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Diversity and Integration in Mycorrhizas: Proceedings of the 3rd International Conference on Mycorrhizas (ICOM3) Adelaide, Australia, 8–13 July 2001 PDF

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DIVERSITY AND INTEGRATION IN MYCORRHIZAS Diversity and Integration in Mycorrhizas Proceedings of the 3rd International Conference on Mycorrhizas (ICOM3) Adelaide, Australia, 8-13 July 2001 Edited by S.E. SMITH and F.A. SMITH Reprinted from Plant and Soil, Volume 244. Nos. 1-2 (2002). SPRINGER-SCIENCE+BUSINESS MEDIA, B.V. Library of Congress Cataloging-in-Publication Data A C.l.P. Catalogue record for this book is available from the library of Congress. ISBN 978-90-481-5933-8 ISBN 978-94-017-1284-2 (eBook) DOI 10.1007/978-94-017-1284-2 Printed on acid-free paper AII Rights reserved © 2002 Springer Science+Business Media Dordrecht Originally published by Kluwer Academic Publishers in 2002 N o part of the material protected by this copyright notice may be reproduced ar utilized in any form or by any means, electronic or mechanical, including photocopying, recording or by any infarmation storage and retrieval system, without written permission from the copyright owner. Contents DIVERSITY AND INTEGRATION IN MYCORRHIZAS Proceedings of the 3rd International Conference on Mycorrhizas (ICOM3) Adelaide, Australia, 8-13 July 2001 Foreword ix Introductory Chapter I. Challenges for mycorrhizal research into the new millennium B. Soderstrom Molecular Diversity and Phylogeny 2. Molecular and functional diversity in soil micro-organisms J .I. Prosser 9 3. Fungal diversity in ectomycorrhizal communities: sampling effort and species detection A.F.S. Taylor 19 4. Molecular approaches to ectomycorrhizal diversity studies: variation in ITS at a local scale T.R. Horton 29 5. Ericoid mycorrhizal fungi: some new perspectives on old acquaintances S. Perotto, M. Girlanda & E. Martino 41 6. Molecular detection, community structure and phylogeny of ericoid mycorrhizal fungi S.M. Berch, T.R. Allen, & M.L. Berbee 55 7. Molecular identification and phylogeny of arbuscular mycorrhizal fungi D. Redecker 67 8. Molecular phylogeny, taxonomy, and evolution of Geosiphon pyriformis and arbuscular mycor rhizal fungi A. SchtiBler 75 Development: from Genes to Structure 9. The application of genetic approaches for investigations of mycorrhizal symbioses S.J. Barker, S. Duplessis, D. Tagu 85 10. The ectomycorrhizal symbiosis: Genetics and development D. Tagu, F. Lapeyrie & F. Martin 97 II. Molecular approaches to understanding mycorrhizal symbioses S.J. Barker & N.J. Larkan 107 12. Fungal gene expression in early symbiotic interactions between Laccaria bicolor and red pine G.K. Podila, J. Zheng, S. Balasubramanian, S. Sundaram, S. Hiremath, J.H. Brand & M.J. Hymes 117 13. Early developmentally regulated genes in the arbuscular mycorrhizal fungus Glomus mosseae: identification of GmGINl, a novel gene with homology to the C-terminus of metazoan hedgehog proteins N. Requena, P. Mann, R. Hampp & P. Franken 129 14. Expression in an arbuscular mycorrhizal fungus of genes putatively involved in metabolism, transport, the cytoskeleton and the cell cycle J. Jun, J. Abubaker, C. Rehrer, P.E. Pfeffer, Y. Shachar-Hill & P.J. Lammers 141 15. Recent developments in the study of orchid mycorrhiza H.N. Rasmussen 149 Functional Relationships 16. An update on nutrient transport processes in ectomycorrhizas M. Chalot, A. Javelle, D. Blaudez, R. Lambilliote, R. Cooke, H. Sentenac, D. Wipf & B. Botton 165 17. The role of the motile tubular vacuole system in mycorrhizal fungi A.E. Ashford & W.G. Allaway 177 18. Tracking metabolism and imaging transport in arbuscular mycorrhizal fungi B. Bago, P.E. Pfeffer, W. Zipfel, P. Lammers & Y. Shachar-Hill 189 19. Cytoskeleton in mycorrhizal symbiosis S. Timonen & R.L. Peterson 199 20. Endosymbiotic bacteria in mycorrhizal fungi: from their morphology to genomic sequences D. Minerdi, V. Bianciotto & P. Bonfante 211 21. P metabolism and transport in AM fungi T. Ezawa, S.E. Smith & F.A. Smith 221 22. Mechanisms of nutrient transport across interfaces in arbuscu1ar mycorrhizas N. Ferro!, J.M. Barea & C. Azc6n-Aguilar 231 23. Phosphate transporters of Medicago truncatula and arbuscular mycorrhizal fungi W.K. Versaw, T.-J. Chiou & M.J. Harrison 239 24. Plant nutrient transporter regulation in arbuscular mycorrhizas S.H. Burleigh & I.E. Bechmann 247 Ecological Interactions 25. Mycorrhizas and global environmental change: research at different scales P.L. Staddon, A. Heinemeyer & A.H. Fitter 253 26. Is there a role for arbuscular mycorrhizal fungi in production agriculture? M.H. Ryan & J.H. Graham 263 27. Inoculation with arbuscular mycorrhizal fungi: the status quo in Japan and the future prospects M. Saito & T. Marumoto 273 28. Host-specificity of AM fungal population growth rates can generate feedback on plant growth J.D. Bever 281 29. Ecological role of hypogeous ectomycorrhizal fungi in Australian forests and woodlands A.W. Claridge 291 30. Effects of mycorrhizal fungi on plant populations R.T. Koide & LA. Dickie 307 31. The role of mycorrhizas in plant community structure and dynamics: lessons from grasslands