The Bacterial Cell Wall

289 Pages·2002·6.602 MB·English

by  Dr. rer. nat. habil. Guntram Seltmann, Prof. Dr. rer. nat. Otto Holst (auth.)

The Bacterial Cell Wall
Springer-Verlag Berlin Heidelberg GmbH
Guntram Seltmann · Otto Holst
The Bacterial Cell Wall
With 114 Figures and 23 Tables
, Springer
Dr. rer. nat. habil. Guntram Seltmann
Karl-Marx-Str·9
38855 Wernigerode
Germany
Prof. Dr. rer. nat. Otto Holst
Forschungszentrum Borstel
Zentrum fur Medizin und Biowissenschaften
Parkallee 1-40
23845 Borstel
Germany
ISBN 978-3-642-07648-0 ISBN 978-3-662-04878-8 (eBook)
DOI 10.1007/978-3-662-04878-8
Library of Congress Cataloging-in-Publication Data
Seltmann, Guntram. The bacterial cell wall I Guntram Seltmann, Otto Holst.
p. cm. Includes bibliographical references and index.
1. Bacterial cell walls. I. Holst, Otto. II. Title.
QRn3 .S45 2001 571.6'8293--dc21
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Contents
Preface ............................................... .
1 Introduction............. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 The Outer Membrane of the Gram-Negative Bacteria and their
Components. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.1 Lipids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.1.1 Classification, Significance . . . . . . . . . . . . . . . . . . . . . . 9
2.1.2 Isolation and Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.1.3 Chemical Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.1.3.1 Fatty Acids. . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.1.3.2 Waxes and Glycerides. . . . . .. ............ 12
2.1.3.3 Phospholipids and Glycolipids. . . . . . . . . . . . . . 14
2.1.3.3.1 Glycerolphospholipids........... 15
2.1.3.3.2 Glycolipids................... 16
2.1.3.3.3 Sphingolipids................. 17
2.1.3.3.4 Hopanoids....... ............ 17
2.1.4 Spatial Structure and Physical Properties . . . . . .. ..... 18
2.1.5 Biosynthesis of the Lipids. . . . . .. ................ 22
2.1.5.1 Biosynthesis of Fatty Acids. . . . . .. ......... 22
2.1.5.2 Biosynthesis of Waxes and Glycerides. . . . . ... 23
2.1.5.3 Biosynthesis of Phospholipids and Glycolipids . . 24
2.1.5.3.1 Biosynthesis of Glycerolphospholipids 24
2.1.5.3.2 Biosynthesis of Glycolipids. . . . . ... 24
2.1.5.3.3 Biosynthesis of Sphingolipids. . . . . . . 24
2.1.5.3.4 Biosynthesis of ACL. . . . . .. ...... 25
2.1.5.3.5 Biosynthesis of Hopanoids. . . . . .. .. 29
2.1.5.4 Incorporation into the Outer Membrane. . . . . ... 29
2.2 Lipopolysaccharides..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
2.2.1 General Remarks. . . .. ......................... 31
2.2.2 General Structure. . . . . . . . . . . . . . . . . . . . . . . .. ..... 33
2.2.3 Isolation and Analysis. . . . . . .. ... . . . . . . . . . . . . . . .. 34
2.2.3.1 Isolation........................ ...... 34
2.2.3.2 Analysis........................ ...... 37
2.2.4 Composition and Structure. . . . . . . . . . . . . . . . . . . . . . .. 40
2.2.4.1 The O-Specific Polysaccharide. . . . . . .. ...... 40
VI
2.2.4.2 The Core Region . . . . . . . . . . . . . . . . . . . . . . . 42
2.2.4.3 The Lipid A. . . . . . . . . . . . . . . . . . . . . . . . . . . 45
2.2.4.4 Physical Properties and Spatial Structure. . . . . . . 48
2.2.5 Biosynthesis................... .......... . . . . 50
2.2.5.1 Biosynthesis of the Precursors. . . . . . . . . . . . . . 50
2.2.5.2 Biosynthesis of Lipid A and the Core Region. . . 52
