Olaf Schmidt Woodand Tree Fungi Biology, Damage, Protection, and Use Olaf Schmidt Wood and Tree Fungi Biology, Damage, Protection, and Use With 74 Figures, 12 in Colors, and 49 Tables 123 Professor Dr. Olaf Schmidt Universität Hamburg Zentrum Holzwirtschaft Abteilung Holzbiologie Leuschnerstraße 91 21031 Hamburg Germany [email protected] Cover: Fruit body of serpula lacrymans and electrophoresis gel demontstrating species-specific priming PCR. Library of Congress Control Number: 2006920787 ISBN-10 3-540-32138-1 Springer Berlin Heidelberg New York ISBN-13 978-3-540-32138-5 Springer Berlin Heidelberg New York This work is subjecttocopyright. All rights reserved, whetherthe whole orpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, 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. Violations are liable for prosecution under the German Copyright Law. Springer is a part of Springer Science + Business Media springer.com © Springer-Verlag Berlin Heidelberg 2006 Printed in Germany 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 protective laws and regulations and therefore free for general use. Editor: Dr. Dieter Czeschlik, Heidelberg, Germany Desk editor: Dr. Andrea Schlitzberger, Heidelberg, Germany Cover design: design & production, Heidelberg, Germany Typesetting and production: LE-TEXJelonek,Schmidt& Vöckler GbR, Leipzig, Germany 31/3100/YL – 543210–Printedonacid-freepaper Foreword Wood, as a raw material and a renewable biomass, has had great importance for thousands of years. Under suitable conditions, however, it is also easily degradable as part of the biological cycle. The processes of decomposition by fungi, and measures for protection against them, have been studied for quite a long time. The resulting knowledge on the causes and effects of wood degradation can hardly be overlooked. For more than 30 years, Olaf Schmidt has investigated the causes and effects of wood degradation by fungi and bacteria. Pioneering contributions have also been made in several fields, such as wood-inhabiting bacteria and molecular methods for fungal identification. Laboratory work is accompanied by teaching the field of wood deterioration by microorganisms, thus contributing to the broad spectrum of information accumulated. “WoodandTreeFungi”byOlafSchmidtpresentsthemostcomprehensive treatise on the fundamentals, causes, and consequences of decomposition of wood as well as measures for its prevention. The 1,400 references give an overlook of the vast amount of information evaluated. For a long time to come this book will be the competent source of knowledge about the fascinating interactions between wood and microorganisms. Walter Liese Preface This book is the updated revision of the German edition “Holz- und Baumpilze” from 1994. Errors were corrected and new results were included. Particularly the chapter “Identification” was supplemented by molecular techniques. I re- alize that a one-author book on a relatively broad topic must include errors and also may have ignored recent literature. Strictly speaking, one should only write about things that they have experienced themselves, in the case of point this only concerns some aspects of bacteria and those fungi which inhabit the xylem of dead wood. Thus, current “secondary literature” was used for those chapters that are “on the edge” of my own research interest. For better readability, the authors of fungal names are not mentioned in the text, but summarized in an appendix. Fungal synonyms are also not given. These are available from Index Fungorum (www.indexfungorum.org/names/ names.asp). Fungal names cited from (older) publications were transferred to the current version. Thanks for general advice go to Prof. Dr. Dr. h.c. mult. Walter Liese, for crit- ical reading to Prof. Dr. Dirk Dujesiefken (Chap. 8.2), Dr. Hubert Willeitner, and Dr. Peter Jüngel (Chap. 7.4), to Mrs. Ute Moreth for providing experimen- tal data, Dr. Tobias Huckfeldt for several photographs, many colleagues for permission to use their photographs, Mrs. Christina Waitkus for transferring many pictures in an electronic version, to Springer-Verlag, particularly Mrs. UrsulaGrammandDr.D.Czeschlik,forco-operation,toMr.JardiMullinaxfor making my English understandable, and to Mrs. Cornelia Gründer for careful printing. Hamburg, December 2005 Olaf Schmidt Contents 1Introduction 1 2 Biology 3 2.1 Cytology and Morphology.................................................... 3 2.2 Growth and Spreading ......................................................... 10 2.2.1 Vegetative Growth ..................................................... 10 2.2.2 Reproduction of Deuteromycetes ................................. 16 2.2.3 Sexual Reproduction.................................................. 18 2.2.4 Fruit Body Formation ................................................ 24 2.2.5 Production, Dispersal and Germination of Spores .......... 25 2.3 Sexuality............................................................................ 26 2.4 Identification...................................................................... 31 2.4.1 Traditional Methods .................................................. 31 2.4.2 Molecular Methods.................................................... 33 2.5 Classification...................................................................... 47 3 Physiology 53 3.1 Nutrients ........................................................................... 53 3.2 Air .................................................................................... 58 3.3 Wood Moisture Content ....................................................... 60 3.4 Temperature....................................................................... 67 3.5 pH Value and Acid Production by Fungi ................................. 70 3.6 Light and Force of Gravity .................................................... 74 3.7 Restrictions of Physiological Data.......................................... 77 3.8 Competition and Interactions Between Organisms................... 79 3.8.1 Antagonisms, Synergisms, and Succession .................... 79 3.8.2 Mycorrhiza and Lichens ............................................. 82 4 WoodCell Wall Degradation 87 4.1 Enzymes and Low Molecular Agents ...................................... 87 4.2 Pectin Degradation.............................................................. 92 4.3 Degradation of Hemicelluloses.............................................. 93 4.4 Cellulose Degradation.......................................................... 95 4.5 Lignin Degradation ............................................................. 99 XContents 5 Damages by Viruses and Bacteria 109 5.1 Viruses .............................................................................. 109 5.2 Bacteria ............................................................................. 109 6 Wood Discoloration 119 6.1 Molding............................................................................. 121 6.2 Blue Stain........................................................................... 125 6.3 Red Streaking ..................................................................... 129 6.4 Protection.......................................................................... 131 7 WoodRot 135 7.1 Brown Rot.......................................................................... 135 7.2 White Rot........................................................................... 138 7.3 Soft Rot ............................................................................. 142 7.4 Protection.......................................................................... 146 8 Habitat of WoodFungi 161 8.1 Fungal Damage to Living Trees.............................................. 161 8.1.1 Bark Diseases............................................................ 163 8.1.2 Wilt Diseases ............................................................ 168 8.2 Tree Wounds and Tree Care .................................................. 173 8.2.1 Wounds and Defense Against Discoloration and Decay ... 173 8.2.2 Pruning.................................................................... 177 8.2.3 Wound Treatment...................................................... 178 8.2.4 Detection of Tree and Wood Damages .......................... 179 8.3 Tree Rots by Macrofungi ...................................................... 183 8.3.1 Armillaria Species ..................................................... 186 8.3.2 Heterobasidion annosum s.l......................................... 189 8.3.3 Stereum sanguinolentum ............................................ 195 8.3.4 Fomes fomentarius ..................................................... 195 8.3.5 Laetiporus sulphureus ................................................ 197 8.3.6 Meripilus giganteus .................................................... 197 8.3.7 Phaeolus schweinitzii.................................................. 198 8.3.8 Phellinus pini ............................................................ 198 8.3.9 Piptoporus betulinus