Hormones and Brain Plasticity Oxford Series in Behavioral Neuroendocrinology Series Editors Gregory F. Ball, Jacques Balthazart, and Randy J. Nelson Hormones and Brain Plasticity LUIS MIGUEL GARCIA-SEGURA 1 2009 1 Oxford University Press, Inc., publishes works that further Oxford University’s objective of excellence in research, scholarship, and education. Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offi ces in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Th ailand Turkey Ukraine Vietnam Copyright © 2009 by Luis Miguel Garcia-Segura Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York 10016 www.oup.com Oxford is a registered trademark of Oxford University Press All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press. Library of Congress Cataloging-in-Publication Data Garcia-Segura, Luis Miguel. Hormones and brain plasticity / Luis Miguel Garcia-Segura. p. ; cm.—(Oxford series in behavioral neuroendocrinology ; 1) Includes bibliographical references and index. ISBN 978-0-19-532661-1 (cloth : alk. paper) 1. Neuroendocrinology. 2. Neuroplasticity. I. Title. II. Series. [DNLM: 1. Brain—physiology. 2. Neuronal Plasticity—physiology. 3. Aging—physiology. 4. Brain—growth & development. 5. Hormones—physiology. WL 300 G216h 2009] QP356.4.G37 2009 612.8—dc22 2008042771 9 8 7 6 5 4 3 2 1 Printed in the United States of America on acid-free paper To my mentor, Alfredo Carrato Ibáñez (1911–1994) For his gift of the light of science in times of deep darkness In memoriam This page intentionally left blank Contents Preface and Acknowledgments xiii Chapter 1 Hormones and the Mutable Brain 3 Introduction: Endocrine Glands, Brain Plasticity, Homeostasis, Allostasis, and Homeodynamics 3 Th e Nervous System is in a Constant State of Remodeling: Cellular Plasticity and Cellular Replacement 8 Th e Classic Concept of Brain Plasticity: Cajal’s Heritage 11 Cellular Replacement: A New Concept 15 Role of Hormones on Brain Mutability: Plasticity and Metaplasticity 16 Chapter 2 Brain Plasticity Regulates Hormonal Homeodynamics 21 Introduction: Brain Plasticity Regulates the Activity of Endocrine Glands 21 Neuroglial Remodeling Associated with Hormonal Secretion by Hypothalamic Magnocellular Neurons 23 Neuroglial Remodeling in the Neurohypophysis 29 Neuroglial Remodeling Associated with Parvocellular Hypothalamic Neurons 30 Neuroglial Remodeling in the Median Eminence 34 Brain Plasticity and the Control of Hormonal Homeodynamics: A Recapitulation 41 Chapter 3 H ormonal Infl uences on Brain Plasticity: I. Melatonin, Th yroid Hormones and Corticosteroids 43 Introduction: A Large Variety of Hormones Regulate Brain Plasticity 43 vii viii Contents Hormones or Paracrine Factors? 47 Melatonin 48 Th yroid Hormones 51 Stress, Corticosteroids, and Brain Plasticity 53 Chapter 4 Hormonal Infl uences on Brain Plasticity: II. Sex Hormones 65 Introduction: Sex Hormones and Brain Remodeling 65 Th e First Evidences 65 Regulation by Estradiol and Progesterone of Synaptic and Glial Remodeling in the Arcuate Nucleus 69 Regulation by Estradiol and Progesterone of Synaptic Remodeling in the Ventromedial Hypothalamic Nucleus 78 Regulation by Estradiol and Progesterone of Synaptic Remodeling in the Hippocampus 79 Regulation by Estradiol and Progesterone of Glial Remodeling in the Hippocampus 86 Regulation by Estradiol and Progesterone of Adult Neurogenesis 88 Role of Brain Estradiol Synthesis in the Regulation of Synaptic Plasticity and Neuronal Replacement 91 Androgens, Synaptic Plasticity, and Cellular Replacement in the Song System of Songbirds 96 Androgens and Synaptic Plasticity in Motoneurons 98 Androgens, Synaptic plasticity, and Cellular Replacement in the Mammalian Brain 99 Chapter 5 Hormonal Infl uences on Brain Plasticity: III. Peptide Hormones 105 Introduction: Peptide Hormones as Regulators of Brain Remodeling 105 Vasopressin and Oxytocin 106 Corticotropin-Releasing Hormone and Urocortins 108 Prolactin 111 Gonadotropin-Releasing Hormone 112 Insulin 113 Growth Hormone/Insulin-Like Growth Factor-I 118 Insulin-Like Growth Factor-I and Synaptic Plasticity 120 Insulin-Like Growth Factor-I and Adult Neurogenesis 122 Interactions of Insulin-Like Growth Factor-I and Estradiol in the Regulation of Brain Plasticity 125 Erythropoietin 131 Angiotensin 133 Contents ix Feeding Hormones: Leptin, Ghrelin and Glucagon-Like Peptide-1 134 Hormonal Infl uences on Brain Plasticity: Recapitulation of the Chapters 3, 4 and 5 139 Chapter 6 Life Stages, Hormones, and Brain Remodeling: Early Hormonal Infl uences on Brain Mutability 141 Introduction: Hormonal Imprinting of the Nervous System 141 Hormonal Orchestration of Neural Development during Prenatal Life 145 Th yroid Hormones 145 Sex Hormones 147 Enduring Eff ects of Prenatal Stress 151 Peptide Hormones 154 Hormonal Regulation of Brain Mutability during Early Postnatal Life 156 Metamorphosis in Vertebrates 156 Th yroid Hormone Regulation of Postnatal Brain Development in Non-Metamorphic Vertebrates 158 Gonadal Hormones 159 Stress Hormones 169 Peptide Hormones 171 Early Hormonal Infl uences on Brain Mutability: Recapitulation 174 Chapter 7 Life Stages, Hormones, and Brain Remodeling: Th e Transition from Childhood to Adulthood 177 Introduction: Prepubertal Maturation, Puberty, and Adolescence 177 Brain Remodeling during the Transition from Childhood to Adulthood 178 Hormones as Regulators of Brain Plasticity during Prepubertal Maturation, Puberty, and Adolescence 182 Brain Remodeling as a Cause of Hormonal Changes during Puberty 186 Chapter Summary 190 Chapter 8 Life Stages, Hormones, and Brain Remodeling: Adult Reproductive Life 193 Introduction: Hormonal Regulation of Brain Plasticity during Adult Life 193
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