BIOS Instant Notes Series Analytical Chemistry Medical Microbiology David Kealey and P. J. Haines Will Irving, Tim Boswell, and Dlawer 978-1-85996-189-6 Ala’Aldeen 978-1-85996-254-1 Biochemistry, 4e David Hames and Nigel Hooper Medicinal Chemistry 978-0-415-60845-9 Graham Patrick 978-1-85996-207-7 Bioinformatics, 2e Charlie Hodgman, Andrew French, Microbiology, 4e and David Westhead Simon Baker, Jane Nicklin, and 978-0-415-39494-9 Caroline Griffiths 978-0-415-60770-4 Chemistry for Biologists, 2e J. Fisher and J. R. P. Arnold Molecular Biology, 3e 978-1-85996-355-5 Phil Turner, Alexander McLennan, Andy Bates, and Michael White Genetics, 3e 978-0-415-35167-6 Hugh Fletcher, Ivor Hickey, and Paul Winter Neuroscience 3e 978-0-415-37619-8 Alan Longstaff 978-0-415-60769-8 Human Physiology Daniel Lydyard, Jonathan Stamford, Organic Chemistry, 2e and David White Graham Patrick 978-0-415-35546-9 978-1-85996-264-0 Immunology, 3e Physical Chemistry Peter Lydyard, Alex Whelan, and Gavin Whittaker, Andy Mount, and Michael Fanger Matthew Heal 978-0-415-60753-7 978-1-85996-194-0 Inorganic Chemistry, 2e Plant Biology 2e Tony Cox Andrew Lack and David Evans 978-1-85996-289-3 978-0-415-35643-5 Mathematics and Statistics for Life Scientists Aulay McKenzie 978-1-85996-292-3 For up-to-date title information and current prices please visit: www.garlandscience.com BIOS INSTANT NOTES Neuroscience THIRD EDITION This page is intentionally left blank. BIOS INSTANT NOTES Neuroscience THIRD EDITION Alan Longstaff Associate Lecturer, Open University This edition published in the Taylor & Francis e-Library, 2011. 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. Garland Science Vice President: Denise Schanck Editor: Elizabeth Owen Editorial Assistant: Louise Dawnay Production Editor: Ioana Moldovan Copyeditor: Sally Huish ©2011 by Garland Science, Taylor & Francis Group, LLC 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. 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—graphic, electronic, or mechanical, including photocopying, recording, taping, or information storage and retrieval systems—without permission of the copyright holder. ISBN 0-203-80829-0 Master e-book ISBN ISBN 978-0-4156-0769-8 (Print Edition) Library of Congress Cataloging-in-Publication Data Longstaff, Alan. Neuroscience / Alan Longstaff. -- 3rd ed. p. cm. -- (Bios instant notes) Includes index. ISBN 978-0-415-60769-8 1. Neurosciences--Outlines, syllabi, etc. I. Title. RC343.6.L66 2011 616.8--dc22 2011003160 Published by Garland Science, Taylor & Francis Group, LLC, an informa business, 270 Madison Avenue, New York NY 10016, USA, and 2 Park Square, Milton Park, Abingdon, OX14 4RN, UK. Visit our web site at http://www.garlandscience.com Preface Neuroscience is multidisciplinary, having contributions from molecular biol- ogy, anatomy, physiology, pharmacology, psychology, and medicine to name the most obvious. Moreover, the neuroscience literature is huge. For these rea- sons textbooks of neuroscience tend to be large, making it hard for students to discriminate the conceptual heart of the subject from exemplars and enrich- ment material. This Third Edition of Instant Notes in Neuroscience is intended as a supplement to lectures that gives rapid and easy access to the core of the subject in an affordable and manageable-sized text. When coming to a new subject, students commonly express three concerns; firstly, how to sort out the important ideas and facts from the wealth of detail; secondly, how to get to grips with the unfamiliar terminology; and thirdly, how to integrate their knowledge across the various disciplines, a necessary step for a good understanding of the subject. The Instant Notes format addresses these issues. Each topic is supported by a “Key Notes” panel which gives a concise summary of the crucial points. Whenever a term appears for the first time it is in bold and immediately followed by a definition or explanation. Extensive cross references are provided between topics so that students can forge the links that are important for integration. Instant Notes in Neuroscience is a much slimmer volume than most neuro- science texts. Several features contribute to this. I have tried to minimize the amount of detail without compromising the need for students to have a data- base for autonomous learning. While many of the methods used by neurosci- entists are included, individual experiments or items of evidence are included only where I thought it essential to illustrate a point. I have