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Hearing. Anatomy, Physiology and Disorders of the Auditory System PDF

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HEARING: ANATOMY, PHYSIOLOGY, AND DISORDERS OF THE AUDITORY SYSTEM Second Edition This page intentionally left blank HEARING: ANATOMY, PHYSIOLOGY, AND DISORDERS OF THE AUDITORY SYSTEM Second Edition A. R. Møller School of Behavioral and Brain Sciences University of Texas at Dallas Texas AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 30 Corporate Drive, Suite 400, Burlington, MA01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Theobald’s Road, London WC1X 8RR, UK This book is printed on acid-free paper. Copyright © 2006, 2000 Elsevier Inc. All rights reserved. No part of this publication 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. Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone: (+44) 1865 843830, fax: (+44) 1865 853333, E-mail: [email protected]. You may also complete your request on-line via the Elsevier homepage (http://elsevier.com), by selecting “Support & Contact” then “Copyright and Permission” and then “Obtaining Permissions.” Front cover design concept by Milda Dorsett. Library of Congress Cataloging-in-Publication Data Møller, Aage R. Hearing : anatomy, physiology, and disorders of the auditory system/A.R. Moller, -- 2nd ed. p. cm. Includes bibliographical references and index. ISBN-13: 978-0-12-372519-6 (casebound : alk. paper) ISBN-10: 0-12-372519-4 (casebound : alk. paper) 1. Hearing--Physiological aspects. 2. Hearing disorders--Pathophysiology. 3. Ear--Anatomy. I. Title. RF290.M58 2006 617.8--dc22 2006014244 British Library Cataloguing-in-Publication Data Acatalogue record for this book is available from the British Library. ISBN 13: 978-0-12-372519-6 ISBN 10: 0-12-372519-4 For information on all Academic Press publications visit our Web site at www.books.elsevier.com Printed in the United States of America 06 07 08 09 10 11 9 8 7 6 5 4 3 2 1 Contents Preface ix 4. Middle Ear 22 Acknowledgements xi 4.1. Middle Ear as an Impedance Introduction xiii Transformer 24 4.2. Transfer Function of the Human Middle Ear 27 S E C T I O N 4.3. Impulse Response of the Human I Middle Ear 29 4.4. Linearity of the Middle Ear 29 THE EAR 4.5. Acoustic Impedance of the Ear 29 4.6. Contributions of Individual Parts of the Middle Ear to Its Impedance 32 CHAPTER 1 Anatomy of the Ear 1. Abstract 3 CHAPTER 3 2. Introduction 3 Physiology of the Cochlea 3. Outer Ear 3 1. Abstract 41 3.1. Ear Canal 5 2. Introduction 41 4. Middle Ear 6 3. Frequency Selectivity of the Basilar 4.1. Tympanic Membrane 6 Membrane 42 4.2. Ossicles 8 3.1. Traveling Wave Motion 43 4.3. Middle-ear Muscles 8 3.2. Basilar Membrane Frequency Tuning 4.4. Eustachian Tube 8 Is Non-linear 44 4.5. Middle-ear Cavities 9 3.3. Frequency Tuning of the Basilar 5. Cochlea 10 Membrane 45 5.1. Organ of Corti 10 3.4. Role of the Outer Hair Cells in Basilar 5.2. Basilar Membrane 13 Membrane Motion 46 5.3. Innervation of Hair Cells 13 3.5. Epochs of Research in Cochlear 5.4. Fluid Systems of the Cochlea 15 Mechanics 47 5.5. Blood Supply to the Cochlea 16 4. Sensory Transduction in the Cochlea 48 4.1. Excitation of Hair Cells 48 4.2. Which Phase of a Sound Excites Hair Cells CHAPTER 2 (Rarefaction or Condensation)? 48 Sound Conduction to the Cochlea 4.3. Molecular Basis for Sensory 1. Abstract 19 Transduction 50 2. Introduction 19 4.4. Endocochlear Potential 52 3. Head, Outer Ear and Ear Canal 20 4.5. Cochlea as a Generator of Sound 53 3.1. Ear Canal 20 4.6. Efferent Control of Hair Cells 55 3.2. Head 20 4.7. Autonomic Control of the Cochlea 55 3.3. Physical Basis for Directional Hearing 21 5. Autoregulation of Blood Flow to the Cochlea 56 v vi Contents CHAPTER 4 3.3. Cochlear Non-linearity Is Reflected in Sound Evoked Electrical Potentials Frequency Selectivity of Auditory Nerve in the Cochlea Fibers 99 3.4. Frequency Tuning in Nuclei of the 1. Abstract 57 Ascending Auditory Pathways 102 2. Introduction 57 3.5. Tonotopic Organization in the Nuclei of the 3. Electrical Potentials in the Cochlea 57 Ascending Auditory Pathways 104 3.1. Cochlear Microphonics 58 3.6. Extraction of Information from Place 3.2. Summating Potential 59 Coding of Frequency 106 3.3. Action Potential 59 4. Coding of Temporal Features 106 3.4. Electrocochleographic Potentials 64 4.1. Coding of Periodic Sounds 107 Section I References 68 4.2. Extraction of Information from the Temporal Pattern of Neural Discharges 111 S E C T I O N 5. Is Temporal or Place Code the Basis for II Discrimination of Frequency? 