Brain Repair ADVANCES IN EXPERIMENTAL MEDICINE AND BIOLOGY Editorial Board: NATHAN BACK, State University of New York at Buffalo IRUN R. COHEN, The Weizmann Institute of Science DAVID KRITCHEVSKY, Wistar Institute ABEL LAJTHA, N.S. Kline Institute for Psychiatric Research RODOLFO PAOLETTI, University of Milan Recent Volumes in this Series Volume 561 CHEMISTRY AND SAFETY OF ACRYLAMIDE IN FOOD Edited by Mendel Friedman and Don Mottram Volume 562 CHOLINERGIC MECHANISMS Edited by Jos6 Gonzalez-Ros Volume 563 UPDATES IN PATHOLOGY Edited by David C. Chhieng and Gene P. Siegal Volume 564 GLYCOBIOLOGY AND MEDICINE: Proceedings of the 7* Jenner Glycobiology and Medicine Symposium Edited by John S. Axford Volume 565 SLIDING FILAMENT MECHANISM IN MUSCLE CONTRACTION: Fifty Years of Research Edited by Haruo Sugi Volume 566 OXYGEN TRANSPORT TO TISSUE XXVI Edited by Paul Okunieff, Jacqueline Williams and Yuhchyau Chen Volume 567 THE GROWTH HORMONE-INSULIN-LIKE GROWTH FACTOR AXIS DURING DEVELOPMENT Edited by Isabel Varela-Nieto and Julie A. Chowen Volume 568 HOT TOPICS IN INFECTION AND IMMUNITY IN CHILDREN II Edited by Andrew J. Pollard and Adam Finn Volume 569 EARLY NUTRITION AND ITS LATER CONSEQUENCES: New Opportunities Edited by Berthold Koletzko, Peter Dodds, Hans Akerbloom and Margaret Ashwell A Continuation Order Plan is available for this series. A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment. For further information please contact the publisher. Brain Repair Edited by Mathias Bahr Department of Neurology University ofGottingen Gottingen, Germany Springer Science+Business Media Landes Bioscience / Eurekah.com Springer Science+Business Media Eurekah.com / Landes Bioscience Copyright ©2006 Eurekah.com and Springer Science+Business Media All rights reserved. No part of this book may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system; for exclusive use by the Purchaser of the work. Printed in the U.S.A. Springer Science+Business Media, 233 Spring Street, New York, New York 10013, U.S.A. http://www.springeronline.com Please address all inquiries to the Publishers: Eurekah.com / Landes Bioscience, 810 South Church Street, Georgetown, Texas, U.S.A. 78626 Phone: 512/ 863 7762; FAX: 512/ 863 0081 http ://www.eurekah.com http://www.landesbioscience.com Brain Repair edited by Mathias Bahr, Landes Bioscience / Springer Science+Business Media dual imprint / Springer series: Advances in Experimental Medicine and Biology ISBN: 0-306-47859-5 While the authors, editors and publisher believe that drug selection and dosage and the specifications and usage of equipment and devices, as set forth in this book, are in accord with current recommend ations and practice at the time of publication, they make no warranty, expressed or implied, with respect to material described in this book. In view of the ongoing research, equipment development, changes in governmental regulations and the rapid accumulation of information relating to the biomedical sciences, the reader is urged to carefully review and evaluate the information provided herein. Library of Congress Catalogmg-in-Publication Data Brain repair / [edited by] Mathias Bahr. p.; cm. ~ (Advances in experimental medicine and biology ; V. 557) Includes bibliographical references and index. ISBN 0-306-47859-5 1. Brain—Physiology. 2. Brain-Regeneration. 3. Cell cycle —Regulation. 4. Apoptosis. I. Bahr, Mathias. II. Series. [DNLM: 1. Nervous System—physiology. 2. Alzheimer Disease. 3. Apoptosis. 4. Cell Cycle. 