restricted examples to mammals, and the anatomy is largely human, even though there is a great deal of remarkable work in birds, fish, and invertebrates. This third edition is over one-fifth shorter than its predecessors. This has been achieved by remov- ing two sections (Developmental Neurobiology and Brain Disorders), by sim- plifying the language of the text throughout, and by re-drafting many of the more complicated diagrams. However, I have not shied away from retaining conceptually challenging material where it is warranted. The Third Edition of Instant Notes in Neuroscience has 15 sections. Section A sets the scene by introducing the cells of the nervous system, and looks at how the nervous system is organized by taking a broad view of neuroanatomy. The next three sections are essentially cellular neuroscience, concerned with elec- trophysiology, the properties of synapses, and an introduction to the principal neurotransmitters and their receptors. Elementary neural computing is intro- duced in Section E which looks at rate and temporal coding and how neurons are connected to make simple circuits. All the material thus far might reason- ably be found in first year courses. The next six sections (F–K) form the core of systems neuroscience. These concentrate on sensory and motor neurobiology, but introduce visual attention and other cognitive aspects of brain function where needed. Section L tackles neuroendocrinology, how the brain controls metabolism, growth, and reproduction, and the autonomic nervous system. vi PREFACE The perspective of the next two sections is rather broader than what has come before. Behaviors, such as emotion, motivation, and sleep are explored in section M. Section N examines learning and memory, homing in on the episodic learning in the hippocam- pus and motor learning in the cerebellum as examples. Finally, Section O looks at the techniques used to explore the brain, neuroimaging, electrophysiology, and some of the remarkable methods being developed in the last few years such as brainbow and opto- genetics. As a student, how should you use this book? Restrict your reading only to the sections and topics covered by your current course. That said, sections A–E are likely to be part of any neuroscience program; you will probably need to work through these first. Later sections can be dipped into in any order. Read the main sections thoroughly first, making sure that you understand the ideas, and use the “Related topics” to make links. This is the stage to incorporate additional material from lectures, and other textbooks, in the gaps at the end of topics. For areas that particularly interest you, turn to “Further Reading” at the end of the book. Although by no means a comprehensive bibliography, following up the refer- ences in them will get you into the rest of the neuroscience literature. Studying Instant Notes “little but often” is a good strategy. The information density in the text is high, so many short, concentrated, bursts is the most effective way to study. The more times you work through a topic, the better your understanding will be, and the more likely you will remember it clearly. When it comes to revision, use the “Key Notes” as a prompt. In addi- tion, you should aim to be able to write, from memory, a few sentences about each of the terms that appear in bold in the main text. Being able to reproduce the simpler diagrams is also an effective way of getting your point across in an exam. I thank Elizabeth Owen and Louise Dawnay of Taylor & Francis for their hard work and encouragement; each helped shape the project in distinctive and important ways. I hope you enjoy studying neuroscience as much as I do. Contents Preface v Section A – Organization of the nervous system 1 A1 Neurons 1 A2 Glial cells 5 A3 Organization of the peripheral nervous system 7 A4 Organization of the central nervous system 13 A5 Meninges and cerebrospinal fluid 19 A6 Blood–brain barrier 23 Section B – Neuron excitation 25 B1 Membrane potentials 25 B2 Voltage-dependent ion channels 30 B3 Action potential conduction 36 Section C – Synapses 40 C1 Synapse structure and function 40 C2 Neurotransmitter release 45 C3 Postsynaptic responses 51 C4 Neurotransmitter inactivation 55 C5 Neural integration 58 Section D – Neurotransmitters 60 D1 Ligand-gated ion channel receptors 60 D2 G-protein-coupled receptors 65 D3 Amino acid transmitters 71 D4 Dopamine 74 D5 Noradrenaline (norepinephrine) 78 D6 Serotonin 81 D7 Acetylcholine 84 D8 Purines and peptides