FUNGI IN ECOSYSTEM PROCESSES - CHAPTER 1 potx

Fungi in Ecosystem Processes John Dighton Institute of Marine and Coastal Sciences Cook College and Rutgers Pinelands Field Station New Lisbon, New Jersey, U.S.A. MARCEL MARCEL DEKKER, INC. NEW YORK • BASEL Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress. ISBN: 0-8247-4244-3 This book is printed on acid-free paper. Headquarters Marcel Dekker, Inc. 270 Madison Avenue, New York, NY 10016 tel: 212-696-9000; fax: 212-685-4540 Eastern Hemisphere Distribution Marcel Dekker AG Hutgasse 4, Postfach 812, CH-4001 Basel, Switzerland tel: 41-61-260-6300; fax: 41-61-260-6333 World Wide Web http://www.dekker.com The publisher offers discounts on this book when ordered in bulk quantities. For more information, write to Special Sales/Professional Marketing at the headquarters address above. Copyright q 2003 by Marcel Dekker, Inc. All Rights Reserved. Neither this book nor any part may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, microfilming, and recording, or by any information storage and retrieval system, without permission in writing from the publisher. Current printing (last digit): 10987654321 PRINTED IN THE UNITED STATES OF AMERICA Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. MYCOLOGY SERIES Editor J. W. Bennett Professor Department of Cell and Molecular Biology Tulane University New Orleans, Louisiana 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 Ecosys- tem, 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 Koltm and Michael J Leibowitz 8 Molecular Industrial Mycology: Systems and Applications for Fila- mentous Fungi, edited by Sally A Leong and Randy M Berka 9 The Fungal Community Its Organization and Role in the Eco- system, 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 Gunther Wmkelmann and Dennis R Wmge 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 Osie- wacz Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. 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 Additional Volumes in Preparation 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 Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. Preface Why did I decide to write this book? As an undergraduate student I could not make up my mind whether I wanted to be a zoologist or a botanist, so I decided to adopt ecology, in its broadest sense, as my area of interest. This led me to think about interactions among organisms and to try to look at ecosystems from a holistic, rather than from an autecological, point of view. As someone with little formal training in mycology, my interest in fungi started during my doctoral research, especially when attending university-wide lectures by C. T. Ingold, given at the University of London. My former job, at the then Institute of Terrestrial Ecology at Merlewood , UK, brought me into contact with the work on ectomycorrhizae, decomposition, and nutrient cycling in temperate and subtropical forest ecosystems. During this time, I had many fruitful discussions with fungal ecologists in both the British Mycological Society and the international world of mycorrhizal researchers. In particular, I remember animated discussions with Alan Rayner, who is a font of knowledge, inspiration, and encouragement. As a result of this history, I have been fascinated by the multifarious ways in which fungi interact with other organisms and function in moderating the processes occurring in ecosystems. With my move to Rutgers Univer sity, I initially taught my first graduate course, Fungi in Ecosystems, during my first semester, while all my books and notes were on a ship crossing the Atlantic. The cont inued development of this course has been the impetus for this book. As I could find no textbook for my students that really approached the subject of myco logy from an ecosystem point of view, I decided to put my ideas on the importance of fungi in ecosy stem processes into a volume that could be used by upper-level students and Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. researchers. This course and its variants have been the main thrust of my teaching at Rutgers. This has not been a sole venture as, along with my interactions with Jim White and Peter Oudemans, we are gradually building a strong and interactive series of mycology courses in our university. In this book I have started with the list of functions that Alan Rayner suggests that fungi perform in ecosystems (introduction to the second edition of The Fungal Community: Its Structure and Function, edited by George C. Carroll and Donald T. Wicklow, (Marcel Dekker, 1992)). I have attempted to elaborate on these functional roles of fungi and tried to show how the world would function less satisfactorily in the absence of fungi. Given the vast range of fungal literature I have not been able to do much more than scratch the surface of the subject. However, I