CO2 0Prels 2/8/01 3:10 pm Page i IUFRO Research Series The International Union of Forestry Research Organizations (IUFRO), with its 14,000 scientistsfrom 700 member institutions in 100 countries, is organized into nearly 300 research units that hold approximately 60 conferences, work- shops and other meetings annually. The individual papers, proceedings and other material arising from these units and meetings are often published, but in a wide array of different journals and other publications. The object of the IUFRO Research Series is to offer a single, uniform outlet for high-quality pub- lications arising from major IUFRO meetings and other products of IUFRO’s research units. The editing, publishing and dissemination experience of CABIPublishing and the huge spread of scientific endeavours of IUFRO combine here to make information widely available that is of value to policy makers, resource man- agers, peer scientists and educators. The Executive Board of IUFRO forms the Editorial Advisory Board for the series and provides the monitoring and unifor- mity that such a high-quality series requires in addition to the editorial work of conference organizers. While adding a new body of information to the plethora currently dealing with forestry and related resources, this series seeks to provide a single, uniform forum and style that all forest scientists will turn to first as an outlet for their conference material and other products, and that the users of information will also see as a reliable and reputable source. Although the official languages of IUFRO include English, French, German and Spanish, the majority of modern scientific papers are published in English. In this series, all books will be published in English as the main language, allow- ing papers occasionally to be in other languages. Guidelines for submitting and CO2 0Prels 2/8/01 3:10 pm Page ii ii IUFRO Research Series publishing material in this series are available from the publisher: Books and Reference Works, CABI Publishing, CAB International, Wallingford, Oxon OX10 8DE, UK; and the IUFRO Secretariat, c/o Federal Forest Research Centre, Seckendorff-Gudent-Weg 8, A-1131, Vienna, Austria. IUFRO Executive Board: J. Burley, Oxford, UK M.N. Salleh, Kuala Lumpur, Malaysia D.P. Dykstra, Portland, Oregon, USA K. Sassa, Uji Kyoto, Japan R. Elena Rosello, Madrid, Spain R. Schlaepfer, Lausanne, Switzerland D.W. Flinn, Heidelberg, Victoria, Australia H. Schmutzenhofer, Vienna, Austria R. Guevara Moncada, Turrialba, Costa Rica L. Sennerby-Forsse, Uppsala, Sweden J. Hong, Beijing, China R. Seppaelae, Helsinki, Finland D.F.Karnosky, Houghton, Michigan, USA B. Solberg, As, Norway P.K. Khosla, Palampur, Himachal Pradesh, V.V. Strakhov, Moscow, Russian Federation India E. Teissier du Cros, Avignon, France N.E. Koch, Hoersholm, Denmark K. Vancura, Praha, Czech Republic F.J. Kruger, Sunnyside, South Africa K. von Gadow, Goettingen, Germany D.K. Lee, Seoul, South Korea J.L. Whitmore, Washington, DC, USA J.A. Prado Donoso, Santiago, Chile C. Winget, Ottawa, Ontario, Canada P.N.Sall, Dakar, Senegal J.A. Youngquist, Madison, Wisconsin, USA IUFRO Research Series Titles available: 1. Forest Dynamics in Heavily Polluted Regions Edited by J.L. Innes and J. Oleksyn 2. Forest History: International Studies on Socioeconomic and Forest Ecosystem Change Edited by M. Agnoletti and S. Anderson 3. Methods and Approaches in Forest History Edited by M. Agnoletti and S. Anderson 4. Air Pollution and the Forests of Developing and Rapidly Industrializing Countries Edited by J.L. Innes and A.H. Haron 5. Forests in Sustainable Mountain Development: a State of Knowledge Report for 2000 Edited by M. Price and N. Butt 6. Forests and Landscapes: Linking Ecology, Sustainability and Aesthetics Edited by S.R.J. Sheppard and H.W. Harshaw 7. Criteria and Indicators for Sustainable Forest Management Edited by J. Raison, A. Brown and D. Flinn 8. The Impact of Carbon Dioxide and Other Greenhouse Gases on Forest Ecosystems Edited by D.F. Karnosky, R. Ceulemans, G.E. Scarascia-Mugnozza and J.L. Innes CO2 0Prels 2/8/01 3:10 pm Page iii The Impact of Carbon Dioxide and Other Greenhouse Gases on Forest Ecosystems Report No. 3 of the IUFRO Task Force on Environmental Change Edited by David F. Karnosky Michigan Technological University, Houghton, Michigan, USA Reinhart Ceulemans University of Antwerp, Wilrijk, Belgium Giuseppe E. Scarascia-Mugnozza University of Tuscia, Viterbo, Italy and John L. Innes University of British Columbia, Vancouver, Canada CABI Publishing in association with The International Union of Forestry Research Organizations (IUFRO) CO2 0Prels 2/8/01 3:10 pm Page iv CABI Publishing is a division of CAB International CABI Publishing CABI Publishing CAB International 10 E 40th Street Wallingford Suite 3203 Oxon OX10 8DE New York, NY 10016 UK USA Tel: +44 (0)1491 832111 Tel: +1 212 481 7018 Fax: +44 (0)1491 833508 Fax: +1 212 686 7993 Email: [email protected] Email: [email protected] Web site: www.cabi.org © CAB International2001. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data The impact of carbon dioxide and other greenhouse gases on forest ecosystems: report no. 3 of the IUFRO Task Force on Environmental Change / edited by David F. Karnosky … [et al.]. p. cm. -- (IUFRO research series ; 8) Includes bibliographical references (p. ). ISBN 0-85199-551-9 (alk. paper) 1. Forest plants--Effect of atmospheric carbon dioxide on. 2. Forest plants--Effect of greenhouse gases on. 3. Trees--Effect of atmospheric carbon di- oxide on. 4. Trees--Effect of greenhouse gases on. 5. Atmospheric carbon dioxide--Environmental aspects. 6. Greenhouse gases--Environmental aspects. 7. Forest ecology. I. Karnosky, David, 1949- II. IUFRO Task Force on Environmental Change. III. Series. QK753.C3 I48 2001 577.3(cid:2)276--dc21 2001018103 Published in association with: The International Union of Forestry Research Organizations (IUFRO) c/o Federal Forest Research Centre Seckendorff-Gudent-Weg 8 A-1131 Vienna Austria ISBN 0 85199 551 9 Typeset in 10/12pt Photina by Columns Design Ltd, Reading. Printed and bound in the UK by Biddles Ltd, Guildford and King’s Lynn. CO2 0Prels 2/8/01 3:10 pm Page v Contents Contributors vii Preface ix 1 The Impact of CO and Other Greenhouse Gases on Forest 2 Ecosystems: an Introduction 1 G.E. Scarascia-Mugnozza, D.F. Karnosky, R. Ceulemans and J.L. Innes 2 Effects of Greenhouse Gases on the Gas Exchange of Forest Trees 17 D. Eamus and R. Ceulemans 3 The Impacts of Greenhouse Gases on Antioxidants and Foliar Defence Compounds 57 G.K. Podila, A.R. Paolacci and M. Badiani 4 Above-ground Growth Responses of Forest Trees to Elevated Atmospheric CO Concentrations 127 2 B.E. Medlyn, A. Rey, C.V.M. Barton and M. Forstreuter 5 Influence of CO on the Growth and Function of Roots and 2 Root Systems 147 M.E. Kubiske and D.L. Godbold 6 Impacts of Greenhouse Gases on the Phenology of Forest Trees 193 M.E. Jach, R. Ceulemans and M.B. Murray v CO2 0Prels 2/8/01 3:10 pm Page vi vi Contents 7 Effects of Elevated CO on Nutrient Cycling in Forests 237 2 D.W. Johnson, R.J. Norby and B.A. Hungate 8 Impacts of Interacting Greenhouse Gases on Forest Ecosystems 253 D.F. Karnosky, E. Oksanen, R.E. Dickson and J.G. Isebrands 9 Modelling the Impacts of Greenhouse Gases on Forests 269 F. Magnani and G. Matteucci 10 FACE Systems for Studying the Impacts of Greenhouse Gases on Forest Ecosystems 297 D.F. Karnosky, B. Gielen, R. Ceulemans, W.H. Schlesinger, R.J. Norby, E. Oksanen, R. Matyssek and G.R. Hendrey 11 Knowledge Gaps in the Study of the Impacts of Elevated Atmospheric CO and Other Greenhouse Gases on 2 Forest Ecosystems 325 D.F. Karnosky, G.E. Scarascia-Mugnozza, R. Ceulemans and J. Innes Index 341 CO2 0Prels 2/8/01 3:10 pm Page vii Contributors M. Badiani,Dipartimento di Agrochimica e Agrobiologia, Universita di Reggio Calabria, P. zza S. Francesco di Sales, 4, I-89061 Gallina di Reggio Calabria, Italy C.V.M. Burton,Institute of Terrestrial Ecology, The Bush Estate, Penicuik, Edinburgh EH26 0QB, UK R. Ceulemans, Department of Biology, University of Antwerp, Univer- siteitsplein 1, B-2610 Wilrijk, Belgium. Email: [email protected] R.E. Dickson, USDA Forest Service, Forestry Sciences Laboratory, 5985 Highway K, Rhinelander, WI 54501, USA D. Eamus,Department of Environmental Sciences, University of Technology, Sydney (UTS), PO Box 123, Broadway, NSW 2040, Australia M. Forstreuter,Institut für Okologie der Gehölze, Technische Universitat Berlin, Königin-Luise-Str. 22, D-1000, Berlin 33, Germany B. Gielen,Department of Biology, University of Antwerp, Universiteitsplein 1, B-2610 Wilrijk, Belgium D.L. Godbold, School of Agricultural and Forest Sciences, University of Wales, Bangor, Gwynedd LL57 2UW, UK. Email: [email protected] G.R. Hendrey,Biosystems and Process Sciences Division, Brookhaven National Laboratory, 1 South Technology Street, Upton, NY 11973, USA B.A. Hungate,Northern Arizona University, Flagstaff, Arizona, USA