5. Neurons—physiology. WL 102 B81335 2006] QP376.B7197 2006 612.8'2-dc22 2005019652 PARTICIPANTS Young Hwan Ahn Thomas Deller Neuronal Survival Unit Institute of Clinical Neuroanatomy Section for Neuroscience Johann Wolfgang Goethe-University Department of Experimental Medical Frankfiirt am Main, Germany Studies Wallenberg Neuroscience Center Jorg Dietrich Lund, Sweden Department of Neurology Massachusetts General Hospital Mathias Bahr Harvard Medical School Department of Neurology Boston, Massachusetts, U.S.A. University of Gottingen Gottingen, Germany Alexandre Dobbertin INSERM U686 Christine E. Bandtlow Centre Universitaire des Saints-Peres Innsbruck Medical University Paris, France Biocenter Innsbruck Division of Neurobiochemistry Andreas Faissner Innsbruck, Austria Department of Cell Morphology and Molecular Neurobiology Patrik Brundin Ruhr-University Neuronal Survival Unit Bochum, Germany Section for Neuroscience Department of Experimental Medical James W. Fawcett Studies Centre for Brain Repair Wallenberg Neuroscience Center Cambridge University Lund, Sweden Cambridge, U.K. Fabio Cavaliere Fulvio Florenzano I.R.C.C.S. Santa Lucia Foundation I.R.C.C.S. Santa Lucia Foundation Rome, Italy Rome, Italy Participants Thomas M. Freiman Susanne Hermanns Department of Neurosurgery NEURAXO BIOTEC GmbH Neurocenter Diisseldorf, Germany University Clinic Freiburg Albert-Ludwigs-University Mathias Jucker Freiburg, Germany Department of Cellular Neurology Hertie-Institute for Clinical Brain Michael Frotscher Research Institute of Anatomy and Cell Biology University of Tubingen University of Freiburg Tubingen, Germany Freiburg, Germany Gerd Kempermann Yutaka Fukuda Max-Delbriick-Center Department of Physiology of Molecular Medicine Graduate School of Medicine Berlin-Buch, Germany Osaka University and Osaka, Japan Department of Neurology Charite University Hospital Jeremy Garwood Humboldt University Berlin Centre de Neurochimie du CNRS Berlin, Germany Strasbourg, France Nicole Klapka Marcia Gasis Molecular Neurobiology Laboratory Molecular Neurobiology Laboratory Department of Neurology Department of Neurology Heinrich-Heine-University Heinrich-Heine-University Diisseldorf, Germany Diisseldorf, Germany Kerstin Krieglstein Carola A. Haas Zentrum Anatomic, Experimental Epilepsy Group Abt. Neuroanatomie Neurocenter Universitat Gottingen University Clinic Freiburg Gottingen, Germany Albert-Ludwigs-University Freiburg, Germany Takuji Kurimoto Department of Ophthalmology Nicolas Heck Hyogo College of Medicine Institute of Physiology Hyogo, Japan and Pathophysiology University of Mainz Mainz, Germany Participants Jia-Yi Li Amie Phinney Neuronal Survival Unit Department of Neuropathology Section for Neuroscience Institute of Pathology Department of Experimental Medical University of Basel Studies Basel, Switzerland Wallenberg Neuroscience Center Lund, Sweden Tommaso Pizzorusso Institute of Neuroscience CNR Rafael Linden and Scuola Normale Superiore Instituto de Biofisica da UFRJ Pisa, Italy CCS Cidade Universitaria Gian Michele Ratto Rio de Janeiro, Brazil Institute of Neuroscience CNR and Scuola Normale Superiore Paul Lingor Pisa, Italy Department of Neurology University of Gottingen Mariana S. Silveira Gottingen, Germany Laboratorio de Neurogenese Instituto de Biofisica da UFRJ Tomomitsu Miyoshi Rio de Janeiro, Brazil Department of Physiology Graduate School of Medicine Maria Teresa Viscomi Osaka University I.R.C.C.S. Santa Lucia Foundation Osaka, Japan Rome, Italy Marco Molinari Cinza Volonte I.R.C.C.S. Santa Lucia Foundation I.R.C.C.S. Santa Lucia Foundation Rome, Italy Rome, Italy and Hans Werner Miiller Institute of Neurobiology Molecular Neurobiology Laboratory and Molecular Medicine (CNR) Department of Neurology Rome, Italy Heinrich-Heine-University Diisseldorf Diisseldorf, Germany Anne D. Zum Department of Experimental Surgery Gesine Paul Lausanne University Hospital Neuronal Survival Unit Faculty of Biology and Medicine Section for Neuroscience Lausanne, Switzerland Department of Experimental Medical Studies Wallenberg Neuroscience Center Lxmd, Sweden CONTENTS 1. CELL DEATH IN THE NERVOUS SYSTEM 1 Kerstin Krieglstein Programmed Cell Death 1 Neuronal Cell Death during Development 1 Cell Death in Early Neural Development 2 Glial CeU Death 2 Molecular Mechanisms of Programmed Cell Death 2 Caspase Independent Pathways 4 Cell Death Receptors 4 Extrinsic Mechanisms 5 Cell Death during