hope that the more than 1300 references that I have cited will act as a means for interested readers to delve further into the literature on any specific subject area. For the upper-level students and researchers to whom this book is targeted, I hope that it will stimulate thoughts beyond the narrow confines of their immediate research questions and allow them to place their work in the wider context of the functioning of ecosystems. I believe that it is only by the greater understanding of the linkages between organisms and the processes they control in the ecosystem that we can appreciate the beauty of the living world around us, appreciate the impacts that we are having on ecosystems, and obtain the understanding of the function of component organisms that will allow us to protect and preserve nature. In attempting to summarize the vast literature, I have written the text in my own words, but have relied heavily on previously published literature for figures and tables to support my commentary. Most tables have been reconstructed and almost all figures have been redrawn from their original in an effort to simplify the information presented in the originals. For the statistical purist, I hope that my removal of error bars and indications of significant differences will be forgiven for the sake of simplicity of presentation. In the main, the data shown in figures represent statistically significant interactions. Although the fig ures and tables represent the key message I wish to convey, they are no substitute for the original data and publications. I encourage the interested reader to explore further by consulting the original publications to obtain more information that I can impart in this book. I dedi cate this book to my uncle, Wally Champkin, whose enthusiasm for natural history fueled my interest in ecology. As a child I was constantly amazed that he could put a name to most of the plants, insects, and birds we saw on our walks. I could watch for hours his cine films and stills of birds, flowers, moths, and butterflies. I owe a debt of gratitude to my parents, who encouraged and supported me in my studies and allowed me to pursue my own interests in biology. I especially thank my wife, Joan, and daughter, Gail, who have supported my career, moved with me to the United States, and encouraged me in Prefaceiv Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. the writing of this book. I could not have completed this task without them. Finally, I wish to thank Bob Evans, who commented on the first drafts of my work here, and to the students and colleagues in my research group, who have both given me encouragement and tolerated my absences during the creation of this book. John Dighton Preface v Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. Contents Preface 1. Introduction 1.1 Why Fungi? 1.2 What Are Ecosystems and Ecosys tem Functions? 1.3 What Are Fungi? 1.4 Specific Ecosystem Services Carried Out by Fungi 1.5 Concluding Remarks References 2. Fungi and Primary Productivity: Making Nutrients Available 2.1 Soils and Nutrient Availability 2.2 Nutrient Availability in Aquatic and Marine Ecosystems 2.3 Concluding Remarks References 3. Fungi and Primary Productivity: Plant Growth and Carbon Fixation 3.1 The Role of Lichens in Primary Production 3.2 The Role of Mycorrhizae in Plant Primary Production 3.3 The Role of Fungal Pathogens in Plant Primary Production Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. 3.4 Interactions Among Mycorrhizae and Plant Pathogens 3.5 Synopsis and Outlook References 4. Fungi, Secondary Productivity, and Other Fungal–Faunal Interactions 4.1 Fungi as Food for Secondary Producers: Positive Impacts on Faunal Productivity 4.2 Effects of Grazing on Fungi and Fungal-Mediated Processes: Negative and Positive Feedbacks 4.3 Specific Fungal–Faunal Interactions 4.4 Fungi as Animal Pathogens: Negative Impacts on Faunal Productivity 4.5 Fungal– Faunal Interactions in Aquatic and Marine Ecosystems 4.6 Summary and Conclusions References 5. Fungi and Population and Community Regulation 5.1 Mycorrhizae and Plant Successions 5.2 Mycorrhizae and Plant Fitness 5.3 Plant Pathogens and Plant Fitness 5.4 Saprotroph–Pathogen Interactions: Biocontrol 5.5 Mycorrhizae–Pathogen Interactions 5.6 Endophytes and Antiherbivore Action 5.7 Nematophagous Fungi and Animal Pathogens 5.8 Allelopathy 5.9 Summary and Discussion References 6. Fungal Interactions with Humans 6.1 Introduction 6.2 Fungi and Acidifying Pollutants 6.3 Fungi and Heavy Metals 6.4 Fungi and Radionuclides 6.5 Fungi and Climate Change 6.6 Concluding Remarks References 7. Synopsis and Outlook to the Future 7.1 Introduction 7.2 The Ecosystem Contentsviii Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. 7.3 The Fungal Organism 7.4 The Fungal Community 7.5 Perturbations 7.6 Fungi in Ecosystems: What Next? References Contents ix Copyright 2003 by Marcel Dekker, Inc. All Rights Reserved. [...]