Fungal cell wall : structure, synthesis, and assembly

194 Pages·2012·42.226 MB·English

by  José Ruiz-Herrera

Fungal Cell Wall
Structure, Synthesis, and Assembly
Second Edition
MYCOLOGY SERIES
Editor
J. W. Bennett
Professor
Department of Plant Biology and Pathology
Rutgers University
New Brunswick, New Jersey
Founding Editor
Paul A. Lemke
1. Viruses and Plasmids in Fungi, edited by Paul A. Lemke
2. The Fungal Community: Its Organization and Role in the Ecosystem, edited by Donald T. Wicklow
and George C. Carroll
3. Fungi Pathogenic for Humans and Animals (in three parts), edited by Dexter H. Howard
4. Fungal Differentiation: A Contemporary Synthesis, edited by John E. Smith
5. Secondary Metabolism and Differentiation in Fungi, edited by Joan W. Bennett and Alex Ciegler
6. Fungal Protoplasts, edited by John F. Peberdy and Lajos Ferenczy
7. Viruses of Fungi and Simple Eukaryotes, edited by Yigal Koltin and Michael J. Leibowitz
8. Molecular Industrial Mycology: Systems and Applications for Filamentous Fungi,
edited by Sally A. Leong and Randy M. Berka
9. The Fungal Community: Its Organization and Role in the Ecosystem, Second Edition,
edited by George C. Carroll and Donald T. Wicklow
10. Stress Tolerance of Fungi, edited by D. H. Jennings
11. Metal Ions in Fungi, edited by Güfcnther Winkelmann and Dennis R. Winge
12. Anaerobic Fungi: Biology, Ecology, and Function, edited by Douglas O. Mountfort and
Colin G. Orpin
13. Fungal Genetics: Principles and Practice, edited by Cees J. Bos
14. Fungal Pathogenesis: Principles and Clinical Applications, edited by Richard A. Calderone and
Ronald L. Cihlar
15. Molecular Biology of Fungal Development, edited by Heinz D. Osiewacz
16. Pathogenic Fungi in Humans and Animals, Second Edition, edited by Dexter H. Howard
17. Fungi in Ecosystem Processes, John Dighton
18. Genomics of Plants and Fungi, edited by Rolf A. Prade and Hans J. Bohnert
19. Clavicipitalean Fungi: Evolutionary Biology, Chemistry, Biocontrol, and Cultural Impacts,
edited by James F. White Jr., Charles W. Bacon, Nigel L. Hywel-Jones, and Joseph W. Spatafora
20. Handbook of Fungal Biotechnology, Second Edition, edited by Dilip K. Arora
21. Fungal Biotechnology in Agricultural, Food, and Environmental Applications,
edited by Dilip K. Arora
22. Handbook of Industrial Mycology, edited by Zhiqiang An
23. The Fungal Community: Its Organization and Role in the Ecosystem, Third Edition,
edited by John Dighton, James F. White, and Peter Oudemans
24. Fungi: Experimental Methods in Biology, Ramesh Maheshwari
25. Food Mycology: A Multifaceted Approach to Fungi and Food, edited by Jan Dijksterhuis and
Robert A. Samson
26. The Aspergilli: Genomics, Medical Aspects, Biotechnology, and Research Methods,
edited by Gustavo H. Goldman and Stephen A. Osmani
27. Defensive Mutualism in Microbial Symbiosis, edited by James F. White, Jr. and Mónica S. Torres
28. Fungi: Experimental Methods In Biology, Second Edition, Ramesh Maheshwari
29. Fungal Cell Wall: Structure, Synthesis, and Assembly, Second Edition, José Ruiz-Herrera
Fungal Cell Wall
Structure, Synthesis, and Assembly
Second Edition
José Ruiz-Herrera
Centro de Investigación y de Estudios Avanzados
Irapuato, Mexico
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Dedication
This book, as was the first edition, is dedicated first to my wife, Carmen, with
love, for all the support, help, and understanding that she has given to me during
the last 51 years. I should also include in this dedication all those who have
brought joy to my life, my daughters and sons, Dr. Ethelia Ruiz-Medrano, José
Javier Ruiz-Medrano (in memoriam), Dr. Roberto Ruiz-Medrano, Dr. Carlos
Rubén Ruiz-Medrano, Psychoanalyst Carmiña Ruiz-Medrano, M.Sc., and Dr.