2.2.5.3 Biosynthesis of O-Specific Polysaccharides. . . . 57
2.2.5.4 Genetic Determination of the LPS Biosynthesis. . 61
2.2.6 The Enterobacterial Common Antigen CECA) . . . . . . . . . 65
2.3 Proteins of the Outer Membrane of Gram-Negative Bacteria. . . . 67
2.3.1 General Remarks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
2.3.1.1 Composition and Structure. . . . . . . . . . . . . . . . . 67
2.3.1.2 Isolation and Analysis. . . . . . . . . . . . . . . . . . . . 73
2.3.2 Proteins of the Outer Membrane. . . . . . . . . . . . . . . . . . . 74
2.3.2.1 Classification.......................... 74
2.3.2.2 Detection and Isolation. . . . . . . . . . . . . . . . . . . 75
2.3.2.3 The Channel-Forming Proteins . . . . . . . . . . . . . 75
2.3.2.3.1 Porin Proteins. . . . . . . . . . . . . . ... 76
2.3.2.3.2 Proteins Forming Specific Channels. . 78
2.3.2.3.3 High-Affinity Receptor Proteins. . . . . 79
2.3.2.4 The Structure Proteins. . . . . . . . . . . . . . . . . . . . 81
2.3.2.4.1 Braun's Lipoprotein. . . . . . . . . . . . . 81
2.3.2.4.2 OmpA-Protein................. 83
2.3.2.5 Further Proteins. . . . . . . . .. .............. 84
2.3.2.6 Biosynthesis of the Outer Membrane Proteins. . . 84
2.3.3 Organisation and Function of the Outer Membrane
Proteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
2.4 The Total Membrane. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
2.4.1 General Structure of Biological Membranes. . . . . . . . . . . 91
2.4.1.1 Composition, Structure, Function. . . . . . . . . . . . 91
2.4.1.2 The Asymmetry of Membranes . . . . . . . . . . . . . 92
2.4.2 Composition of the Outer Membrane. . . . . . . . . . . . . . . . 94
2.4.3 Structure of the Outer Membrane . . . . . . . . . . . . . . . . . . 95
2.4.4 The Fluidity of the Outer Membrane. . . . . . . . . . . . . . . . 98
2.4.5 Assembly of the Outer Membrane. . . . . . . . . . . . . . . . .. 100
3 Periplasmic Space and Rigid Layer. . . . . . . . . . . . . . . . . . . . .. 103
3.1 The Periplasmic Space. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 103
3.2 The Rigid Layer of Gram-Positive and Gram-Negative Bacteria.. 105
3.2.1 Significance, Isolation, Composition, Structure. . . . . . . .. 105
3.2.1.1 Isolation of the Peptidoglycan. . . . . . . . . . . . .. 105
3.2.1.2 Chemical Structure of Peptidoglycan.. . . . . . .. 106
3.2.1.2.1 Chemical Structure of the
Polysaccharide Moiety. . . . . . . . . . .. 107
3.2.1.2.2 Chemical Structure of the Peptide
Moiety. . . . . . . . . . . . . . . . . . . . . .. 109
VII
3.2.1.3 The Spatial Structure of the Peptidoglycan. . . .. 111
3.2.1.4 Peptidoglycan as a Transport Barrier. . . . . . . . .. 117
3.2.2 Biosynthesis of Peptidoglycan. . . . . . . . . . . . . . . . . . . .. 118
3.2.2.1 Biosynthesis of UDP-N-Acetylmuramyl
Pentapeptide. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 119
3.2.2.2 Biosynthesis of the Complete Subunit and of the
Bridge Peptide. . . . . . . . . . . . .. ...... . . . .. 120
3.2.2.3 Transfer to the Growing Terminus of the
Polysaccharide Chain. . . . . . . . . . . . .. ...... 123
3.2.2.4 Cross-Linking.......................... 123
3.2.2.5 Regulation of Peptidoglycan Biosynthesis. . . . .. 125
3.2.2.6 Possibilities to Influence the Biosynthesis of
Peptidoglycan. . . . . . . . . . . . . . . . . . . . . . . . .. 128
4 Further Cell Wall Components of Gram-Positive Bacteria. . . .. 133
4.1 Polysaccharides (Except Capsular Polysaccharides) . . . . . . . . .. 133
