Vitamin C Vitamin C Function and biochemistry in animals and plants Edited by Han Asard Department of Biochemistry, Beadle Center, University of Nebraska-Lincoln, 1901 Vine Street, Lincoln, NE 68588, USA James M.May 715 Preston Research Building, Vanderbilt University School of Medicine, 2220 Pierce Ave., Nashville, TN 37232–6303, USA Nicholas Smirnoff School of Biological Sciences, University of Exeter, Washington Singer Laboratories, Perry Road, Exeter EX4 4QG, UK LONDON AND NEW YORK © Garland Science/BIOS Scientific Publishers, 2004 First published 2004 This edition published in the Taylor & Francis e-Library, 2005. “To purchase your own copy of this or any of Taylor & Francis or Routledge's collection of thousands of eBooks please go to www.eBookstore.tandf.co.uk.” All rights reserved. No part of this book may be reprinted or reproduced or utilised in any form or by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying and recording, or in any information storage or retrieval system, without permission in writing from the publishers. A CIP catalogue record for this book is available from the British Library. ISBN 0-203-50000-8 Master e-book ISBN ISBN 0-203-59762-1 Adobe e-Reader Format ISBN 1 85996 293 9 (Print Edition) Garland Science/BIOS Scientific Publishers 4 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN, UK and 29 West 35th Street, New York, NY 10001–2299, USA World Wide Web home page: www.bios.co.uk Garland Science/BIOS Scientific Publishers is a member of the Taylor & Francis Group. Distributed in the USA by Fulfilment Center Taylor & Francis 10650 Toebben Drive Independence, KY 41051, USA Toll Free Tel.: +1 800 634 7064; E-mail: [email protected] Distributed in Canada by Taylor & Francis 74 Rolark Drive Scarborough, Ontario MIR 4G2, Canada Toll Free Tel.: +1 877 226 2237; E-mail: [email protected] Distributed in the rest of the world by Thomson Publishing Services Cheriton House North Way Andover, Hampshire SP1O 5BE, UK Tel.: +44(0)1264 332424; E-mail: [email protected] Library of Congress Cataloging-in-Publication Data Vitamin C : its function and biochemistry in animals and plants / edited by Han Asard, James May, Nicholas Smirnoff. p. cm. Includes bibliographical references and index. ISBN 1-85996-293-9 (hardcover : alk. paper) 1. Vitamin C. I. Asard, Han. II. May, James. III. Smirnoff, N. QP772.A8V567 2003 572′.58–dc22 2003017985 Production Editor: Andrew Watts Table of contents Contributors viii Acknowledgements xi Abbrevations xii Introduction 1 1. Ascorbate biosynthesis: a diversity of pathways 8 N.Smirnoff, J.A.Running and S.Gatzek 2. Ascorbic acid catabolism: breakdown pathways in animals and 35 plan ts G.Bánhegyi and F.A.Loewus 3. The biotechnology of ascorbic acid manufacture 55 J.A.Running, S.Peng and R.A.Rosson 4. The role of ascorbic acid in defense networks and signaling in plants 73 C.H.Foyer 5. Ascorbate and plant growth: from germination to cell death 92 L.De Gara 6. Vitamin C transport in animals and plants 107 J.X.Wilson 7. Membrane redox proteins involved in ascorbate-mediated reactions 131 J.M.Villalba, M.del Carmen Córdoba-Pedregosa and J.A.González-Reyes 8. Ascorbate recycling 153 F.May and H.Asard 9. How does ascorbic acid prevent scurvy? A survey of the nonantioxidant 176 functions of vitamin C M.C.De Tullio 10. Ascorbate as an antioxidant 191 G.R.Buettner and F.Q.Schafer 11. Vitamin C and oxidative damage to DNA 209 H.E.Poulsen, P.MØller, F.Lykkesfeldt, A.Weimann and S.Loft 12. Vitamin C status declines with age 224 A.F.Michels and T.M.Hagen 13. Ascorbic acid in the central nervous system: uptake, distribution and 253 functions R.J.Reiter 14. Physiology of vitamin C in neutrophils during inflammation 272 J.W.Heinecke 15. Ascorbic acid in atherosclerosis 287 P.K.Witting and R.Stocker 16. Dietary allowances for vitamin C: Recommended Dietary Allowances and 321 optimal nutrient ingestion M.Levine, S.Padayatty, A.Katz, O.Kwon, P.Eck, C.Corpe, J.