Table Of ContentEcological Studies, Vol. 157
Analysis and Synthesis
Edited by
I.T. Baldwin, Jena, Germany
M.M. Caldwell, Logan, USA
G. Heldmaier, Marburg, Germany
0.1. Lange, Wiirzburg, Germany
H.A. Mooney, Stanford, USA
E.-D. Schulze, Jena, Germany
U. Sommer, Kiel, Germany
Ecological Studies
Volumes published since 1997 are listed at the end of this book.
Springer-Verlag Berlin Heidelberg GmbH
M.G.A. van der Heijden I.R. Sanders (Eds.)
Mycorrhizal Ecology
1s t Edition 2002, 2nd Printing 2003
With 59 Figures, 3 in Color, and 16 Tables
Springer
Dr. Marcel G.A. van der Heijden
Free University of Amsterdam
Department of Systems Ecology
Faculty of Earth and Life Sciences
De Boelelaan 1087
1081 HV Amsterdam
The Netherlands
Dr. Ian R. Sanders
University of Lausanne
Institute of Ecology
Biology Building
1015 Lausanne
Switzerland
Cover illustration: Marcel G.A. van der Heijden
ISSN 0070-8356
ISBN 978-3-540-00204-8 ISBN 978-3-540-38364-2 (eBook)
DOI 10.1007/978-3-540-38364-2
Originally published as hardcover edition in 2002 ISBN 978-3-540-00204-8
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Preface
Plants collaborate with many micro-organisms in the rhizosphere to form
mutualistic associations. One of the best examples is the mycorrhizal symbio
sis between plants and fungi. Here, fungi support plants with mineral nutri
ents and other services and the fungi, in turn, receive photosynthates from the
autotrophic plants. Mycorrhizal associations are common in almost all eco
systems and 80 % of all land plants associate with these mutualistic soil fungi.
There is an increasing awareness among biologists, ecologists and mycolo
gists that mycorrhizal associations need to be considered in order to under
stand the ecology and evolution of plants, plant communities and ecosystems.
In the last decade, many advances and breakthroughs have been made in
mycorrhizal ecology. We aim to summarise these advances in this Volume,
with special emphasis given to the ecological function of the mycorrhizal
symbiosis.
This Volume is divided into six sections. The first section gives an intro
duction to the mycorrhizal symbiosis and discusses the progress that has
been made in understanding the ecological function of this association. The
second section deals with the eco-physiology of mycorrhizal plants. It also
covers the influence of global changes on the symbiosis. The third section dis
cusses the influences of mycorrhizal fungi on biodiversity and ecosystem
functioning. It also discusses factors that influence the diversity and structure
of mycorrhizal fungal communities. The fourth section shows the multi
trophic nature of the mycorrhizal symbiosis. It shows that mycorrhizal fungi
interact with individuals and communities of different trophic levels includ
ing plants, herbivores, insects and other soil fungi. The fifth section takes a
more evolutionary approach to looking at host specificity between plants and
mycorrhizal fungi. Finally, the last section gives a summary of this Volume.
We aim to bring together these major topics in mycorrhizal ecology and
this also reflects our personal interest in integrating the study of the mycor
rhizal symbiosis within the much wider field of plant ecology. Moreover, the
field of mycorrhizal ecology has expanded greatly in recent years. It was,
therefore, not possible to include all disciplines and choices had to be made.
In addition, during the preparation of this Volume, several exciting findings in
our field were published. Several authors have included these findings in their
VI Preface
contributions but of course not all could be covered. We aimed to cover these
last-minute breakthroughs in Chapte 17, "Mycorrhizal Ecology: Synthesis and
Perspectives" .
The editors are thankful to many persons who contributed to this Volume.
First, we would like to thank Dieter Czechlik and Andrea Schlitzberger from
Springer Verlag for their invitation to start this Volume. We would also like to
thank Prof. 0.1. Lange, the Series Editor of Ecological Studies. I am especially
thankful to Rien Aerts, Head of the Department of Systems Ecology at the
Vrije Universiteit Amsterdam, for his continuous support and for giving me
the opportunity to accomplish this exciting project (MvdH). Many thanks also
to David Read for his help, suggestions and contribution to this Volume. Each
of the chapters was peer-reviewed by two independent referees. We are, there
fore, grateful to all the people who reviewed the chapters. Last, but not least,
MvdH would like to thank his closest friends and relatives for their encour
agement, support and interest in his work. IS thanks the Swiss National Sci
ence Foundation for financial support of a professorial fellowship during the
period in which this Volume was edited.