4.1.1 The Teichoic Acid Family. . . . . . . . . . . . . . . . . . . . . . .. 133
4.1.1.1 Teichoic Acids. . . . . . . . . . . . . . . . . . . . . . . .. 134
4.1.1.1.1 Poly-(Polyolphosphate)-Teichoic
Acids. . . . . . . . . . . . . . . . . . . . . . .. 134
4.1.1.1.2 Poly-(Glycosylpolyolphosphate)-
Teichoic Acids. . . . . . . . . . . . . . . .. 135
4.1.1.1.3 Heterogeneity of Teichoic Acids. . .. 136
4.1.1.1.4 Linkage Units to Peptidoglycan. . . .. 136
4.1.1.2 Teichuronic Acids . . . . . . . . . . . . . . . . . . . . .. 136
4.1.1.3 Lipoteichoic Acids. . . . . . . . . . . . . . . . . . . . .. 137
4.1.1.4 Functions of the Teichoic Acids. . . . . . . . . . . .. 139
4.1.2 Acidic Polysaccharides. . . . . . . . . . . . .. ........... 140
4.1.3 Biosynthesis................................. 141
4.1.3.1 Biosynthesis of Teichoic Acids . . . . . . . . . . . .. 141
4.1.3.2 Biosynthesis of Teichuronic Acids .......... 142
4.1.3.3 Biosynthesis of Lipoteichoic Acids. . . . . . . . .. 142
4.1.3.4 Control of the Biosyntheses. . . . . . . . . . . . . . .. 142
4.2 Proteins....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
4.2.1 Isolation and General Structure. . . . . . . . . . . . . . . . . . .. 144
4.2.2 Streptococcal Cell Wall Proteins. . . . . . . . . . . . . . . . . .. 146
4.2.3 Staphylococcal Protein A and Similar Proteins. . . . . . . .. 147
4.2.4 Clumping Factor of Staphylococci. . . . . . . . . . . . . . . . .. 149
4.2.5 Further Proteins of the Gram-Positive Cell Wall. . . . . . .. 149
4.2.6 Biosynthesis................................. 150
4.3 Cell Wall Components of Mycolata. . . . . . . . . . . . .. ....... 151
4.3.1 The Mycoloyl Arabinogalactan Complex. . . . . . . . . . . .. 152
4.3.1.1 The Mycolic Acid Moiety. . . . . . . . . . . . . . . .. 152
VIII
4.3.1.1.1 Properties, Structure. . . . . . . . . . . . . 152
4.3.1.1.2 Biosynthesis................... 154
4.3.1.2 The Arabinogalactan Moiety. . . . . . . . . . . . ... 155
4.3.1.3 The Complete Molecule. . . . . . . . . . . . .. .... 155
4.3.2 Lipoarabinomannan............................ 157
4.3.3 Extractable Lipids. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 158
4.3.3.1 Lipooligosaccharides (LOS) . . . . . . . . . . . . . .. 158
4.3.3.2 Phenolic Glycolipids (PGL) . . . . . . . . . . . . ... 159
4.3.3.3 Glycopeptidolipids (GPL) . . . . . . . . . . . . .. .. 160
4.3.4 Waxes and Sulfolipids. . . . . . . . . . . . . . . . . . . . . . . . .. 161
4.3.5 Proteins.................................... 161
5 Cell Wall Components of Archaea. . . . . . . . . . . . . . . . . . . . . . .. 162
5.1 Pseudomurein. . . . . . . . . . . . . . . . . . . . . . . . . . . .. ....... 163
5.2 Methanochondroitin. . . . . . . . . . . . . . . . . . . . . . . . . . . .. ... 164
5.3 Heteropolysaccharides. . . . . . . . . . . . . . . . . . . . . . . . . . . ... 166
5.4 Glutaminylglycan.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
5.5 Lipids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. ... 167
5.6 S-Layers and Sheaths. . . . . . . . . . . . . . . . . . . . . . . . . . . .. .. 169
6 Components Outside the Cell Wall. . . . . . . . . . . . . . . . . . . . . . .. 171
6.1 Capsules and Slime Layers. . . . . . . . . . . . . . . . . . . . . . . . . . .. 171
6.1.1 General Characterisation. . . . . . . . . . . .. .......... 171
6.1.2 Detection of Capsules. . . . . . . . . . . . . . . . . . . . . . . . . .. 172
6.1.3 Isolation and Chemical Analysis of the Capsule Material.. 173