D.C. Hartnett, G.W.T. Wilson 319 Plant and Soil 244: ix-x, 2002. IX © 2002 Kluwer Academic Publishers. Foreword Proceedings of ICOM3: Diversity and Integration in Mycorrhizas This book contains chapters that were derived from invited Keynote and Symposium presentations at the 3rd International Conference on Mycorrhizas (ICOM3), held in Adelaide, Australia, from 8-13 July 200 I. However, it is more than 'a book of the conference': it also contains chapters by authors who did not speak at the conference, but who were invited in order to provide additional depth and balance. We are grateful to all for the efforts that they have put into their contributions. Not all of those invited to write chapters were able to do so, but we hope that the book provides a flavour both of ICOM3 itself and of current research on mycorrhizal symbioses-a field that is rapidly developing. All chapters were reviewed independently, following the normal procedures of Plant and Sui/. About 450 delegates from 50 countries attended ICOM3 and Abstracts of all presentations-talks and posters-are available on the ICOM3 Website: http://www. waite.adelaide.edu.au/Suil_ Water/3icom.html As organizers of ICOM3, it is not for us to judge its success or impact. We are pleased that informal feedback and published reports have been very positive ('Mycorrhizal Research on the Move', Burleigh and Jakobsen, 2002 and 'ICOM3: A Landmark in Mycorrhiza Research!', Colpaert, 2002). The conference theme was 'Diversity and Integration in Mycorrhizas' and, to ensure that this theme could be appreciated by all participants, our approach was to bring together major topics in mycorrhizal research that crossed some of the traditional boundaries often reflected in mycorrhizal conferences, such as divisions between different classes of mycorrhizas or divisions based on techniques. We also ensured that a number of up-and-coming younger researchers were invited to present their work, particularly since, in the main, they are the people who are applying novel techniques to unravelling details of symbiotic processes. This approach was evidently well appreciated, as emphasized by Professor David Read in his closing overview of ICOM3. The themes of the conference are reflected in the way in which chapters are grouped in this book. After the scene is set by Bengt Soderstrom in his thoughtful introductory chapter 'Challenges for mycorrhizal research into the new millennium', the chapters are presented in four groups: 'Molecular diversity and phylogeny', 'Development: from genes to structure', 'Functional relationships' and 'Ecological interactions'. The arrangement of chapters in the book is not exactly the same as that of the presentations at the conference. Some of the chapters lie at the boundaries between themes and others are wide-ranging and cover more than one theme; there is also a connecting loop because of the way molecular tools resulting from studies of 'Molecular diversity and phylogeny' are becoming increasingly important in unravelling 'Ecological interactions'. We hope that in the new arrangement no papers seem entirely out of place. No classification can ever be perfect and- as with mycorrhizal fungal taxonomists - we have done our best. We are conscious of topics that were covered in concurrent sessions at the meeting and are not included in the book, such as those on 'Problem soils and extreme environments', 'Mycorrhizas in the tropics', the 'Mycor rhizosphere', and 'Mycorrhizas in vegetation restoration', as well as the informal workshops on a wide range of specialist topics. We mention those here to emphasize the wide-ranging influence of mycorrhizas on plant-soil interfaces, and the fact that even an international meeting such as ICOM cannot emphasise diversity without be coming much larger, in which case it would lose the integration that is obviously highly appreciated by participants. The integration is also one of the keys to grasping the complexities of the roles of mycorrhizas in ecosystems. In conclusion, ICOM3 (and hence this book) would not have been possible without generous financial support from our major sponsors: Kluwer Academic Publishers/Plant and Soil, Samuel Roberts Noble Foundation, New Phytologist, Canadian Journal of Botany, Forest & Wood Products Research and Development Corporation, Grains Research and Development Corporation, the Australian Society of Plant Physiologists (now the Australian Society of Plant Scientists), and the University of Adelaide's Centre for Plant Root Symbioses. We also received financial support from other sources within the University of Adelaide (the Vice-Chancellor's Fund, the Faculty of Science and the Department of Soil and Water) and from AusAID, Biorize, the Cooperative Research Centre for Molecular Plant Breeding, Premiertech, Springer-Verlag, the Australian Academy of Science, the Australian Institute of X Biology and last but not least Mrs Lindsay Harley. We thank the team at Kluwer who have played a major role in bringing the publication to completion and in good time, including Arno Flier for his strong encouragement to take on the project, Pieter Poot as managing editor of the volume, Peter-Paul Delsman as desk editor and particularly Otilia