112 5.1. Temporal Hypothesis for Frequency Discrimination of Complex Sounds 112 THE AUDITORY NERVOUS SYSTEM 5.2. Place Hypothesis for Frequency Discrimination of Complex Sounds 113 5.3. Preservation of the Temporal Code of CHAPTER 5 Frequency 115 Anatomy of the Auditory Nervous System 5.4. Preservation of the Place Code 117 1. Abstract 75 5.5. Robustness of the Temporal Code 117 2. Introduction 76 5.6. Robustness of the Place Code of 3. Classical Ascending Auditory Pathways 76 Frequency 117 3.1. Auditory Nerve 76 5.7. Coding of Speech Sounds 117 3.2. Cochlear Nucleus 80 5.8. ADuplex Hypothesis of Frequency 3.3. Superior Olivary Complex 81 Discrimination 118 3.4. Lateral Lemniscus and Its Nuclei 81 5.9. Cochlear Spectral Filtering May Be 3.5. Inferior Colliculus 81 Important in other Ways than Frequency 3.6. Medial Geniculate Body 81 Discrimination 118 3.7. Auditory Cerebral Cortex 82 5.10. Speech Discrimination on Spectral 3.8. Differences between the Classical Auditory Information Only 118 Pathways in Humans and in Animals 84 5.11. Conclusion 119 4. Non-classical Ascending Auditory Pathways 85 6. Coding of Complex Sounds 119 5. Parallel Processing and Stream Segregation 87 6.1. Response to Tone Bursts 120 5.1. Parallel Processing 88 6.2. Coding of Small Changes in 5.2 Stream Segregation 88 Amplitude 121 5.3. Connections to Non-auditory Parts of the 6.3. Response to Tones with Changing Brain 89 Frequency 128 6. Descending Pathways 89 6.4. Selectivity to Other Temporal Patterns of Sounds 138 6.5. Coding of Sound Intensity 140 CHAPTER 6 6.6. Conclusion 140 Physiology of the Auditory Nervous 7. Directional Hearing 142 System 7.1. Physical Basis for Directional Hearing 142 1. Abstract 93 7.2. Neurophysiologic Basis for Sound 2. Introduction 94 Localization 143 3. Representation of Frequency in the Auditory 7.3. Localization in the Vertical Plane 146 Nervous System 95 7.4. Representation of Auditory Space 3.1 Hypotheses about Discrimination (Maps) 146 of Frequency 95 8. Efferent System 149 3.2. Frequency Selectivity in the Auditory 9. Non-classical Pathways 149 Nervous System 97 10. Effect of Anesthesia 150 Contents vii CHAPTER 7 3. Pathologies of the Sound Conducting Evoked Potentials from the Nervous Apparatus 206 System 3.1. Ear Canal 207 3.2. Middle Ear 207 1. Abstract 151 3.3. Impairment of Sound Conduction in the 2. Introduction 152 Cochlea 213 3. Near-field Potentials from the Auditory 3.4. Accuracy of Measurements of Conductive Nervous System 152 Hearing Loss 213 3.1. Recordings from the Auditory Nerve 152 3.5. Implications of Impairment of Conduction 3.2. Recordings from the Cochlear of Sound to the Cochlea 214 Nucleus 160 4. Pathologies of the Cochlea 215 3.3. Recordings from More Central Parts of the 4.1. General Audiometric Signs of Cochlear Ascending Auditory Pathways 163 Pathologies 215 4. Far-field Auditory Evoked Potentials 163 4.2. Age-related Hearing Loss (Presbycusis) 216 4.1. Auditory Brainstem Responses 165 4.3. Noise Induced Hearing Loss 219 4.2. Middle Latency Responses 175 4.4. Implications of Hearing Loss on Central 4.3. Far-field Frequency Following Responses Auditory Processing 226 in Humans 176 4.5. Modification of Noise Induced Hearing 4.4. Myogenic Auditory Evoked Potentials 177 Loss 226 4.6. Hearing Loss Caused by Ototoxic Agents (Drugs) 227 CHAPTER 8 4.7. Diseases that Affect the Function of the Acoustic Middle-ear Reflex Cochlea 229 1. Abstract 181 4.8. Congenital Hearing Impairment 233 2. Introduction 