... the future Appl Soil Ecol 11 :11 1 – 12 6 Hyde, K D., Gareth Jones, E B., Leano, E., Pointing, S B., Poonyth, A D., Vrijmoed, L L P (19 98) Role of fungi in marine ecosystems Biodiv Conserv 7 :11 47 11 61 Jennings, D H (19 76) Transport and translocation in filamentous fungi In: Smith, J E., Berry, D R., eds The Filamentous Fungi Vol 2 London: Edward Arnold, pp 32–64 Jennings, D H (19 82) The movement of Serpula... M (19 95) Soil organisms as engineers: microsite modulation of macroscale processes In: Jones, C G., Lawton, J H., eds Linking Species and Ecosystems New York: Chapman & Hall, pp 94– 10 6 Andrews, J H (19 92) Fungal life-history strategies In: Carrol, G C., Wicklow, D T., eds The Fungal Community: Its Organization and Role in the Ecosystem New York: Marcel Dekker, pp 11 9– 14 5 Arnolds, E (19 91) Decline... corals, clonal plants 14 Chapter 1 (production of grazing animal biomass) in ecosystems The functional role of fungi in a terrestrial ecosystem is represented in Fig 1. 4, which shows the interactions between above- and below-ground components of the ecosystem, plants, animals, and the abiotic environment FIGURE 1. 4 Representation of a plant – soil – fungal interaction in a terrestrial ecosystem The model... development in Pinus sylvestris L seedlings New Phytol 11 9:405 – 411 Wright, S F., Updahaya, A (19 98) A survey of soils for aggregate stability and glomalin, a glycoprotein produced by hyphae of arbuscular mycorrhizal fungi Plant Soil 19 8:97– 10 7 Zak, J C., Willig, M R., Moorhead, D L., Wildman, H G (19 94) Functional diversity of microbial communities: a quantitative approach Soil Biol Biochem 26 :11 01 11 08... the nonliving components of the environment that make fungi a key group of organisms in the regulation of ecosystem processes Fungi are important in driving or controlling the mineral and energy cycling within ecosystems and in uencing the composition of other organisms within the ecosystem Copyright 2003 by Marcel Dekker, Inc All Rights Reserved Introduction 7 needed to build cytoplasm and structural... their role in water transport Nature 287:834– 836 Ekschmitt, K., Griffiths, B S (19 98) Soil biodiversity and its implications for ecosystem functioning in a heterogeneous and variable environment Appl Soil Ecol 10 :2 01 215 Fogel, R (19 76) Ecological studies of hypogeous fungi II Sporocarp phenology in a western Oregon Douglas-fir stand Can J Bot 54 :11 52 – 11 62 Frankland, J.C (19 92) Mechanisms in fungal... fruiting structures (represented by mushrooms) are a food source for grazing animals, thus in uencing the fitness of individuals and regulating animal populations Source: Adapted from Dighton and Boddy (19 89) Copyright 2003 by Marcel Dekker, Inc All Rights Reserved Introduction 15 1. 4 SPECIFIC ECOSYSTEM SERVICES CARRIED OUT BY FUNGI 1. 4 .1 Making Nutrients Available As with other micro-organisms, fungi. .. Carreiro, M M (19 97) Soil heavy metal concentrations, microbial biomass and enzyme activities in a contaminated grassland ecosystem Soil Biol Biochem 29 :17 9– 19 0 Lavelle, P (19 97) Faunal activities and soil processes: adaptive strategies that determine ecosystem function Adv Ecol Res 27:93 13 2 Lawton, J H., Jones, C G (19 95) Linking species and ecosystems: organisms as ecosystem engineers In: Jones, C... (19 91) Heirarchy theory as a guide to mycorhizal research on large-scale problems Environ Pollut 73:2 71 – 284 Pianka, E R (19 70) On r- and K-selection Am Nat 10 4:592– 597 Pirozynski, K A., Malloch, D W (19 75) The origin of land plants: a matter of mycotropism Biosystems 6 :15 3– 16 4 Rayner, A D M (19 91) The challenge of the individualistic mycelium Mycologia 83:48 – 71 Rayner, A D M (19 92) Introduction In: ... in uences of water chemistry Ecology 76 :14 33 – 14 45 Copyright 2003 by Marcel Dekker, Inc All Rights Reserved Introduction 25 Tilman, D., Wedin, D., Knops, J (19 96) Productivity and sustainability in uenced by biodiversity in grassland ecosystems Nature 379: 718 – 720 Trappe, J M (19 88) Lessons from alpine fungi Mycologia 80 :1 10 Trappe, J M., Maser, C (19 76) Germination of spores of Glomus macrocarpus . Reserved. Contents Preface 1. Introduction 1. 1 Why Fungi? 1. 2 What Are Ecosystems and Ecosys tem Functions? 1. 3 What Are Fungi? 1. 4 Specific Ecosystem Services Carried Out by Fungi 1. 5 Concluding Remarks References 2. Fungi. 4, Postfach 812 , CH-40 01 Basel, Switzerland tel: 4 1- 6 1- 2 6 0-6 300; fax: 4 1- 6 1- 2 6 0-6 333 World Wide Web http://www.dekker.com The publisher offers discounts on this book when ordered in bulk quantities Congress. ISBN: 0-8 24 7-4 24 4-3 This book is printed on acid-free paper. Headquarters Marcel Dekker, Inc. 270 Madison Avenue, New York, NY 10 016 tel: 21 2-6 9 6-9 000; fax: 21 2-6 8 5-4 540 Eastern Hemisphere