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Nine years have elapsed since the publication of the second edition and we app-
roached the preparation of this new edition with much trepidation. First, there has
been a vast expansion of information and, secondly, much of this information has
been derived from the application of new techniques of molecular biology, which had
barely commenced at the time of preparation of the second edition. Now the task of
writing is completed to the best of our abilities we can only agree with the statement
made in 1983 (first edition) that it is 'almost impossible for one person or even two
to keep up with all the experimental work and speculation on the subject of mycor-
rhizas'! We have done our best, but apologize to those readers who feel that our treat-
ments of some areas lack sufficient coverage.
In this edition, we have retained part of the text from the second edition but, again,
there has been so much new work that detailed reference to some research included
in the first two editions has had to be reduced. We have, however, attempted to
retain a feeling for the way the subject has developed and to highlight major con-
tributions of early researchers and we urge current and future researchers to delve
into early (pre-electronic) literature because it contains many ideas and much valu-
able information which should not be wasted or describes experiments that need not
be repeated.
Again, it has been impossible to review all topics in detail and this third edition
once more reflects our interests and provides personal views of the subject. Where
the first edition explicitly avoided evolutionary discussions, on the grounds that
insufficient information was available to make this profitable, we have capitalized
on new information based on molecular phylogeny of fungi and of plants (intro-
duced in the second edition), to reveal likely evolutionary pathways by which
mycorrhizas have arisen. We have also maintained the emphasis on the extraradi-
cal mycelium, both with respect to development in all types of mycorrhizas and to
function. In most areas of research, the structural and functional diversity among
mycorrhizas has continued to be revealed. Accordingly, we highlight this aspect of
mycorrhizal biology and show how it may be important in ecological situations.
The second edition was written when the very first results of molecular bio-
logical investigation of mycorrhizas were being published. In the intervening time
there has been an explosion of such research, both targeted to increasing our under-
standing of well-known features of symbiosis, such as nutrient transfer, as well as
to revealing 'unknown' aspects of the symbioses. The roles of mycorrhizas as the
normal nutrient absorbing organs of the vast majority of plant species in nature is
increasingly being revealed and appreciated by ecologists. Throughout the book we
have emphasized the experimental approaches that have proved useful and that
may need to be applied in the future.
viii Preface

We have retained the same general structure of the book, with three sections pro-
viding general accounts of the main types of mycorrhiza, including information
on the identity of the symbionts, structure and development of the mycorrhizas
formed by them, as well as their function and ecological significance. The fourth
section is devoted to the functioning of mycorrhizas in broader contexts, in which
we believe integration of ideas and information is essential for clear understanding.
These chapters build on the material presented earlier, but so that they can be read
separately we have deliberately included some repetition and have also tried to
improve the cross-referencing in the text. We concentrate on the presence and activ-
ity of mycorrhizal symbioses in the major global biomes, on their roles in ecological
interactions and on applications related to agriculture, horticulture and forestry. The
essays in the first edition have served their purpose. Information on general aspects
of nutrient transfers (second edition) has now been integrated as far as possible into
chapters on the different mycorrhizal types. Readers interested in these topics will
need to use the earlier editions.
Throughout the text we have tried to indicate where future experimental research
might be directed, although we realize that our personal views may have intro-
duced some bias, which will be recognized by those who know us.
Many friends and colleagues have helped us in countless ways, especially by dis-
cussions, by reading drafts of chapters and by supplying illustrations and results
(some unpublished). We have not repeated the list from the second edition, but list
alphabetically all those to whom we are particularly indebted in the preparation of
this third edition: Susan Barker, Martin Bidartondo, Tom Bruns, Duncan Cameron,
Tim Cavagnaro, Michel Chalot, Mark Chase, Sandy Dickson, Jeff Duckett, Evelina
Facelli, Jose Facelli, Andrea Genre, Manuela Giovannetti, Emily Grace, Lisa Grubisha,
Sarah Hambleton, Maria Harrison, Jan Jansa, Jonathan Leake, Francis Martin, Maria
Manjarrez, Randy Molina, Hugues Massicotte, Joe Morton, Jesus Perez-Moreno,
Larry Peterson, Arthur Sch(if~ler, Andrew Smith, Lee Taylor, Sari Timonen, Jean-
Patrick Toussaint, Andrew Smith, Michael Weiss and Katya Zimmer. We are grate-
ful to them all for their critical and helpful comments, many of which have been
incorporated into the text. However, we must take responsibility for all the ideas and
views expressed whether good or bad; the errors are (once again) all ours.
We have also received invaluable help in preparation of the manuscript. Jayne
Young typed many drafts of the chapters and produced the final manuscript;
Rebecca Upson and Chris Read provided us with much-needed support in obtain-
ing copyright clearances and cross-checking information and Glyn Woods provided
assistance with production of many images.
We have used much previously published material in both tables and figures. The
sources are acknowledged in each case, but we here wish to give special thanks to
all the authors and publishers who generously allowed us to use their copyright
Last, but by no means least, our thanks and love go to the 'long-sufferers',
Andrew Smith and Chris Read, who provided invaluable moral support which
helped to see us through the many months of largely self-centred devotion to writ-
ing. They have had to go through this for a second time, so we are doubly grateful.
The book is again dedicated to the memory of Jack Harley, whose influence upon
the development of our subject was so enormous. To expand the list of those who
have made highly significant contributions to mycorrhizal research would have
Preface ix

opened a Pandora's box. Who should we choose and what would happen if we
inadvertently omitted crucial names? We hope that our attempts at historical treat-
ment will make abundantly clear where credit for particular advances should be
Among Jack Harley's many contributions, two stand out as being of pre-eminent
importance. Frustrated in his earlier career as an ecologist by much woolly think-
ing about the biology of mycorrhiza, Jack determined to subject the topic to rigorous
physiological analysis. Over several decades and using as his research material the
ectomycorrhizal roots of beech, he and his collaborators evaluated the basic processes
whereby nutrients are exchanged between partners in the symbiosis. The second and
arguably more important contribution arose out of his skill as a communicator. The
Biology of Mycorrhiza, published in 1959, was the first ever attempt to synthesize the
many and sometimes disparate strands of thought which had developed in over 100
years of research. With characteristic incisiveness he cut through much, often pedan-
tic, debate to focus upon those questions which were in need of resolution. The work
was of inestimable importance to many who, like ourselves, were struggling to take
their first steps in research on this subject. These combined contributions provided
impetus to a further expansion of research to the extent that, by 1983, a new and even
more substantial volume, the first edition of Mycorrhizal Symbiosis, was required. One
of us had the privilege to collaborate in that enterprise.
The influence of Jack Harley goes on. Although, sadly, he has not been here to
assist us in updating the book through two editions, we have both been conscious
of his legacy. Without him it is doubtful whether the subject would be in the pre-
eminent position it enjoys today, increasingly recognized by physiologists and
ecologists and now molecular biologists as being of central importance in plant and
fungal biology.