The Mycota Edited by K. Esser and IW. Bennett Springer-Verlag Berlin Heidelberg GmbH The Mycota I Growth, Differentiation and Sexuality Ed. by IGH. Wesseis and F. Meinhardt II Genetics and Biotechnology Ed. by U. Kück III Biochemistry and Molecular Biology Ed. by R. Brambl and G Marzluf IV Environmental and Microbial Relationships Ed. by D. Wicklow and B. Sädersträm V Plant Relationships Ed. by G. Carroll and P. Tudzynski VI Human and Animal Relationships Ed. by D.H. Howard and ID. Miller VII Systematics and Evolution Ed. by D.I McLaughlin, E.G. McLaughlin, and P.A. Lemke t VIII Biology of the Fungal Cell Ed. by R.I Howard and N.A.R. Gow IX Fungal Associations Ed. by B. Hock X Industrial Applications Ed. by H.D. Osiewacz XI Agricultural Applications Ed. by F. Kempken XII Human Fungal Pathogens Ed. by IE. Domer and GS. Kobayashi The Mycota A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research Edited by K. Esser and IW. Bennett XI Agricultural Applications Volume Editor: F. Kempken With 44 Figures, 2 in Color, and 29 Tables Springer Series Editors Professor Dr. Dr. h.c. mult. KARL ESSER Allgemeine Botanik Ruhr-Universität 44780 Bochum, Germany Tel.: +49(234)32-22211 Fax: +49(234)32-14211 e-mail: [email protected] Professor Dr. JOAN W. BENNETI Department of Cell and Molecular Biology Tulane University New Orleans, Louisiana 70118 USA Tel.: +1(504)865-5546 Fax: +1(504)865-6785 e-mail: [email protected] Volume Editor Professor Dr. Frank Kempken Christian-Albrechts-Universität zu Kiel Botanisches Institut Abteilung Botanische Genetik und Molekularbiologie Olshausenstr. 40 24098 Kiel, Germany Tel: +49(431) 8804274 Fax: +49(431) 880 4248 e-mail: [email protected] ISBN 978-3-642-07650-3 ISBN 978-3-662-03059-2 (eBook) DOI 10.1007/978-3-662-03059-2 Library of Congress Cataloging-in-Publication Data The Mycota. Includes bibliographical references and index. Contents: 1. Growth, differentiation, and sexuality/editors, J.G.H. Wesseis and F. Meinhardt - 2. Genetics and biotechnology. 1. Mycology. 2. Fungi. 3. Mycology - Research. 4. Research. 1. Esser, Karl, 1924-. H. Lemke, Paul A., 1937- . QK603.M87 1994 589.2 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concemed, specifi cally 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 provi sions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. http://www.springer.de © Springer-Verlag Berlin Heidelberg 2002 Originally published by Springer-Verlag Berlin Heidelberg New York in 2002. Softcover reprint ofthe hardcover 1st edition 2002 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 there fore free for general use. Production Editor: PRO EDIT GmbH, Heidelberg, Germany Cover design: Springer-Verlag, E. Kirchner Typesetting: SNP Best-set Typesetter Ltd., Hong Kong Printed on acid-free paper SPIN: 10664995 3113130/So 5 4 3 2 1 0 Series Preface Mycology, the study of fungi, originated as a subdiscipline of botany and was a descrip tive discipline, largely neglected as an experimental science until the early years of this century. A seminal paper by Blakeslee in 1904 provided evidence for self incompatibility, termed "heterothallism", and stimulated interest in studies related to the control of sexual reproduction in fungi by mating-type specificities. Soon to follow was the demonstration that sexually reproducing fungi exhibit Mendelian inheritance and that it was possible to conduct formal genetic analysis with fungi. The names Burgeff, Kniep and Lindegren are all associated with this early period of fungal genetics research. These studies and the discovery of penicillin by Fleming, who shared a Nobel Prize in 1945, provided further impetus for experimental research with fungi. Thus began a period of interest in mutation induction and analysis of mutants for biochemical traits. Such fundamental research, conducted largely with Neurospora crassa, led to the one gene: one enzyme hypothesis and to a second Nobel Prize for fungal research awarded to Beadle and Tatum in 1958. Fundamental research in biochemical genetics was extended to other fungi, especially to Saccharomyces cerevisiae, and by the mid-1960s fungal systems were much favored for studies in eukaryotic molecular biology and were soon able to compete with bacterial systems in the molecular arena. The experimental achievements in research on the genetics and molecular biology of fungi have benefited more generally studies in the related fields of fungal bio chemistry, plant pathology, medical mycology, and systematics. Today, there is much interest in the genetic manipulation of fungi for applied research. This current interest in biotechnical genetics has been augmented by the development of DNA media ted transformation systems in fungi and by an understanding of gene expression and regulation at the molecular level. Applied research initiatives involving fungi extend broadly to areas of interest not only to industry but to agricultural and envi ronmental sciences as weB. It is this burgeoning interest in fungi as experimental systems for applied as weIl as basic research that has prompted publication of this se ries of books under the title The Mycota. This title knowingly relegates fungi into aseparate realm, distinct from that of either plants, animals, or protozoa. For consistency throughout this Series of Volumes the names adopted for major groups of fungi (representative genera in paren theses) are as folIows: Pseudomycota Division: Oomycota (Achlya, Phytophthora, Pythium) Division: Hyphochytriomycota Eumycota Division: Chytridiomycota (Allomyces) Division: Zygomycota (Mucor, Phycomyces, Blakeslea) Division: Dikaryomycota VI