181 4.9. Infectious Diseases 234 3. Neural Pathways of the Acoustic Middle-ear 4.10. Perilymphatic Fistulae 234 Reflex 182 4.11. Changes in Blood Flow in the Cochlea 234 4. Physiology 183 4.12. Injuries to the Cochlea from Trauma 234 4.1. Responses to Stimulation with Tones 184 4.13. Sudden Hearing Loss 234 4.2. Functional Importance of the Acoustic 5. Implications of Hearing Loss on Central Auditory Middle-ear Reflex 187 Processing 235 4.3. Non-acoustic Ways to Elicit Contraction of the 5.1 Neural Components of Hearing Loss 236 Middle-ear Muscles 190 5.2. Role of Expression of Neural Plasticity 237 4.4. Stapedius Contraction May Be Elicited before 6. Pathologies from Damage to the Auditory Vocalization 190 System 239 5. Clinical Use of the Acoustic Middle-ear Reflex 190 6.1. Auditory Nerve 239 Section II References 192 6.2. Other Space-occupying Lesions 243 7. Pathologies of the Central Auditory Nervous System 243 S E C T I O N 7.1. Disorders of the Brainstem Auditory III Pathways 244 7.2. Auditory Cortices 244 DISORDERS OF THE AUDITORY 7.3. Efferent System 246 7.4. Pathologies that Can Affect Binaural SYSTEM AND THEIR Hearing 246 PATHOPHYSIOLOGY 7.5. Viral Infections 246 7.6. Ototoxic Drugs 247 7.7. Sudden Hearing Loss 247 CHAPTER 9 8. Role of Neural Plasticity in Disorders of the Hearing Impairment Central Auditory Nervous System 247 1. Abstract 205 8.1. What Is Neural Plasticity? 247 2. Introduction 206 8.2. What Can Initiate Expression of Neural Plasticity? 250 viii Contents CHAPTER 10 3.1. Function and Design of Auditory Hyperactive Disorders of the Auditory Brainstem Implants 277 System 3.2. Physiological Basis for Auditory Brainstem Implants 277 1. Abstract 253 3.3. Success of Auditory Brainstem Implants 278 2. Introduction 253 3.4. Patient Selection for Auditory Brainstem 3. Subjective Tinnitus 254 Implants 279 3.1. Assessment of Tinnitus 254 4. Role of Neural Plasticity 279 3.2. Disorders in which Tinnitus Is Section III References 280 Frequent 255 3.3. Causes of Subjective Tinnitus and Other Hyperactive Symptoms 255 APPENDIX A 3.4. Role of Expression of Neural Plasticity Definitions in Anatomy 289 in Tinnitus 259 4. Abnormal Perception of Sounds 260 4.1. Hyperacusis 261 4.2. Phonophobia 262 APPENDIX B 4.3. Misophonia 262 Hearing Conservation Programs 291 4.4. Recruitment of Loudness 262 1. Introduction 291 5. Treatment of Subjective Tinnitus 263 2. Purpose and Design of Hearing Conservation 5.1. Medical Treatment 264 Programs 292 5.2. Electrical Stimulation 264 2.1. Basis for Hearing Conservation Programs 292 5.3. Surgical Treatment 265 3. Establishment of Noise Standards 295 5.4. Desensitization 266 3.1. Noise Level and Exposure Time 296 6. Treatment of Hyperacusis 266 3.2. Effect of Age-related Hearing Loss 296 3.3. What Degree of Hearing Loss is Acceptable? 296 CHAPTER 11 4. Measurement of Noise 297 Cochlear and Brainstem Implants 4.1. Sound Level Meters 297 1. Introduction 267 4.2. Noise Dosimeters 298 2. Cochlear Implants 268 5. Personal Protection 299 2.1. Development of Cochlear Implants 268 5.1. Earplugs and Earmuffs 299 2.2. Function and Design of Cochlear 5.2. Active Noise Cancellation 300 Implants 268 5.3. Other Means of Reducing the Risk of Noise 2.3. Physiological Basis for Cochlear Implants 273 Induced Hearing Loss 300 2.4. Coding of Sound Intensity 275 6. Non-occupational Noise Exposure 300 2.5. Functions that Are Not Covered by Modern 7. Effect of Noise on Bodily Functions 300 Cochlear Implants 275 7.1. Effect of Ultrasound and Infrasound 300 2.6. Success of Cochlear Implants 276 Appendices References 301 2.7. Selection Criteria for Cochlear Implant List of Abbreviations 303 Candidates 277 Index 305 3. Cochlear Nucleus Implants 277 Preface This book is intended for otologists, audiologists, by Academic Press, and the second, Hearing: Its neurologists and researchers in the field of hearing. The Physiology and Pathophysiology, published in 2000, also book will also be of interest to psychologists and psy- by Academic Press. chiatrists who