December 2001 Marcel G.A. van der Heijden,
Amsterdam, The Netherlands
Ian R. Sanders,
Lausanne, Switzerland
Foreword: Mycorrhizae and "Joined-Up" Ecology
In the second half of the 20th century, rapid progress was achieved in many
fields of biological research by the development of specialist approaches.
Ecology was no exception to this trend and much has been achieved by the
concentrated attack that is possible when groups of scientists co-operate to
address well-defined objectives.
As the end of the last century approached, it became apparent that to serve
the best interests of ecological research, this reductionist agenda would
require more sophisticated development than was necessary in neighbouring
biological sciences such as genetics, molecular biology and biomedical sci
ence. In particular, it became necessary to remind ourselves that the ultimate
objectives of ecology are to understand the assembly and dynamic properties
of communities, ecosystems, landscapes and biomass. To achieve this "scal
ing-up", I propose that the most significant challenge for ecologists at the pre
sent time is to continue to exploit the many current opportunities for pene
tration into detailed mechanisms but simultaneously to actively promote
integration of sub-disciplines within ecological research. Mycorrhizal ecology
is uniquely positioned and qualified to provide a lead in this difficult task.
Quite clearly, as the expert authors of the various chapters in this Volume
explain, their subject has been an interdisciplinary pursuit throughout its his
tory with important and continuing involvement of plant physiologists,
mycologists, soil scientists, animal ecologists and, more recently, molecular
biologists and geneticists. Equally important, since its inception, mycorrhizal
ecology has recognised the diversity of phenomena represented by the vari
ous associations between mycorrhizal fungi and vascular plants and the very
different consequences of this diversity for the survival and functioning of
plant communities and ecosystems in contrasted environments.
Ever since my first encounter with Jack Harley, I have been aware that a
motivating force for mycorrhizal ecologists and their subterranean allies has
been the corporate and deeply held conviction that research below ground
has been a neglected and undervalued part of ecology. The exciting achieve
ments and expanding perspectives reviewed in this Volume are a vindication
of the long campaign to direct sharp intelligence and adequate funding to the
many unsolved problems that lie below the soil surface.
VIII Foreword: Mycorrhizae and "Joined-Up" Ecology
With the case for the importance of mycorrhizal ecology now firmly estab
lished in terrestrial studies, what will be the future role of research on this
topic? This Volume provides several pointers to particular developments in
both fundamental and applied ecology. In addition to these, however, I would
not be surprised to see a more general effect on the conduct of ecological
research and the development of ecological theory. For the purposes of man
agement and conservation, it has become essential to develop an array of
dynamic models that are driven by the biology of component organisms and
capture the functioning of ecosystems of widely contrasting types. As ecolo
gists of many persuasions converge upon the ecosystem as a subject for mod
elling and as a vehicle for advice to policy makers, the pivotal position of
dominant plant species and their mycorrhizal associates, as mediators of
ecosystem productivity and biogeochemistry, is likely to be widely recog
nised. I am confident that in such circumstances the continuation of a
research tradition embracing in situ observation, experimental manipulation,
quantitative biochemistry and mechanistic interpretation will assure an
influential role for mycorrhizal experts of the future.
1.P. Grime
Contents
Section A: Introduction
1 Towards Ecological Relevance -
Progress and Pitfalls in the Path Towards
an Understanding of Mycorrhizal Functions in Nature 3
D.J.READ
1.1 Summary . . . . . . . . . . . . . . . . . . . . . . . . 3
1.2 Introduction . . . . . . . . . . . . . . . . . . . . . . 4
1.3 The Platform Provided by Reductionist Approaches 4
1.4 Approaches to Examination of Mycorrhizal Function
at the Community Level 8
1.4.1 The Microcosm Approach 10
1.4.2 The Field Approach 21
1.5 Conclusion 23
References . . . . . . . . . . . 24
Section B: Ecophysiology, Ecosystems Effects and Global Change
2 Carbon and Nutrient Fluxes Within
and Between Mycorrhizal Plants 33
S.W. SIMARD, D. DURALL, M. JONES
2.1 Summary ............... 34
.............
2.2 Introduction 34
2.3 Ecto-mycorrhiza and Nutrient Uptake 35
2.3.1 Quantitative Effects of Ecto-mycorrhiza
on Nutrient Uptake by Plants . . . . . . ....... 35
......
2.3.2 Mechanisms of Increased Nutrient Uptake 39
2.3.3 Transport of Nutrients Within the Ecto-mycorrhizal
Fungal Mycelium ............. 43
2.4 Carbon Flux in Ecto-mycorrhizal Plants 44
