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Trace Elements in the Rhizosphere 1535/frame/FM Page 2 Tuesday, August 22, 2000 9:08 AM Trace Elements in the Rhizosphere Edited by George R. Gobran Walter W. Wenzel Enzo Lombi CRC Press Boca Raton London New York Washington, D.C. Library of Congress Cataloging-in-Publication Data Trace elements in the rhizosphere / George R. Gobran, Enzo Lombi & Walter W. Wenzel, editors. p. cm. Papers from a special symposium held during the Fifth International Conference on Biogeochemistry of Trace Elements, July 11–15, 1999 in Vienna, Austria. Includes bibliographical references and index. ISBN 0-8493-1535-2 1. Soils—Trace element content—Congresses. 2. Trace element in plant nutrition—Congresses. 3. Plant-soil relationships—Congresses. 4. Rhizosphere—Congresses. I. Gobran, George R. II. Lombi, Enzo, 1968– III. Wenzel, Walter W. IV. International Conference on the Biogeochemistry of Trace Elements (5th : 1999 : Vienna, Austria) S592.6.T7 T73 2000 631.4′16—dc21 00-058577 CIP This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials or for the consequences of their use. 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 or retrieval system, without prior permission in writing from the publisher. All rights reserved. Authorization to photocopy items for internal or personal use, or the personal or internal use of specific clients, may be granted by CRC Press LLC, provided that $.50 per page photocopied is paid directly to Copyright Clearance Center, 222 Rosewood Drive, Danvers, MA 01923 USA. The fee code for users of the Transactional Reporting Service is ISBN 0-8493-1535-2/01/$0.00+$.50. The fee is subject to change without notice. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. The consent of CRC Press LLC does not extend to copying for general distribution, for promotion, for creating new works, or for resale. Specific permission must be obtained in writing from CRC Press LLC for such copying. Direct all inquiries to CRC Press LLC, 2000 N.W. Corporate Blvd., Boca Raton, Florida 33431. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation, without intent to infringe. © 2001 by CRC Press LLC No claim to original U.S. Government works International Standard Book Number 0-8493-1535-2 Library of Congress Card Number 00-058577 Printed in the United States of America 1 2 3 4 5 6 7 8 9 0 Printed on acid-free paper Preface The term rhizosphere was introduced by Hiltner in 1904. Since then, the “hidden half of the hidden half” has been investigated in its many aspects. Initially, the interest of scientists and practitioners was directed toward plant nutrition, in partic- ular, as to whether rhizosphere processes influence the uptake of mineral nutrients by crop and tree species. With the general move of soil and plant scientists from agricultural to ecological/environmental issues, the emphasis of rhizosphere research has also increasingly shifted to studying the effect of plant–microbial associations on the bioavailability, uptake, and transformation of inorganic and organic contam- inants in soils. Most recently, contaminant–rhizosphere interactions have attracted renewed attention as green plants have been proposed as being useful in the reme- diation of contaminated soils (phytoremediation). While potential practical applications are a driving force in rhizosphere research, it is equally important to develop methodological tools employed to elucidate the fundamental physical, chemical, and biological processes that create the highly dynamic and reactive microenvironment around plant roots. Basically, two approaches have been taken in the study of the rhizosphere: experimental investi- gation and modeling. While a relatively large number of publications dealing with various aspects of the rhizosphere is available, the information is still scattered and we are far from a comprehensive understanding of the manifold interactions and feedbacks that occur in the rhizosphere. In particular, only a few examples of studies that link modeling to experimental work (model validation) are available, and little is available on the fate of trace elements in the rhizosphere, even though such information is urgently needed to improve food quality and to develop phytoreme- diation technologies. Needless to say, the rhizosphere plays a key role in controlling trace element contamination of the terrestrial food chain. Therefore, it is one of the hot spots in sustaining ecosystem health. Interest in this topic led to the organization of a special symposium entitled, Fate of Trace Elements in the Rhizosphere, held during the Fifth International Conference on the Biogeochemistry of Trace Elements, July 11–15, 1999, in Vienna, Austria. This special symposium provided a forum to bring together scientists involved in various aspects of rhizosphere research. Most of the chapters in this book comprise work that was presented by the outstanding scientists invited to this special symposium. The book Trace Elements in the Rhizosphere is divided in three sections. Section I (Chapters 1 to 5) emphasizes the bioavailability of trace elements, including radionuclides in the rhizosphere and their significance in developing management strategies for phytoremediation. Section II (Chapters 6 to 8) deals with the complex interactions of root–microbial associations, in particular mycorrhizae, and the trans- formation of trace elements. Tools and methods available for study of the dynamics and fate of trace elements in the rhizosphere are discussed in Section III (Chapters 9 to 12), presenting examples of experimental techniques and modeling approaches. Chapter 1 presents a review of the role that rhizosphere processes play in the mobilization or immobilization of trace elements and summarizes the state-of-the- art knowledge on metal hyperaccumulators as compared to nonaccumulating plant species. Special reference is given to the application of rhizosphere processes to phytoremediation, and the potential of manipulating the rhizosphere to control trace element availability for phytoremediation of crops is explored. Chapter 2 provides a review of complementary in-depth information on trace metal