treat patients with tinnitus and other The book has 11 chapters that are organized in three hyperactive auditory disorders. The book provides the sections. Chapters from earlier editions have been basis for a broad understanding of the anatomy and re-organized and most parts have been re-written and function of the ear and the auditory nervous system, new information has been added. Aseparate chapter and it discusses the cause and treatment of hearing is devoted to an extended coverage of hyperactive disorders. Most books on hearing focus either on the disorders, most importantly tinnitus, the cause and anatomy and function of the ear, the auditory nervous treatment of which is discussed in detail. Anew chap- system or on peripheral or central hearing disorders. ter describes cochlear and brainstem implants and This book covers both anatomy and physiology of the hearing conservation programs are discussed in an ear and the nervous system. The book also provides a appendix. comprehensive coverage of disorders of the auditory The four chapters of Section I cover anatomy and system emphasizing the interaction between patholo- physiology of the middle ear and the cochlea, includ- gies of the middle ear and the cochlea and the function ing a chapter on the electrical potentials that are gen- of the nervous system and vice versa. Hyperactive dis- erated by the cochlea. Section II has two chapters that orders of the auditory nervous system and the role of cover anatomy and physiology of the nervous system. expression of neural plasticity in causing auditory Both the classical and the less known non-classical symptoms are also topics of the book. An extensive list (extralemniscal) auditory pathways are covered exten- of references makes it possible for the reader to find sively. The latter is involved in some forms of tinnitus original work on the different subjects. and may be activated in other disorders also. Athird Understanding of the anatomy and the function of chapter is devoted to evoked potentials from the nerv- the auditory system together with knowledge about ous system. The neural generators of the ABR are dis- the pathophysiology of the auditory system are essen- cussed in detail. The anatomy and physiology of the tial for all clinicians who are involved in diagnosis and acoustic middle-ear reflex is covered in a fourth chapter treatment of disorders of the auditory system. The in this section. book prepares the clinician and the clinical researcher The final section (Section III) discusses disorders of for the challenges of the modern clinical auditory disci- the auditory system. Two chapters regard hearing pline. The book also provides basic information about impairment and hyperactive disorders, focusing on tin- the auditory system in a form that is suitable for the sci- nitus, its etiology, and treatment. These two chapters entist who does basic research on the auditory system. stress the role of expression of neural plasticity. Athird The book thus aims at cross-fertilization between clini- chapter in this section concerns cochlear implants and cians, clinical researchers and basic scientists. It is my auditory brainstem implants. The basic design and hope that such knowledge can guide basic auditory function of the processors in these modern auditory research into clinically relevant questions. prostheses are described and the physiologic basis for The book is the third edition of books on the auditory the function of these prostheses is discussed. An system, the first, Auditory Physiology, published in 1983 appendix discusses hearing conservation programs. ix

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Hearing: Anatomy, Physiology and Disorders of the Auditory System provides detailed information about the anatomy and physiology of the entire auditory system and it describes important aspects of disorders of the middle ear, the cochlea, and the nervous system in a comprehensive manner. Most other
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