6.1.4 The Capsules of Gram-Negative Bacteria. . . . . . . . . . . .. 174
6.1.4.1 Escherichia coli. . . . . . . . . . . . . . . . . . . . . . . .. 174
6.1.4.2 The Capsules of other Gram-Negative Bacteria. .. 178
6.1.4.2.1 Gram-Negative Rods. . . . . . . . . . . .. 178
6.1.4.2.2 Gram-Negative Cocci. . . . . . . . . . .. 179
6.1.5 Capsules of Gram-Positive Bacteria. . . . . . . . . . . . . . . .. 179
6.1.5.1 Streptococci and Staphylococci. . . . . . . . . . . .. 179
6.1.5.2 Mycobacteria.......................... 180
6.1.6 Biosynthesis of CPS. . . . . . . . . . . . . . . . . . . . . . . . . . .. 180
6.1.7 Capsules in Immunogenicity and Virulence. . . . . . . . . .. 182
6.2 S-Layers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... 183
6.2.1 General Remarks. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 183
6.2.2 Composition and Structure. . . . . . . . . . . . . . . . . . . . . . . 183
6.2.2.1 Archaea.............................. 185
6.2.2.2 Bacteria.............................. 187
6.2.3 Biosynthesis................ ................. 187
6.2.4 Fixation of S-Layers in the Cell Wall. . . . . . . . . . . . . . .. 190
6.2.5 Functions of S-Layers. . . . . . . . . . . . . . . . . . . . . . . . . .. 190
6.3 Sheaths. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. .. 193
IX
6.4 Filamentous Proteins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 194
6.4.1 Flagella..................................... 194
6.4.2 Fimbriae and Fibrils. . . . . . . . . . . . . . . . . . . . . . . . . . .. 198
6.4.3 Sex Pili. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 202
7 Cell Wall Models. . . . . . . . . . . . .. ........................ 204
7.1 Gram-Negative Bacteria. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 206
7.2 Gram-Positive Bacteria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
7.2.1 "Typical" Gram-Positive Bacteria. . . . . . . . . . . .. .... 211
7.2.2 "Atypical" Gram-Positive Bacteria. . . . . . . . . . . . . . . . .. 215
8 Cell Wall Functions. . . . . . . . . . . . .. ...................... 219
8.1 The Cell Wall as a Transport Organ. . . . . . . . . . . . . . . . . . . .. 219
8.1.1 General Remarks. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 219
8.1.2 Transport Across the Bacterial Cell Wall. . . . . . . . . . . .. 221
8.1.2.1 Transport from the Exterior to the Interior. . . . .. 221
8.1.2.2 Transport from the Interior to the Exterior. . . . .. 227
8.1.3 The Transport (Transduction) of Signals. . . . . . . . . . . .. 231
8.2 Protective Function of the Bacterial Cell Wall. . . . . . . . . . . . .. 233
8.3 The Importance of the Cell Wall for Pathogenicity and Virulence 235
8.3.1 Adherence of Bacteria at Surfaces. . . . . . . . . . . . . . . . .. 238
8.3.2 Survival at Surfaces and Penetration Into or Across the
Tissue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 239
8.3.3 Growth in the Host. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 241
8.3.4 Undermining, Repulse or Elimination of the Defence
Mechanisms of the Host. . . . . . . . . . . . . . . . . . . . . . . .. 242
8.3.5 Damaging of the Host. . . . . . . . . . . . . . . . . . . . . . . . .. 244
8.3.6 The Roles of Particular Cell Wall Components. . . . . . . .. 244
8.3.6.1 Polysaccharides and Related Compounds. . . . .. 245
8.3.6.2 Lipids (Including Lipid A) . . . . . . . . . . . . . . .. 247
8.3.6.3 Proteins.............................. 248
8.3.6.4 Peptidoglycan.......................... 250