Salvador Francisco Ruiz-Medrano, plus all the members of the G2 generation:
Ivan, Carlos Adrián, Diego, Aurora, Roberto, Gilberto, Mariana Mercedes, and Ethelia

Contents
Preface...............................................................................................................................................xi
Acknowledgments.......................................................................................................................xiii
The.author.......................................................................................................................................xv
Chapter 1 Introduction ..............................................................................................................1
References.........................................................................................................................................5
Chapter 2 Cell wall composition .............................................................................................7
2.1. Introduction............................................................................................................................7
2.2. Cell.wall.composition...........................................................................................................8
2.2.1. Cell.wall.polysaccharides.......................................................................................8
2.2.2. Cell.wall.proteins...................................................................................................11
2.2.3. Lipids.from.the.cell.walls.....................................................................................12
2.2.4. Pigments.................................................................................................................13
2.2.5. Inorganic.salts.in.the.walls..................................................................................14
2.3. Differentiation.of.the.cell.wall.chemical.composition...................................................15
References.......................................................................................................................................21
Chapter 3 Structure of the fungal cell wall .........................................................................29
3.1. Introduction..........................................................................................................................29
3.2. Methods.for.the.study.of.cell.wall.structure...................................................................29
3.3. Organization.of.the.cell.wall..............................................................................................31
3.4. The.cell.wall.surface............................................................................................................32
3.5. Microfibrilar.structure.of.the.cell.wall.............................................................................35
3.6. Associations.of.the.components.of.the.cell.wall.............................................................37
3.7. Mechanical.characteristics.of.the.fungal.cell.wall..........................................................39
References.......................................................................................................................................40
Chapter 4 Chitin .......................................................................................................................45
4.1. Introduction..........................................................................................................................45
4.2. Structure.of.chitin................................................................................................................45
4.3. Chitin.distribution.in.nature.............................................................................................46
4.4. Chitin.biosynthesis..............................................................................................................47
4.4.1. General.properties.of.fungal.chitin.synthases..................................................47
4.4.2. CHS.gene.multiplicity.in.fungi............................................................................49
4.4.3. Classification.and.possible.evolution.of.fungal.chitin.synthases...................51
4.4.4. Structure.of.chitin.synthases...............................................................................52
vii
viii Contents
4.4.5. Regulation.of.chitin.synthases............................................................................53
4.4.6. The.mechanism.of.chitin.biosynthesis...............................................................55
4.5.. Inhibitors.of.chitin.synthases............................................................................................56
4.6. Cytology.of.chitin.biosynthesis.........................................................................................57
References.......................................................................................................................................61
Chapter 5 Chitosan ...................................................................................................................69
5.1. General.characteristics.and.distribution.of.chitosan.....................................................69
5.2. The.functions.of.chitosan...................................................................................................69