Prins who has been at the interface between authors and editors. We cannot thank individually all those who helped arrange ICOM3: the members of the Organizing Committee and the Australasian Advisory Committee, the organizers of Symposia, Workshops and Concurrent sessions, and many others. Nevertheless we will once again gladly thank the conference organizer Sally Jay, her co-organizer Jenny Blanchard, and Mark Stevens the audiovisual coordinator, for everything that they did to bring ICOM3 into being, and the members and associates of the Mycorrhiza Group in the Department of Soil and Water, who helped us in many ways. Sally Smith (SE Smith) Andrew Smith (FA Smith) Department of Soil and Water and the Centre for Plant Root Symbioses, Waite Campus, The University of Adelaide, PMB 1, Glen Osmond 5064 South Australia, AUSTRALIA References Burleigh. Sand Jakobsen, I 2002. Mycorrhizas on the move. Trends in Plant Science, 7. &-7. Colpaert, J 2002. ICOM3: A Landmark in Mycorrhiza Research! Mycorrhiza, 12. 5!. Plant and Soil 244: 1-7, 2002. © 2002 Kluwer Academic Publishers. Challenges for mycorrhizal research into the new millennium Bengt Soderstrom Department o{Microbial Ecology, Ecology Building, Lund University, SE-223 62 Lund, Sweden Key words: carbon dioxide, carbon flow, climate change, genomic biology, mycorrhiza Abstract Some future directions for mycorrhizal research are discussed. Data from The IPCC Third Assessment Reports 2001 on elevated carbon dioxide the concentration and consequences regarding global climate change, form the basis for arguments that a better understanding of the mycorrhizal activities with regard to controlling carbon flow and plant nutrient flow is essentiaL Mycorrhiza can be regarded as energy and nutrient switches in the plant roots and it is suggested that more studies on mycorrhizal carbon relations are necessary. The impact of the genomic research era on mycorrhizal research is suggested to be of major importance during the years to come, and it is claimed that stronger interactions between different disciplines will be essentiaL Introduction corrhizal research even further. This paper exemplifies some areas of research which have the potential for The importance of mycorrhizal symbioses in nature exciting and important development, and some of the cannot be overestimated. The significance of these prerequisites for such development are discussed. interactions in the nutrition and well-being of the in dividual plant partner is well established (Smith and Read, 1997) and recent evidence also indicates that The mycorrhizal research community they may also have major effects on the structure of the plant community (van der Heijden et a!., 1998). Mycorrhizal researchers have traditionally been re Their ubiquitous existence and impact on plant and cruited from a number of established disciplines such soil biology make it necessary to consider them in all as botany, microbiology or ecology. This is not sur terrestrial ecological research. Since this form of sym prising when one considers that mycorrhizal sym biosis was first described in the mid 19th century, the biosis in itself is a very integral phenomenon. For fundamental questions surrounding these intriguing the advancement of our knowledge and understanding biological systems have attracted a large number of of these complex systems, the international meet researchers, as has the question regarding potential ings dedicated to the study of mycorrhizas have un applications in agriculture and forestry to enhance doubtedly been of great importance. During the past plant production and sustainability. Most work has 20 years, these meetings have grown considerably. been carried out on the ectomycorrhizal and arbuscu Based on the number of abstracts submitted to the in lar mycorrhizal systems, a fact which is not surprising ternational conferences with mycorrhiza as the main considering their dominance in nature. theme, it seems that the number of researchers in It has been predicted that the 21st century will be volved in mycorrhizal investigations has increased come the century of biology as the 20th century was four times the past 20 years. The North American Con the century of physics, the so called 'new biology' ferences on Mycorrhiza (NACOM) had a strong inter being the justification for such a prediction. Bearing in national participation, and in 1979 the 4th NACOM mind the critical importance of the mycorrhizal sym attracted I 06 abstracts. Since International Confer bioses in nature and their strong influence on many ences of Mycorrhizas have been organized (!COM, ecological processes, mycorrhizal researchers are in a 1996, 1998, 2001 ), they have attracted between 400 very favourable position to develop the field of my- and 500 contributions and the NACOMs and the

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This book is highly recommended on the basis of the following points: - The editors are highly regarded in the field of mycorrhizal biology and one is co-author of the most comprehensive textbook on mycorrhizas; - Chapters by international experts based on invited presentations at the 3rd Internatio
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