................................................................................................................716 TibA, ShdA, and MisL ...........................................................................................................................................716 AutA and AutB ........................................................................................................................................................718 CLUSTERS 5 AND 8: SERINE PROTEASE AUTOTRANSPORTERS .............................................................718 IgA1 Proteases.........................................................................................................................................................718 H. influenzae Adhesion and Penetration Protein (Hap) ....................................................................................720 N. meningitidis App and MspA Autotransporter Proteins.................................................................................720 Serine Protease Autotransporters of the Enterobacteriaceae.............................................................................721 Tsh ........................................................................................................................................................................721 SepA......................................................................................................................................................................722 EspC .....................................................................................................................................................................722 EspP......................................................................................................................................................................722 Pet .........................................................................................................................................................................723 * Corresponding author. Mailing address: Division of Immunity and Infection, The Medical School, University of Birmingham, Birming- ham B15 2TT, United Kingdom. Phone: 44 121 414 4368. Fax: 44 121 414 3599. E-mail: [email protected]. 692 VOL. 68, 2004 TYPE V SECRETION 693 Pic .........................................................................................................................................................................723 SigA.......................................................................................................................................................................725 Sat .........................................................................................................................................................................725 Vat.........................................................................................................................................................................725 Others...................................................................................................................................................................725 CLUSTER 6: BORDETELLA AUTOTRANSPORTERS ........................................................................................726 Identification of Autotransporters in the Bordetella Genomes .........................................................................726 Pertactin...................................................................................................................................................................727