bioavailability in the rhizosphere. Chapter 3 contributes to an understanding of the specific conditions that modify trace metal interactions in the rhizosphere. Chapter 4 extends the scope to radiocesium. Chapter 5 presents a comprehensive review of the fate of U and Pu radionuclides in soil–plant systems. The role of rhizospheric interactions and their application to the phytoremediation of U- and Pu-polluted soil and water environ- ments are emphasized. Chapter 6 provides current information on exudates and metal-organic complexes associated with root–microbial associations. Chapter 7 emphasizes the importance of metal speciation in symbiotic fungi (ectomycorrhiza), presenting data generated by micro X-ray absorption spectroscopy, a promising technique that might be more widely applied in rhizosphere research. Chapter 8 discusses bioavailability of trace elements in mycorrhizal rhizosphere systems, hence, linking the information provided in Sections I and II. Chapter 9 explores the application of a sequential chemical extraction technique as a means to study trace metal fractionation in rhizosphere soils. Chapter 10 presents a technique to study the effect of micronutrients or toxic trace elements on plant growth, using a steady- state approach. This methodology may prove useful in rhizosphere studies, since experimental conditions of microelement nutrition and their interactions with plant physiological processes can modify root exudation and related processes in the rhizosphere. Chapter 11 presents a model that describes cation exchange on plant roots, a process that has been neglected in most models so far. Chapter 12 describes a model of aluminum chemistry with reference to acidic forest soils. It is hoped that this book will contribute to the body of knowledge on the secrets of the rhizosphere. The book is suitable for advanced undergraduate and graduate courses in soil environmental chemistry, biogeochemistry, and ecology. It should interest professionals in those areas, as well as consulting engineers and decision makers involved in bioremediation technology and sustaining ecosystem health. We greatly acknowledge the authors of this book and their cooperation and support. We also wish to extend our special thanks to many anonymous reviewers of each chapter, especially G. Ågren and R. Finlay (Swedish University of Agricul- tural Sciences, SLU, Sweden), and K. Turnau (Jagiellonian University, Krakow, Poland). The book’s manuscript was carefully read by M.B. Kirkham (Kansas State University, U.S.A.) and M.P. Huang (University of Saskatchewan, Canada), who deserve our special thanks and appreciation. We are also grateful for the financial support received from the Faculty of Forestry, SLU, Sweden, and the Swedish Council for Forestry (SJFR); the Agricultural Research, IACR-Rothamsted grant- aided support received from the Biotechnology and Biological Sciences Research Council of the United Kingdom; and the support of the University of Agricultural Sciences, Vienna, Fonds zur Förderung der Wissenschaftlichen Forschung, Austrian Federal Ministry of Science and Transport. April 2000 George R. Gobran, Walter W. Wenzel, and Enzo Lombi Editors George R. Gobran is Professor of Ecology-Soil Sciences with specialization in nutrient dynamics in the rhizosphere. Since 1985, Dr. Gobran has been working at the Department of Ecology and Environmental Research, Swedish University of Agriculture Sciences, Uppsala, Sweden. Dr. Gobran received his Ph.D. in soil chem- istry from the Catholic University of Louvain-la-Neuve, Belgium in 1980. He received his M.S. in soil chemistry in 1975 and his B.S. in soil and water sciences in 1969 from Alexandria University, Egypt. Dr. Gobran was also awarded a post- doctoral fellowship from Texas A & M University, U.S.A. (1982–1984). Professor Gobran has significant experience in research dealing with soil chem- ical processes, with special interest in soil–plant interactions and rhizospheric pro- cesses. Currently, he focuses his research and teaching efforts on the reciprocal effects of soil–plant interactions, especially in ecosystems under environmental stress. Dr. Gobran has written many papers and book chapters, and has participated in several international conferences, workshops, and symposia. Additionally, he is a reviewer for several international journals and programs and is included in the 1999 edition of Who’s Who in the World. With his coeditors, Drs. Wenzel and Lombi, Dr. Gobran organized a symposium entitled, Fate of Trace Elements in the Rhizo- sphere, which was presented during the Fifth International Conference on the Bio- geochemistry of Trace Elements, Vienna, July 11–15, 1999. Walter W. Wenzel is Professor of Soil Science at the University of Agricultural Sciences in Vienna, Austria. He has been working on various fundamental aspects of trace element biogeochemistry, with emphasis on several cationic metals and the metalloids As and F. Currently, Professor Wenzel’s work is centered around exper- imental assessment and modeling of rhizosphere processes and metal hyperaccumu- lation. His involvement in applied research has been largely related to risk assessment and the development of phytoremediation technologies for the cleanup of soils contaminated with metals and organic pollutants. Professor Wenzel is coeditor of the Journal of Environmental Quality and the Journal of Plant Nutrition and Soil Science. He organized the Fifth International Conference on the Biogeochemistry of Trace Elements, Vienna, July 11–15, 1999. Additionally Professor Wenzel is one of the initiators and founders of the International Society for Trace Element Bio- geochemistry, for which he currently serves on the board of directors. Enzo Lombi was born in Piacenza, Italy, 1968. After graduation in 1993, he earned a Ph.D. in agricultural chemistry and conducted his research in Italy, Austria, and the United States. During 1997–1998 he was Scientific Researcher at the Institute of Soil Science, University of Agricultural Sciences, Vienna, and at the Austrian Research Centre, Seibersdorf. His activities focused on the development and appli-

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