8.4 Significance of the Cell Wall for the Maintenance of the Bacterial
Shape. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 251
8.4.1 Rod-Shaped Bacteria. . . . . . . . . . . . . . . . . . . . . . . . . .. 251
8.4.2 Spherical (Coccoid) Bacteria. . . . . . . . . . . . . . . . . . . . .. 253
8.5 Interactions between the Bacterial Cell Wall and Bacteriophages. 257
8.6 Interactions between the Bacterial Cell Wall and Antibiotics. . .. 261
Uncommon Abbreviations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 266
Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. .. 268
Preface
The bacterial cell wall represents a very complex structure disconnecting the
interior of single-cell organisms from the environment, thus protecting, but also
enabling, them to interact with the surrounding milieu and to exchange both
substances and information. Knowledge of the biochemistry of the cell wall
(components) and the genetic background helps to understand their significance
with regard to microbiology and immunology of bacteria.
This book represents the second edition of a publication which was presented
nearly 20 years ago in the German language (Die bakterielle Zellwand). Since that
time our knowledge in this field has been significantly enlarged. Therefore, the
manuscript had to be completely revised and updated. To maintain both the size
and the introductory character of the book at least to a great extent, the authors had
to restrict the presented material to that which appears basic and most important.
This requirement must inevitably bring about many subjective factors.
As pointed out in the first edition, the term cell wall was not taken too strictly.
Since the constituents located outside the cytoplasmic membrane are frequently
difficult to divide in structure, localisation, and/or function into true cell wall
components and supplementary substances, they are all at least briefly mentioned.
For this reason capsules, S-layers and appendices like flagellae, fimbriae and pili
are discussed. In addition, sometimes deeper regions of the bacteria like
cytoplasmic membrane or cytoplasm are crucial for the course of processes
necessary for the cell wall (components), e.g. for their biosynthesis or transport. In
such cases, these also are discussed.
The progress of natural sciences depends mainly on the development and
introduction of modern methods. Therefore in the text important methods are
mentioned and briefly discussed. In some cases it appeared useful, by historical
and didactic reasons, to discuss older approved methods.
Science does not only live in the present. Current developments are mostly
based on results obtained in the past. Thus, it was both tempting and attractive to
sketch or outline historical developments in particular cases.
In most cases the binary taxonomic designation of the species is used
unabbreviated. In the case of well-known bacteria (e.g. Escherichia coli) the first
term is abbreviated. In agreement with the commonly used practice for Salmonella
serovars of subspecies I (Salmonella enterica subsp. enterica), the old names were
maintained, however, not italicised, and the second term capitalised (e.g.
Salmonella Typhi).
Some decades ago (1977) archaebacteria (archaea) were recognised as a third
form of life besides (eu-)bacteria and eukaryotes. They are prokaryotes whose cell
walls in many cases resemble those of the "classical" bacteria, but which may