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Ectomycorrhizal Fungi Key Genera in Profile PDF

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John W. G. Cairney· Susan M. Chambers (Eds) Ectomycorrhizal Fungi: Key Genera in Profile Springer-Verlag Berlin Heidelberg GmbH John W. G. Cairney . Susan M. Chambers (Eds) Ectotnycorrhizal Fungi Key Genera in Profile With 30 Figures Springer Dr. John W.G. Cairney Dr. Susan M. Chambers Mycorrhiza Research Group School of Science University of Western Sydney (Nepean) PO Box 10 Kingswood NSW 2074 Australia ISBN 978-3-642-08490-4 Library of Congress CIP-data applied for Die Deutsche Bibliothek - CIP-Einheitsaufnahme Ectomycorrhizal fungi : key genera in profile I J. W. G. Cairney ; S. M. Chambers (eds.). ISBN 978-3-642-08490-4 ISBN 978-3-662-06827-4 (eBook) DOI 10.1007/978-3-662-06827-4 This work is subject to copyright. Ali rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitations, broad casting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag Berlin Heidelberg GmbH. Violations are liable for prosecution under the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1999 Originally published by Springer-Verlag Berlin Heidelberg New York in 1999 Softcover reprint of the hardcover 1s t edition 1999 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant pro tective laws and regulations and therefore free general use. Typesetting: BestSet Typesetter Ltd. Hong Kong Cover design: design & production GmbH, Heidelberg SPIN: 10643460 31/3136 - 5 4 3 2 1 0-printed on acid-free paper Preface It is becoming increasingly clear that communities of ectomycorrhizal fungi in natural and man-managed habitats are far more diverse than has pre viously been recognised. The development of morphotype analysis in the 1980s led to an appreciation that above-ground sporocarp diversity is a poor reflection on below-ground diversity at the root-tip level. The availability of the much higher resolving power of molecular methods in more recent years has revealed still further diversity in below-ground ectomycorrhizal fungal communities. Indeed, from data presented in a number of papers at the Second International Conference on Mycorrhiza in Uppsala, Sweden, in July 1998, it is apparent that we currently have a very poor appreciation of ecto mycorrhizal fungal diversity in most habitats. As mycorrhiza researchers struggle to come to grips with the functional significance of such diversity, we thought it timely to bring together a series of chapters, each dedicated to a particular ectomycorrhizal fungal genus. While previous reviews of mycor rhizal biology have sought to generalise aspects of the symbioses, the empha sis in this book is very much upon highlighting diversity in taxonomy, ecology, physiology and, where applicable, practical aspects of ectomycor rhizal fungi. We hope that this approach will facilitate a wider appreciation of ectomycorrhizal functioning, and stimulate further research into the func tional significance of fungal diversity in forest ecosystems. It will be clear to the reader that only a few genera have, for reasons of eco nomics or ease of laboratory manipulation, been investigated extensively. We thus have a reasonable knowledge of the biology of Cenococcum, Hebeloma, Laccaria, Paxillus, Pisolithus, Rhizopogon, Suillus, Thelephora and Tuber. Even within these genera, however, work has often concentrated on only single or a few taxa, and in many cases on a limited number of isolates of each. Several other genera, which are clearly widespread and important in ecological and/or economic terms, such as Amanita, Cantharellus, Hysterangium, Lactarius and Scleroderma, have received rather less attention so far. In each case, we con sidered that present knowledge was sufficient for a useful genus-specific chapter. Many other taxa such as Cortinarius and Russula, although probably key genera in many habitats, have not been included in this volume. While these taxa are clearly geographically widespread, we currently have an insufficient understanding of their biology to merit a broad review. In this context, some may question the inclusion of a chapter on resupinate genera. VI Preface Recent work, however, indicates that taxa forming resupinate sporocarps are important components of below-ground communities and we hope that their inclusion here will encourage further work on their biology. Finally, in the interests of saving space, and in order to allow as much data as possible to be presented in each chapter, we have used the abbreviation ECM throughout the book to replace ectomycorrhiza, ectomycorrhizas, ectomycorrhizae and ectomycorrhizal. October 1998 John W. G. Cairney Susan M. Chambers Contents Preface .................................................... V Chapter 1 Pisolithus 1 1.1 Introduction .......................................... 