Series Preface Subdivision: Ascomycotina Class: Saccharomycetes (Saccharomyces, Schizosaccharomyces) Class: Ascomycetes (Neurospora, Podospora, Aspergillus) Subdivision: Basidiomycotina Class: Heterobasidiomycetes (Ustilago, Tremella) Class: Homobasidiomycetes (Schizophyllum, Coprinus) We have made the decision to exclude from The Mycota the slime molds whieh, although they have traditional and strong ties to mycology, truly represent nonfungal forms insofar as they ingest nutrients by phagocytosis, lack a cell wall during the assim ilative phase, and clearly show affinities with certain protozoan taxa. The Series throughout will address three basic questions: what are the fungi, what do they do, and what is their relevance to human affairs? Such a focused and com prehensive treatment of the fungi is long overdue in the opinion of the editors. A volume devoted to systematies would ordinarily have been the first to appear in this Series. However, the scope of such a volume, coupled with the need to give serious and sustained consideration to any reclassification of major fungal groups, has delayed early publication. We wish, however, to provide a preamble on the nature of fungi, to acquaint readers who are unfamiliar with fungi with certain characteristics that are representative of these organisms and which make them attractive subjects for experimentation. The fungi represent a heterogeneous assemblage of eukaryotic microorganisms. Fungal metabolism is characteristically heterotrophic or assimilative for organic car bon and some none lernen tal source of nitrogen. Fungal cells characteristically imbibe or absorb, rather than ingest, nutrients and they have rigid cell walls. The vast major ity of fungi are haploid organisms reproducing either sexually or asexually through spores. The spore forms and details on their method of production have been used to delineate most fungal taxa. Although there is a multitude of spore forms, fungal spores are basically only of two types: (i) asexual spores are formed following mitosis (mitospores) and culminate vegetative growth, and (ii) sexual spores are formed fol lowing meiosis (meiospores) and are borne in or upon specialized generative struc tures, the latter frequently clustered in a fruit body. The vegetative forms of fungi are either unicellular, yeasts are an example, or hyphal; the latter may be branched to form an extensive mycelium. Regardless of these details, it is the accessibility of spores, especially the direct recovery of meiospores coupled with extended vegetative haploidy, that have made fungi especially attractive as objects for experimental research. The ability of fungi, especially the saprobic fungi, to absorb and grow on rather simple and defined substrates and to convert these substances, not only into essential metabolites but into important secondary metabolites, is also noteworthy. The meta bolic capacities of fungi have attracted much interest in natural products chemistry and in the production of antibiotics and other bioactive compounds. Fungi, especially yeasts, are important in fermentation processes. Other fungi are important in the production of enzymes, citric acid and other organic compounds as weH as in the fermentation of foods. Fungi have invaded every conceivable ecological niche. Saprobie forms abound, especiaHy in the decay of organie debris. Pathogenic forms exist with both plant and animal hosts. Fungi even grow on other fungi. They are found in aquatie as weH as soil environments, and their spores may pollute the air. Some are edible; others are poi sonous. Many are variously associated with plants as copartners in the formation of lichens and mycorrhizae, as symbiotic endophytes or as overt pathogens. Association with animal systems varies; examples include the predaceous fungi that trap nema todes, the microfungi that grow in the an aerobic environment of the rumen, the many Series Preface VII insectassociated fungi and the medically important pathogens affiicting humans. Yes, fungi are ubiquitous and important. There are many fungi, conservative estimates are in the order of 100000 species, and there are many ways to study them, from descriptive accounts of organisms found in nature to laboratory experimentation at the cellular and molecular level. All such studies expand our knowledge of fungi and of fungal processes and improve our ability to utilize and to control fungi for the benefit of humankind. We have invited leading research specialists in the field of mycology to contribute to this Series. We are especially indebted and grateful for the initiative and leadership shown by the Volume Editors in selecting topics and assembling the experts. We have all been a bit ambitious in producing these Volumes on a timely basis and therein lies the possibility of mistakes and oversights in this first edition. We encourage the readership to draw our attention to any error, omission or inconsistency in this Series in order that improvements can be made in any subsequent edition. Finally, we wish to acknowledge the willingness of Springer-Verlag to host this project, which is envisioned to require more than 5 years of effort and the publication of at least nine Volumes. Bochum, Germany KARL ESSER Auburn, AL, USA PAUL A. LEMKE April 1994 Series Editors Addendum to the Series Preface In early 1989, encouraged by Dieter Czeschlik, Springer-Verlag, Paul A. Lemke and I began to plan The Mycota. The first volume was released in 1994, other volumes followed in the subsequent years. Also on behalf of Paul, I would like to take this opportunity to thank Dieter Czeschlik, his