Microbial Extracellula r Polymeric Substance s

Transcription

1 Jost Wingender Thomas R. Neu Hans-Curt Flemming Microbial Extracellula r Polymeric Substance s Characterization, Structure and Function

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3 What are Bacterial Extracellular Polymeric Substances? Jost Wingender, Thomas R. Neu, Hans-Curt Flemming 1 1 Introduction 1 2 Definitions of EPS 2 3 Composition, Secretion and Spatial Arrangement of EPS 4 4 Industrial and Clinical Importance of EPS 9 5 Bacterial Alginate - an Example of Bacterial EPS Functions of EPS Ecological Aspects of EPS 1 3 References 1 5 In Situ Characterization of Extracellular Polymeric Substances (EPS ) in Biofilm Systems Thomas R. Neu, John R. Lawrence Introduction Traditional Approaches for Studying Microbial Polysaccharides Change of Paradigms In situ Methods 23 2 Destructive Analysis of EPS Chemical Techniques Extraction Analysis Electron Microscopy Scanning Probe Techniques Non-Destructive Analysis of EPS Infrared Spectroscopy (FT-IR) Magnetic Resonance Spectroscopy (NMR) Confocal Laser Scanning Microscopy (CLSM) CLSM Approach Probes for Analysis of EPS Polysaccharides Proteins Nucleic Acids Physicochemical Characterization of EPS Matrix 32

4 4.2.1 Charge Distribution Hydrophobicity/Hydrophilcity Permeability EPS Bound and Associated Molecules Image Analyses and Three-Dimensional Data Presentation Analyses Techniques Three-Dimensional Imaging Three-Dimensional Display Case Studies : In Situ Characterization of EPS Limitations of Non-Destructive Analyses FT-IR/ATR NMR CLSM 4 1 References 42 Extraction of EP S Per H. Nielsen, Andreas Jahn 49 1 Introduction Definition of EPS Composition of EPS Extraction Procedure Sampling and Pretreatment Sampling and Storage Homogenization Washing 54 3 Extraction of EPS Selection of an Appropriate Extraction Method Physical Methods Chemical Methods Combination of Physical and Chemical Methods Contamination by Intracellular Macromolecules Disruption of Macromolecules 64 6 Extraction Efficiency Purification and Analysis Conclusion 6 8 References 6 9 Biofilm Exopolysaccharide s Ian W. Sutherland Introduction Polysaccharide Composition and Structure Promotion of Order and Gelation Ionic Interactions 83 5 Synergistic Effects 83

5 6 Other Functions of Biofilm Exopolysaccharides Dissolution of Physical Structures 87 8 Conclusions 8 8 References 8 9 Regulation of Matrix Polymer in Biofilm Formation and Dispersion David G. Davies 93 1 Alginate Biosynthesis in Pseudomonas aeruginosa Regulation of Alginate Biosynthetic Genes During Biofilm Development Control of Biofilm Dispersion Activation of Alginate Biosynthesis and Chemical Communication in Biofilms P. aeruginosa Biofilm Life Cycle 11 0 References 11 2 Exopolymers of Sulphate-Reducing Bacteri a Iwona B. Beech, Rudi C. Tapper 11 9 Analysis and Function of the EPS from the Strong Acidophil e Thiobacillus ferrooxidan s Wolfgang Sand, Tilman Gehrke Introduction Leaching Bacteria Metal Sulfide Dissolution EPS Analysis Function of EPS Conclusions 13 8 References 140 Physical and Chemical Properties of Extracellular Polysaccharide s Associated with Biofilms and Related Systems Bjorn E. Christensen Introduction Biopolymer Gels Physical Characterization of Biofilms Chemical and Biochemical Characterization of Biofilms Bacterial Alginates 149 References 153

6 Chemical Communication Within Microbial Biofilms: Chemotaxis and Quorum Sensing in Bacterial Cell s Alan W. Decho Introduction Bacterial Chemotaxis and Initial Aggregations of Cells Microspatial Patterns and Movement of Bacteria Within Biofilms Bacterial Quorum Sensing : Induction and Regulation of Activities Model Systems Homoserine Lactone (Autoinducer) Signals The Vibrio fischeri System Other Chemical Signals Conjugation and Pheromone-Inducible Gene Expression in Enterococcus faecalis Cyanobacteria Systems Functional Roles of Quorum Sensing Specificity of Autoinduction Process Enhancement of Chemotaxis and Quorum Sensing Processe s by the Biofilm Matrix The Biofilm as a Microbial Organism 16 4 References 16 5 Function of EPS Gideon M. Wolfaardt, John R. Lawrence, Darren R. Korber Introduction Role of EPS in Cellular Associations Suspended and Attached Cells Coaggregation, Consortial Behavior, and Floc Formation Biofilm Formation Genetic Transfer Role of EPS in Nutrition Production of EPS as a Nutrient Reserve Accumulation of Nutrients by EPS Role of EPS in the Interaction of Microorganisms with their Biological, Physical, and Chemical Environment EPS and the Micro-Environment EPS as a Physical Barrier to Solute Translocation Role in Enzyme Reactions Role in Pathogenicity and Protection Against Host Defens e Mechanisms Protective Role Against Predation/Digestion Microbial-Plant Associations Effect of Environmental Conditions EPS and the Macro-Environment Stabilization of the Environment 190

7 5.2 Microbial-Mineral Interactions Controls on Flow Role of EPS in the Bio-Accumulation of Contaminants Conclusions 19 4 References 19 5 Polysaccharases in Biofilms - Sources - Action - Consequences! Ian W. Sutherland Polysaccharases in Biofilms Nature of Polysaccharases Sources of Polysaccharases The Endogenous Production of Polysaccharases The Exogenous Production of Polysaccharases Bacteriophage Effects of Enzymes on Biofilms Application of Enzymes to Biofilms 21 2 References 21 3 Extracellular Enzymes Within Microbial Biofilm s and the Role of the Extracellular Polymer Matri x Monica Hoffman, Alan W. Decho Introduction: Importance of Extracellular Enzymes to Bacteria l Cells and Organic Matter Processing The Microbial Biofilm and Extracellular Polymers Extracellular Polymers as a "Sorptive Sponge" and "Colloidal Trap " for High-Molecular-Weight Organic Matter Localization of Enzymes by Extracellular Polymers : the "Lectin-Localization Model" "Extracellular Polymer Microdomains" and the Stabilit y of Extracellular Enzymes Environmental Influences on Enzyme Activity Biofilm Induction and Regulation of Extracellular Enzymes Induction of Extracellular Enzymes at the Physiological Level Regulation of Enzyme Activity by Extracellular Factors 22 6 References 22 7 Interaction Between Extracellular Polysaccharides and Enzyme s Jost Wingender, Karl-Erich Jaeger, Hans-Curt Flemming Extracellular Enzymes in Biofilms Mechanisms of Enzyme and Polysaccharide Secretion Secretion of Enzymes The ABC Pathway 236

8 2.1.2 The General Secretory Pathway The Contact Site-Dependent Pathway Secretion of Polysaccharides Interaction of Enzymes with Extracellular Polysaccharides Consequences of Enzyme-Polysaccharide Interactions in Biofilms 246 References 247