1 l.2 Taxonomy and Phylogenetic Relationships .................. 1 1.3 Ecology ............................................... 5 1.4 ECM Formation ........................................ 6 l.4.1 Host-Fungus Specificity ............................ 6 l.4.2 Genetics of the Interaction .......................... 6 l.4.3 Developmental Aspects of the Fungus-Host Interaction .................... :.................. 7 l.4.4 Symbiotic Functioning ............................. 11 l.5 Host Plant Growth Responses and Fungus-Derived Benefits ... 12 l.5.1 Growth Responses ................................. 12 l.5.2 Nutritional Benefits to the Host ...................... 14 l.5.3 Mineral Transformations ........................... 16 l.5.4 Influence on Host Plant Carbon Economy ............. 17 l.5.5 Non-nutritional Benefits to the Host .................. 18 l.6 Conclusions ........................................... 20 References ................................................ 21 Chapter 2 Suillus 33 2.1 Introduction ........................................... 33 2.2 Taxonomy and Phylogenetic Relationships .................. 33 2.3 Geographic Distribution and Host Specificity ............... 35 2.3.1 Geographic Distribution ............................ 35 2.3.2 Host Specificity ................................... 37 2.4 Community Structure, Population Biology and Genetics ...... 39 2.4.1 Community Structure ............................. 39 2.4.2 Population Biology ................................ 40 2.4.3 Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 2.5 ECM Morphology and Colonisation ....................... 44 2.5.1 ECM Morphology .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 2.5.2 ECM Colonisation ................................. 44 VIII Contents 2.6 Physiology ............................................ 45 2.6.1 Nutrient and Water Translocation .................... 45 2.6.2 Nitrogen Metabolism .............................. 46 2.6.3 Enzymatic Activities of Mycelia ...................... 47 2.6.4 Siderophore Production and Chemical Weathering 48 2.6.5 Carbon Metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 2.7 Anthropogenic Influences ................................ 49 2.7.1 Accumulation of Radionuclides ...................... 49 2.7.2 Nitrogen Deposition ............................... 50 2.7.3 Effects of Toxic Metals ............................. 50 2.7.4 Herbicides ....................................... 52 2.7.5 Bioremediation .................................... 52 2.8 Inoculation Experiments and Applications ................. 52 2.9 Interactions with Other Organisms ........................ 53 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 Chapter 3 Laccaria 65 3.1 Introduction .......................................... 65 3.2 Taxonomy and Ecology .................................. 66 3.2.1 Taxonomy........................................ 66 3.2.2 Ecology and Distribution ........................... 67 3.2.3 Population Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 3.2.4 Markers for Population Studies ...................... 70 3.3 ECM Formation ........................................ 73 3.3.1 Host-Fungus Specificity ............................ 73 3.3.2 Production of Sporocarps .......................... 73 3.3.3 Sexual Incompatibility and Behavior of Monokaryotic Strains. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 3.3.4 Genetics of ECM Formation ......................... 75 3.3.5 Molecular Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 3.3.6 ECM Development ................................ 78 3.4 Host Plant Growth Responses and Fungus-Derived Benefits ... 79 3.4.1 Nursery and Field Application ....................... 79 3.4.2 Interactions with Other Organisms ................... 81 References ................................................. 83 Chapter 4 Hebeloma 89 4.1 Introduction .......................................... 89 4.2 Taxonomy and Ecology ................................. 90 4.2.1 Taxonomy ....................................... 90 4.2.2 Saprophytic and ECM Hebeloma Species .............. 90 4.2.3 Host Range ....................................... 91 4.2.4 Geographical Distribution .......................... 92 4.2.5 Hebeloma Species as Successional Pioneers ............ 94 4.3 Life Cycle and Breeding Systems .......................... 95 Contents IX 4.3.1 Basidiospore Germination .......................... 95 4.3.2 Nuclear Status of Mycelia ........................... 96 4.3.3 Breeding Systems ............... . . . . . . . . . . . . . . . . . . 97 4.3.4 Sporocarp Formation .............................. 98 4.4 Quantitative Variation ................. . . . . . . . . . . . . . . . . . 100 4.4.1 Interspecific Variation .............................. 