coHeague Andrea Schlitzberger, and Springer-Verlag for their help in realizing the enterprise and for their exceHent co operation for many years. Unfortunately, after a long and serious illness, Paul A. Lemke died in November 1995. Without his expertise, his talent for organization and his capability to grasp the essentials, we would not have been able to work out a concept for the volumes of the series and to acquire the current team of competent volume editors. He was an out standing scientist interested in many fields. Together with the volume editors, authors, and Springer-Verlag, I mourn the loss of a very good and reliable friend and colleague. Since the first Volumes of The Mycota were weH accepted by the scientific com munity, the publisher suggested to extend this series. For Volumes X, XI and XII I was able to win Joan W. Bennett as serial co-editor. Bochum, Germany KARL ESSER New Orleans, LA, USA JOAN W. BENNETI July 2001 Volume Preface The development of agriculture was an essential prerequisite to the establishment of permanent settlements and eventually complex human society. At all times, and pos sibly even more so now, humanity depended on the annual crop yield, which may be influenced by weeds, pathogens or poor weather conditions. Fungi are important plant pathogens and can reduce yield significantly. In fact, many examples can be cited where fungal pathogens have actually made history, e.g., the infections by the ergot fungus Claviceps purpurea causing ergotismen in the middle ages or the disastrous Phytoph thora infections of potato in nineteenth century Ireland, leading to the emigration of millions of Irish people to the United States of America. In addition, the occurrence of mildew can severally spoil food and fodder and mycotoxins produced by these fungi may cause illness or even death. Clearly, fungi had and still have a tremendous impact on humanity. However, while only a minority of fungi are pathogens, many others can be quite useful, e.g., to nutritionally enrich straw or to ferment food and drink. In this volume the relevance of fungi for agriculture is discussed in 18 chapters, wh ich are divided into four sections: (1) food and fodder production, (2) mycotoxins and detoxification, (3) disease control, diagnostic, and management, and finally (4) update on host-parasite interactions. Chapters 1-3 discuss various aspects of food and fodder production, featuring the application and potential of mushrooms, straw enrichment and food or crop spoilage. The first article by Paul Horgan and Alan Castle provides insight into the use and genetics of mushrooms, especially Agaricus. Dusan Jale contributed a chapter about straw enrichment by fungi, giving many details about this field of research. Jan Dijksterhuis and Rob Samson present the current knowledge about the important problem of food and crop spoilage. I should add that other aspects of food production have already been reviewed in Vol. X (Industrial Applications) of The Mycota and are therefore not covered again here. The second section contains two chapters which are devoted to knowledge about the biosynthesis of mycotoxins and the use of fungi in organopollutant degradation. Contamination with mycotoxins of course is a problem, particularly in humid climates, and may increase when anti-fungal agents are not available or are rejected, as is the case in organic farming. Nancy Keller and colleagues provide a detailed insight into the synthesis of some mycotoxins. Organopollutant degradation, the second chapter in this section, has a high potential for future pollution management and was written by Daniel Cullen. As fungal phytopathogens are of great concern in agriculture, a large section of this volume deals with various aspects of biological control (three chapters), diagnos tics (two chapters) and disease management (three chapters). Fungal biological control is covered by Yigal Elad and Stanley Freeman, insect control by Tariq Butt and weed control by Harry Evans. Disease control is another focus with emphasis on the example of Magnaporthe grisea given by Nicholas Talbot. Another chapter, by Diana Fernandez and Thierry Langin, deals with the use of repeated DNA and transposons as diagnostic tools, a rather recent development. Finally, disease management is covered in three chapters XII \1alurne Preface dealing with important fungal pathogens such as Phoma, by Kerstin Voigt and Johannes Wöstemeyer, Fusarium, by Kerstin Voigt, and rusts and powdery mildew by Holger Deising and collaborators. In the fourth section, host-parasite interactions are the main topic. This section has been named "update on ..." to acknowledge work presented in a previous volume of The Mycota (Vol. V). In this volume, five chapters present the current knowl edge in this important field discussing relevant issues like signal transduction by Michael Bölker, avirulence determinants by Wolfgang Knogge, phytotoxins by Dan Panacchione and colleagues, and cell wall degradation, by Jan van Kan and collabora tors. The final chapter, written by Jacques Mugnier, gives insight into the co-evolution of pathogenic fungi and grass hosts. As this volume is restricted in size, certainly not all aspects of fungal applications in agriculture are covered. However, the 18 chapters provide an important insight into this area of research and, I sincerely hope that it will serve as a guide for readers from outside the field and as a valuable reference for those to this type of research. Finally, I wish to express my gratitude to all contributors to this volume. Kiel, Germany, April 2002 FRANK KEMPKEN Volume Editor