J.L. Innes,Department of Forest Resources Management, University of British Columbia, Forest Sciences Centre 2045, 2424 Main Hall, Vancouver, BC, Canada V6T 1Z4. Email: [email protected] J.G. Isebrands,USDA Forest Service, Forestry Sciences Laboratory, 5985 Highway K, Rhinelander, WI 54501, USA vii CO2 0Prels 2/8/01 3:10 pm Page viii viii Contributors M.E. Jach, University of Antwerp, UIA, Department of Biology, Universiteitsplein 1, B-2610 Wilrijk, Belgium D.W. Johnson, Department of Environmental and Resource Sciences, Fleischmann Agriculture Building/370, University of Nevada, Reno, NV 89557, USA. Email: [email protected] D.F. Karnosky,School of Forestry and Wood Products, Michigan Technological University, 101 U.J. Noblet Forestry Building, 1400 Townsend Drive, Houghton, MI 49931-1295, USA. Email: [email protected] M.E. Kubiske, USDA Forest Service, Forestry Sciences Laboratory, 5985 Highway K, Rhinelander, WI 54501, USA. Email: mkubiske@ cfr.msstate.edu F. Magnani,IMGPF–CNR, Institute of Forest Tree Breeding, National Research Council, via A. Vannucci 13, I-50134 Firenze, Italy. Email: federico@imgpf.fi.cnr.it G. Matteucci, DISAFRI, University of Tuscia, Department of Forest Environment and Resources, Via San Camillo de Lellis s.n.c., I-01100 Viterbo, Italy. Email: [email protected] R. Matyssek,Lehrstuhl für Forstbotanik, Technische Universität München, Am Hochanger 13, D-85354 13 Freising, Germany B.E. Medlyn,IERM, University of Edinburgh, King’s Buildings, Mayfield Road, Edinburgh EH9 3JU, UK; present address: INRA Pierroton, Station de Recherges Forestières, Laboratoire d’Ecophysiologie et Nutrition, BP 45, 33611 Gazinet Cedex, France M.B. Murray,Institute of Terrestrial Ecology, Edinburgh Research Station, Bush Estate, Penicuik, Midlothian EH26 0QB, UK R.J. Norby,Oak Ridge National Laboratory, Bethel Valley Rd, Building 1059, PO Box 2008, Oak Ridge, TN 37831-6422, USA E. Oksanen,Department of Ecology and Environmental Science, University of Kuopio, PO Box 1627, 70211 Kuopio, Finland A.R. Paolacci,Dipartimento di Agrobiologia e Argochimica, Universita della Tuscia, Via S.C. De Lellis, I-01100 Viterbo, Italy G.K. Podila, Department of Biological Sciences, Michigan Technological University, 1400 Townsend Drive, Houghton, MI 49931–1295, USA A. Rey, IERM, University of Edinburgh, King’s Buildings, Mayfield Road, Edinburgh EH9 3JU, UK G.E. Scarascia-Mugnozza,Department of Forest Environment and Resources, University of Tuscia, Via S.C. de Lellis, I-01100 Viterbo, Italy. Email: [email protected] W.H. Schlesinger,Department of Botany and Division of Earth and Ocean Sciences, Nicholas School of Environment, Duke University, Durham, NC 27708-0340, USA CO2 0Prels 2/8/01 3:10 pm Page ix Preface With much effort currently focusing on the role of forests as a sink for atmos- pheric carbon, it is often forgotten that carbon dioxide and other greenhouse gases can have a major effect on trees themselves. A number of atmospheric gases generated by human activities are having an impact on the environment. The most widely recognized is carbon dioxide, but we must not forget that methane and ozone are also greenhouse gases. While we know that carbon dioxide can stimulate tree growth, at least in the short term, ozone has the opposite effect, and in some areas the growth of trees has been severely affected by this gas. In this book, the third report of the IUFRO Task Force on Environmental Change, the state-of-the-art of our knowledge on the effects of greenhouse gases on trees is presented. I am particularly pleased to see that several chapters have authors from more than one country. With experiments designed to investigate the impacts of greenhouse gases becoming increasingly realistic, they are also becoming increasingly complex, and teamwork is essential if we are to utilize fully the technology that has become available. Many of the results presented here represent ‘big science’ at its best, and I am delighted that IUFRO has been able to help bring together some of the different scientists working in this impor- tant field. The fostering of international collaboration between scientists is the raison d’êtreof IUFRO, but another important aspect of its work is the commu- nication of scientific results to those in decision-making positions. I hope that this report will provide such policy makers with some of the basic information that they require to make wise decisions about the stewardship of our environ- ment. Jeff Burley President IUFRO ix