101 4.4.2 Intraspecific Variation .............................. 101 4.5 Population Structure and Dynamics ....................... 103 4.6 Genetics and Molecular Biology .......................... 107 4.6.1 Protoplasts and Mutagenesis ........................ 107 4.6.2 Gene Cloning and Genetic Transformation ............ 108 4.7 ECM Formation ......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 4.7.1 Developmental Aspects ....... . . . . . . . . . . . . . . . . . . . . . 109 4.7.2 Role of Fungal Auxin .............................. 112 4.8 Nutritional Interactions ................................. 114 4.8.1 Effect on Host Plant Growth. . . . . . . ... . . . . . . . . . . . . . . . 114 4.8.2 Influence on Host Carbon Economy .................. 114 4.8.3 Phosphorus Metabolism ............................ 115 4.8.4 Nitrogen Metabolism .............................. 116 4.9 Non-nutritional Interactions ............................. 119 4.10 Conclusions .......................................... 119 References 120 Chapter 5 Rhizopogon 129 5.1 Introduction .......................................... 129 5.2 Taxonomy ............................................ 129 5.3 Morphology and Anatomy of Rhizopogon ECM .............. 133 5.3.1 Pseudotsuga menziesii ECM ......................... 133 5.3.2 Pinus spp. ECM ................................... 135 5.3.3 Tsuga heterophylla ECM and Arbutoid Mycorrhizas ..... 136 5.4 Ecology .............................................. 137 5.4.1 Geographical Distribution and Host Specificity ......... 137 5.4.2 Habitats and Community Dynamics .................. 140 5.4.3 Sporocarp Productivity ............................ 142 5.4.4 Spore Biology .................................... 143 5.4.5 Mycophagy of Sporocarps .......................... 144 5.4.6 Interactions with Other Micro-organisms ............. 145 5.5 Physiology ............................................ 146 5.5.1 Growth in Culture and Physiological Traits ............ 146 5.5.2 Nutrient Translocation ............................. 148 5.5.3 Enzyme and Hormone Activities ..................... 149 5.6 Applications in Forestry ................................. 149 5.6.1 Seedling Inoculation ............................... 150 5.6.2 Benefits to Hosts .................................. 151 References 152 X Contents Chapter 6 Tuber 163 6.1 Introduction........................................... 163 6.2 Taxonomy and Life Cycle ................................ 163 6.3 Ecology............................................... 165 6.3.1 General Ecology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165 6.3.2 Tuber ECM and Bacteria ........................... 166 6.3.3 Spore Germination and Mycelial Growth .............. 167 6.4 ECM Formation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169 6.4.1 Host-Fungus Specificity ............................ 169 6.4.2 Developmental Aspects of the Fungus-Host Interaction ....................................... 170 6.4.3 Identification of ECM .............................. 172 6.5 Host Plant Growth Responses and Fungus-Derived Benefits ............................................... 175 6.5.1 Growth Responses ................................. 175 6.5.2 Nutritional Benefits to the Host ...................... 176 6.5.3 Non-nutritional Benefits to the Host .................. 177 6.6 Conclusions ........................................... 178 References ................................................ 179 Chapter 7 Scleroderma 187 7.1 Introduction .......................................... 187 7.2 Taxonomy and Ecology ................................. 187 7.3 ECM Formation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190 7.3.1 Host-Fungus Specificity ............................ 190 7.3.2 Developmental Aspects of the Fungus-Host Interaction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190 7.3.3 Symbiotic Functioning ............................. 191 7.4 Host Plant Growth Responses and Fungus-Derived Benefits ... 192 7.5 Nutritional Benefits to the Host and Influence on Host Carbon Economy .................................. 195 7.6 Non-nutritional Benefits to the Host ...................... 196 References ................................................ 197 Chapter 8 Amanita 201 8.1 Introduction........................................... 201 8.2 Taxonomy............................................. 201 8.2.1 Key to Subgenera and Sections ..................... 202 8.2.2 Subgenus Amanita ............................... 207 8.2.2.1 Section Amanita ................................ 207 8.2.2.2 Section Caesareae ................................ 207 8.2.2.3 Section Vaginatae ............................... 207 8.2.3 Subgenus Lepidella ............................... 209

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