Neurodegenerative Disorders and Aging 5 2. THE GLL\L RESPONSE TO INJURY AND ITS ROLE IN THE INHIBITION OF CNS REPAIR 11 James W. Fawcett What Is the Glial Scar? 11 Control of Glial Scar Formation 12 The Glial Scar and Axon Regeneration 12 Inhibitory Glial Boundaries 12 Inhibitory Molecules in the Damaged CNS 13 Inhibitory Molecules Produced by Oligodendrocytes 13 Inhibitory Molecules Produced in the Glial Scar 13 Individual CSPGs 15 Glial Boundaries in the CNS 18 Strategies for Repairing the CNS 20 3. DSD-1-PROTEOGLYCAN/PHOSPHACAN AND RECEPTOR PROTEIN TYROSINE PHOSPHATASE-BETAISOFORMS DURING DEVELOPMENT AND REGENERATION OF NEURAL TISSUES 25 Andreas Faissner, Nicolas Heck, Alexandre Dobbertin and Jeremy Garwood General Introduction 25 DSD-1-PG/Phosphacan/RPTPbeta 27 Contents X 4. REGENERATION FAILURE IN THE CNS: CELLULAR AND MOLECULAR MECHANISMS 54 Anne D. Zum and Christine E. Bandtlow Intrinsic Neuronal Properties 54 The CNS Environment Is Non-Permissive for Axonal Growth 57 Experimental Strategies to Promote Nerve Regeneration Following CNS Injury 65 5. THE ROLE OF lONOTROPIC PURINERGIC RECEPTORS (P2X) IN MEDIATING PLASTICITY RESPONSES IN THE CENTRAL NERVOUS SYSTEM 77 Fulvio Florenzano, Maria Teresa Viscomi, Fabio Cavaliere, Cinza Volonte and Marco Molinari Neurobiology of P2 Receptors 77 6. LESION-INDUCED AXONAL SPROUTING IN THE CENTRAL NERVOUS SYSTEM 101 Thomas Deller, Carola A. Haas, Thomas M. Freiman, Amie Phinney, Mathias Jucker and Michael Frotscher Reorganization of the CNS after Entorhinal Lesioning 103 Molecular Regulation of Axonal Sprouting 103 Extracellular Matrix Molecules Direct Axonal Growth Processes 106 Axonal Sprouting in Alzheimer's Disease 107 Axonal Sprouting in Temporal Lobe Epilepsy 113 What Is the Functional Significance of Axonal Sprouting? 115 7. A KINASE WITH A VISION: ROLE OF ERK IN THE SYNAPTIC PLASTICITY OF THE VISUAL CORTEX 122 Gian Michele Ratto and Tommaso Pizzorusso Critical Period for Ocular Dominance 122 Factors Critical for Ocular Dominance: Electrical Activity and Neurotrophins 123 Activation of ERK 1/2 Is Necessary for BDNF-Induced Phosphorylation of CREB 125 ERK 1/2 Is Phosphorylated by Activity and Visual Stimulation in the Cortex 127 ERK Activation Is Required for Synaptic Plasticity in Vitro and in Vivo 128 Beyond ERK: Mechanisms Controlling Synaptic Plasticity in the Visual Cortex .... 129 Contents xi 8. ATTEMPTS TO RESTORE VISUAL FUNCTION AFTER OPTIC NERVE DAMAGE IN ADULT MAMMALS 133 Tomomitsu Miyoshi, Takuji Kurimoto and Yutaka Fukuda Optic Nerve Regeneration and Functional Recovery of Vision in Rodents 133 Attempts to Promote RGC Survival and Their Axonal Regeneration in Rodents.... 134 RGC Survival and Their Axonal Regeneration in Adult Cats 137 Neuroprotection of RGCs by Electrical Stimulation 141 9. BRAIN REPAIR: EXPERIMENTAL TREATMENT STRATEGIES, NEUROPROTECTIVE AND REPAIR STRATEGIES IN THE LESIONED ADULT CNS 148 Mathias Bahr and Paul Linger Optic Nerve Transsection as Exemplary Model for CNS Lesions 148 Post-Traumatic and Ontogenetic Cell Death in the Retino-Tectal System 149 Key Players in Apoptotic Cell Death: Possible Targets for Therapeutic Strategies 150 Therapeutic Agents 154 Ways to Enter the Brain 156 10. NEUROPROTECTION BY CAMP: ANOTHER BRICK IN THE WALL 164 Mariana S. Silveira and Rafael Linden Programmed CeU Death and Signal Transduction Pathways 164 Cell Death by Apoptosis and the Role of cAMP-Dependent Protein Kinase in Apoptotic Execution Pathways 165 Cyclic AMP and CeU Death 166 Neuroprotection by Cyclic AMP 167 Signal Transduction Pathways Regulated by cAMP * 170 11. THE COLLAGENOUS WOUND HEALING SCAR IN THE INJURED CENTRAL NERVOUS SYSTEM INHIBITS AXONAL REGENERATION 177 Susanne Hermanns, Nicole Klapka, Marcia Gasis and Hans Werner Miiller Peripheral and Central Nervous System Responses to Axotomy 177 Factors in the Lesion Area Impede Axonal Regeneration 177 Basement Membrane Formation in the Injured CNS Coincides with Axonal Growth Arrest 178