5.3. Chitosan.biosynthesis.and.genetic.control.and.phylogenetic.relationships.of.
chitin.deacetylases...............................................................................................................70
References.......................................................................................................................................72
Chapter 6 Fungal glucans........................................................................................................73
6.1. Introduction..........................................................................................................................73
6.2. Structure.of.glucans............................................................................................................73
6.3. Role.of.fungal.glucans........................................................................................................76
6.4. Synthesis.of.glucans............................................................................................................77
6.4.1. Introduction............................................................................................................77
6.4.2. Introduction.to.the.synthesis.of.β-1,3-glucans..................................................78
6.4.3. Stimulators.and.inhibitors.of.β-1,3-glucan.synthases......................................80
6.4.4. Purification.of.β-1,3-glucan.synthases................................................................81
6.4.5. Genes.encoding.β-1,3-glucan.synthases.............................................................82
6.4.6. Structure.and.phylogenetic.characteristics.of.β-1,3-glucan.synthases..........84
6.4.7. Activation.of.β-1,3-glucan.synthases.by.a.small.GTPase.................................85
6.4.8. Cytological.aspects.of.the.synthesis.of.β-1,3-glucans.......................................86
6.5. Other.glycosyl.transferases.involved.in.the.synthesis.of.β-glucans............................87
6.6. Synthesis.of.β-1,6-glucans...................................................................................................89
6.7. Synthesis.of.α-1,3-glucans...................................................................................................91
References.......................................................................................................................................95
Chapter 7 Proteins ..................................................................................................................103
7.1. Introduction........................................................................................................................103
7.2. Chemical.structure.of.glycoproteins..............................................................................104
7.3. Different.classes.of.covalently.bound.glycoproteins....................................................106
7.3.1. GPI-proteins..........................................................................................................107
7.3.2. Proteins.with.internal.repeats...........................................................................107
7.3.3. Proteins.associated.with.the.wall.through.disulfide.bonds.........................108
7.4. Biosynthesis.of.glycoproteins..........................................................................................108
7.4.1. Synthesis.of.the.inner.core,.or.the.“dolichol.pathway”..................................109
7.4.2. Transfer.of.the.inner.core.to.proteins...............................................................111
7.4.3. Processing.and.maturation.of.the.N-linked.oligosaccharides......................111
7.4.4. Further.glycosylation.of.the.inner.core.to.form.the.outer.chain.of.
glycoproteins........................................................................................................112
7.4.5. O-glycosylation.of.mannoproteins...................................................................113
7.5. Synthesis.of.GPI-proteins.................................................................................................114
7.6. Nonclassical.protein.secretion.........................................................................................116
References.....................................................................................................................................117
Contents ix
Chapter 8 Cytological aspects of cell wall synthesis ......................................................123
8.1. Introduction........................................................................................................................123
8.2. Synthesis.of.secretory.and.membrane-bound.proteins...............................................123
8.3. Translocation.of.proteins.to.the.endoplasmic.reticulum.............................................125
8.4. The.importance.of.protein.glycosylation.in.translocation..........................................127
8.5. Transfer.of.proteins.along.the.endomembrane.system................................................129
8.5.1. From.ER.to.Golgi.................................................................................................131
8.5.2. Out.from.Golgi.....................................................................................................134
8.6. The.secretion.mechanism.in.filamentous.fungi...........................................................135
8.7. The.Spitzenkörper.............................................................................................................135
8.8. Vesicles.and.chitosomes.in.fungal.cell.wall.growth....................................................136
8.9. The.porosome.....................................................................................................................142
References.....................................................................................................................................143
Chapter 9 Cell wall growth and expansion .......................................................................149
9.1. Introduction........................................................................................................................149
9.2. Expansion.of.the.cell.wall.................................................................................................150
9.2.1. Introduction..........................................................................................................150
9.2.2. The.physical.stages.of.nascent.and.mature.wall.and.their.relation.to.
cell wall.expansion.and.fungal.growth............................................................150
9.2.3. Intermittent.growth.of.the.cell.wall..................................................................152
9.2.4. Mobilization.of.microvesicles.and.vesicles.involved.in.wall.growth..........153
9.2.5. Reinitiation.of.wall.growth................................................................................156
9.3. Polarization.of.cell.wall.synthesis.and.apical.growth..................................................162
9.3.1. Introduction..........................................................................................................162
9.3.2. Establishment.of.polarity...................................................................................162
9.3.3. Development.of.the.hyphal.shape.....................................................................165
References.....................................................................................................................................168