Bacterial Modulins: a Novel Class of Virulence Factors Which Cause Host Tissue Pathology by Inducing Cytokine Synthesis
B. HENDERSON, S. POOLE, AND M. WILSON



INTRODUCTION......................................................................................................................................................317
CYTOKINE BIOLOGY.............................................................................................................................................317
Cytokines—a Working Definition........................................................................................................................318
Biological Actions of Cytokines............................................................................................................................318
Cytokine Nomenclature.........................................................................................................................................319
Pathological Actions of Cytokines ........................................................................................................................319
Anti-Inflammatory Cytokines ...............................................................................................................................321
Cytokine Gene Knockouts and Susceptibility to Infection ...............................................................................322
LPS AND CYTOKINE INDUCTION......................................................................................................................322
LPS and Endotoxin: Definition and Composition..............................................................................................322
Standard for LPS...............................................................................................................................................323
Structure of LPS....................................................................................................................................................323
Lipid A.................................................................................................................................................................323
Core region .........................................................................................................................................................323
O-antigenic side chain.......................................................................................................................................323
LPS Receptor..........................................................................................................................................................323
LPS-Induced Transmembrane Signalling ...........................................................................................................325
CYTOKINE-STIMULATING CELL COMPONENTS OTHER THAN LPS......................................................326
Proteins ...................................................................................................................................................................326
Outer membrane proteins ................................................................................................................................326
(i) Porins.........................................................................................................................................................327
(ii) Lipid A-associated proteins ....................................................................................................................327
(iii) Other outer membrane proteins ...........................................................................................................328
Fimbrial proteins ...............................................................................................................................................328
Other cell wall-associated proteins ..................................................................................................................328
(i) Unidentified surface-associated proteins ...............................................................................................328
(ii) Protein A ..................................................................................................................................................329
Heat shock proteins...........................................................................................................................................329
Lipoproteins............................................................................................................................................................329
Glycoproteins..........................................................................................................................................................329
Lipids.......................................................................................................................................................................329
Polysaccharides ......................................................................................................................................................329
Cell surface polysaccharides .............................................................................................................................329
Peptidoglycan......................................................................................................................................................330
Peptidoglycan fragments...................................................................................................................................330
Teichoic Acids.........................................................................................................................................................331
Mycobacterial Cell Wall Components..................................................................................................................331
Lipoarabinomannan ..........................................................................................................................................331
Other mycobacterial components .....................................................................................................................332
EXTRACELLULAR PRODUCTS............................................................................................................................332
Proteases .................................................................................................................................................................332
Exotoxins.................................................................................................................................................................332
Superantigens .........................................................................................................................................................333
Other Extracellular Proteins................................................................................................................................333
BACTERIAL ADHESION.........................................................................................................................................333
BACTERIAL INVASION ..........................................................................................................................................333
SUMMARY AND CONCLUSIONS.........................................................................................................................334
ACKNOWLEDGMENTS ..........................................................................................................................................335
REFERENCES ...........................................................................................................................................................335

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