-H.Lee and Y.Wang Index 353 Contributors Asard, H. Department of Biochemistry, Beadle Center, University of Nebraska-Lincoln, 1901 Vine Street, Lincoln, NE 68588, USA Bánhegyi, G. Department of Medical Chemistry, Molecular Biology and Pathobiology, Semmelweis University, 9 Puskin Street, 1088 Budapest, Hungary Buettner, G.R. University of lowa, Free Radical Research Group, EMRB 68 lowa City,IA 52242–1101, USA Córdoba-Pedregosa, M.del Carmen Departamento de Biologia Celular, Fisiologia e Immunologia, Universidad de Córdoba, Campus de Rabanales, Edificio C-6, Planta 3a, 14014 Córdoba, Spain Corpe, C. Molecular and Clinical Nutrition Section, Bld 10, Rm 4D52, MSC 1372 National Institute of Diabetes, Digestive, and Kidney Diseases, National Institutes of Health, Bethesda, MD 20892–1372, USA De Gara, L. Dipartimento di Biologia e Patologia Vegetale, V.E Orabona, 4I-70126 Bari, Italy De Tullio, M. Dipartimento di Biologia e Patologia Vegetale, V.E Orabona, 4I-70126 Bari, Italy Eck, P. Molecular and Clinical Nutrition Section, Bld 10, Rm 4D52, MSC 1372 National Institute of Diabetes, Digestive, and Kidney Diseases, National Institutes of Health, Bethesda, MD 20892–1372, USA Foyer, C.H. Crop Performance and Improvement Division, Rothamsted Research, Harpenden, Herts AL5 2JQ, United Kingdom Frei, B. Linus Pauling Institute, 571 Weniger Hall, Oregon State University, Corvallis, OR 97331–6512, USA Gatzek, S. Max Planck Institute of Molecular Plant Physiology, Am Mühlenberg 1, D- 14476 Golm, Germany González-Reyes, J.A. Departamento de Biologia Celular, Fisiologia e Immunologia, Universidad de Córdoba, Campus de Rabanales, Edificio C-6, Planta 3a, 14014 Córdoba, Spain Hagen, T.M. Linus Pauling Institute, Oregon State University, Corvallis, OR 97331, USA Heinecke, J. Medicine/Metabolism, Box 356426, University of Washington, 1959 NE Pacific Street, HSB BB512, Seattle, WA 98195–6426, USA Higdon, J.V. Linus Pauling Institute, Oregon State University, Corvallis, OR 97331, USA Katz, A. Molecular and Clinical Nutrition Section, Bld 10, Rm 4D52, MSC 1372 National Institute of Diabetes, Digestive, and Kidney Diseases, National Institutes of Health, Bethesda, MD 20892–1372, USA Kwon, U. Molecular and Clinical Nutrition Section, Bld 10, Rm 4D52, MSC 1372 National Institute of Diabetes, Digestive, and Kidney Diseases, National Institutes of Health, Bethesda, MD 20892–1372, USA Lawson, S. Linus Pauling Institute, Oregon State University, Corvallis, OR 97331, USA Lee, J.-H. Molecular and Clinical Nutrition Section, Bld 10, Rm 4D52, MSC 1372 National Institute of Diabetes, Digestive, and Kidney Diseases, National Institutes of Health, Bethesda, MD 20892–1372, USA Levine, M. Molecular and Clinical Nutrition Section, Bld 10, Rm 4D52, MSC 1372 National Institute of Diabetes, Digestive, and Kidney Diseases, National Institutes of Health, Bethesda, MD 20892–1372, USA Loewus, F.A. 1700 NE Upper Drive, Pullman, WA 99163–4624, USA Loft, S. Department of Clinical Pharmacology, Q-7642, Rigshopitalet, Copenhagen University Hospital, 20 Tagensvej, DK-2200 Copenhagen N, Denmark Lykkesfeldt, J. Department of Clinical Pharmacology, Q-7642, Rigshopitalet, Copenhagen University Hospital, 20 Tagensvej, DK-2200 Copenhagen N, Denmark May, J.M. 715 Preston Research Building, Vanderbilt University School of Medicine, 2220 Pierce Ave., Nashville, TN 37232–6303, USA Michels, AJ. 571 Weniger Hall, Linus Pauling Institute, Oregon State University Corvallis,OR 97331, USA MØller, P. Department of Clinical Pharmacology, Q-7642, Rigshopitalet, Copenhagen University Hospital, 20 Tagensvej, DK-2200 Copenhagen N, Denmark Padayatty, S. Molecular and Clinical Nutrition Section, Bld 10, Rm 4D52, MSC 1372 National Institute of Diabetes, Digestive, and Kidney Diseases, National Institutes of Health, Bethesda, MD 20892–1372, USA Peng, S. Bio-Technical Resources, 1035 South 7th Street, Manitowoc, WI 54220, USA Poulsen, H.E. Department of Clinical Pharmacology, Q-7642, Rigshopitalet, Copenhagen University Hospital, 20 Tagensvej, DK-2200 Copenhagen N, Denmark Reiter, RJ. Department of Cellular and Structural Biology, The University of Texas Health Science Center, San Antonio, TX 78284–7762, USA Rosson, R.A. Bio-Technical Resources, 1035 South 7th Street, Manitowoc, WI 54220, USA Running, J.A. Bio-Technical Resources, 1035 South 7th Street, Manitowoc, WI 54220, USA Schafer, F.Q. University of lowa, Free Radical & Radiation Biology, Dept of Radiation Oncology, EMRB 68, lowa City, IA 52242–1101, USA Smirnoff, N. School of Biological Sciences, University of Exeter, Washington Singer Laboratories, Perry Road, Exeter EX4 4QG, United Kingdom Stocker, R. Centre for Thrombosis and Vascular Research, School of Medical Sciences, University of New South Wales, Sydney, UNSW 2052, Australia Villalba, J.-M. Departamento de Biologia Celular, Fisiologia e Immunologia, Universidad de Córdoba, Campus de Rabanales, Edificio C-6, Planta 3a, 14014 Córdoba, Spain Wang, Y. Molecular and Clinical Nutrition Section, Bld 10, Rm 4D52, MSC 1372 National Institute of Diabetes, Digestive, and Kidney Diseases, National Institutes of Health, Bethesda, MD 20892–1372, USA Weimann, A. Department of Clinical Pharmacology, Q-7642, Rigshopitalet, Copenhagen University Hospital, 20 Tagensvej, DK-2200 Copenhagen N, Denmark