SPRINGER HANDBOOK OF AUDITORY RESEARCH. Series Editors: Richard R. Fay and Arthur N. Popper

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1 SPRINGER HANDBOOK OF AUDITORY RESEARCH Series Editors: and Arthur N. Popper

2 SPRINGER HANDBOOK OF AUDITORY RESEARCH Volume 1: The Mammalian Auditory Pathway: Neuroanatomy Edited by Douglas B. Webster, Arthur N. Popper, and Volume 2: The Mammalian Auditory Pathway: Neurophysiology Edited by Arthur N. Popper and Volume 3: Human Psychophysics Edited by William Yost, Arthur N. Popper, and Volume 4: Comparative Hearing: Mammals Edited by and Arthur N. Popper Volume 5: Hearing by Bats Edited by Arthur N. Popper and Volume 6: Auditory Computation Edited by Harold L. Hawkins, Theresa A. McMullen, Arthur N. Popper, and Volume 7: Clinical Aspects of Hearing Edited by Thomas R. Van de Water, Arthur N. Popper, and Forthcoming Volumes (partial list) Development of the Auditory System Edited by Edwin Rubel, Arthur N. Popper, and The Cochlea Edited by Peter Dallos, Arthur N. Popper, and Plasticity in the Auditory System Edited by Edwin Rubel, Arthur N. Popper, and

3 Douglas B. Webster Arthur N. Popper Editors The Mammalian Auditory Pathway: Neuroanatomy With 110 Illustrations Springer-Verlag New York Berlin Heidelberg London Paris Tokyo Hong Kong Barcelona Budapest

4 Douglas B. Webster Kresge Hearing Research Laboratory of the South Department of Otorhinolaryngology Louisiana State University New Orleans, LA Arthur N. Popper Department of Zoology University of Maryland College Park, MD Parmly Hearing Institute and Department of Psychology Loyola University of Chicago Chicago, IL Series Editors: and Arthur N. Popper Cover illustration: Golgi-impregnated neurons from the medial geniculate body of a cat. The form and distribution can be related to the neurochemical circuitry of the auditory thalamus. This is a detail from Figure 6.4A, p Library of Congress Cataloging-in-Publication Data The mammalian auditory pathway: Neuroanatomy I Douglas B. Webster, Arthur N. Popper,, editors. p. cm. - (Springer handbook of auditory research; v. 1) Includes bibliographical references and index. 1. Auditory pathways-anatomy. 2. Neuroanatomy. I. Webster, Douglas B., II. Popper, Arthur N. III. Fay, Richard R. IV. Series. [DNLM: 1. Anatomy, Comparative. 2. Auditory Pathways-anatomy & histology. WV 272 N4932] QM451.N '.048-dc20 DNLM/DLC for Library of Congress CIP Printed on acid-free paper Springer-Verlag New York, Inc. All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag New York, Inc., 175 Fifth Avenue, New York, NY 10010, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use of general descriptive names, trade names, trademarks, etc., in this publication, even if the former are not especially identified, is not to be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Production managed by Terry Kornak; manufacturing supervised by Jacqui Ashri Typeset by Impressions, Madison, Wisconsin, a division of Edwards Brothers, Inc ISBN-l3: e-isbn-13: DOl: /

5 Series Preface The Springer Handbook of Auditory Research presents a series of comprehensive and synthetic reviews of the fundamental topics in modem auditory research. It is aimed at all individuals with interests in hearing research including advanced graduate students, postdoctoral researchers, and clinical investigators. The volumes will introduce new investigators to important aspects of hearing science and will help established investigators to better understand the fundamental theories and data in fields of hearing that they may not normally follow closely. Each volume is intended to present a particular topic comprehensively, and each chapter will serve as a synthetic overview and guide to the literature. As such, the chapters present neither exhaustive data reviews nor original research that has not yet appeared in peer-reviewed journals. The series focusses on topics that have developed a solid data and conceptual foundation rather than on those for which a literature is only beginning to develop. New research areas will be covered on a timely basis in the series as they begin to mature. Each volume in the series consists of five to eight substantial chapters on a particular topic. In some cases, the topics will be ones of traditional interest for which there is a solid body of data and theory, such as auditory neuroanatomy (Vol. 1) and neurophysiology (Vol. 2). Other volumes in the series will deal with topics which have begun to mature more recently, such as development, plasticity, and computational models of neural processing. In many cases, the series editors will be joined by a co-editor having special expertise in the topic of the volume. Arthur N. Popper v

6 Preface When students or colleagues have asked us where they could learn about the organization ofthe mammalian central auditory system, we have had no good place to send them. Neuroanatomy texts treat the subject superficially, and often incorrectly. Review articles assume a previous sophisticated knowledge. The original literature is too specialized for the novice. We saw a need for a different source of information. This volume is meant to answer that need. It is intended for basic researchers, graduate and postdoctoral students, and for our colleagues from other fields who wish to learn about auditory neuroanatomy-including basic scientists as well as audiologists, otologists, and speechlanguage pathologists. Chapter 1 presents an overview of the entire neural auditory system, from the periphery through the auditory cortex, with an emphasis on the human system. The following six chapters treat. each region of the auditory pathway in detail. Each chapter is authored by an expert who not only understands the specialty but can write about it clearly for the nonspecialist. This volume is the first in the series; it presents the structural basis for what will follow. The companion Volume 2 has parallel chapters presenting the neurophysiology of the mammalian auditory system. Succeeding volumes will deal specifically with the cochlea, human and animal psychoacoustics, development, and the auditory neurophysiology and neuroanatomy of nonmammalian vertebrates. As editors we are indebted to the chapter authors for their time and effort, and to the staff of Springer-Verlag for their enthusiastic cooperation. Douglas B. Webster Arthur N. Popper Vll

7 Contents Series Preface... Preface... Contributors... Chapter 1 An Overview of Mammalian Auditory Pathways with an Emphasis on Humans.... DOUGLAS B. WEBSTER Chapter 2 The Auditory Nerve: Peripheral Innervation, Cell Body Morphology, and Central Projections DAVID K. RYUGO Chapter 3 The Cochlear Nucleus: Neuronal Types and Their Synaptic Organization NELL BEATTY CANT Chapter 4 The Superior Olivary Complex and Lateral Lemniscal Nuclei ILSA R. SCHWARTZ Chapter 5 Inferior and Superior Colliculi DOUGLAS L. OLIVER AND MICHAEL F. HUERTA Chapter 6 The Functional Architecture of the Medial Geniculate Body and the Primary Auditory Cortex JEFFERY A. WINER Chapter 7 Organization of Olivocochlear Efferent Systems in Mammals W. BRUCE WARR Subject Index Author Index v vn xi ix

8 Contributors Nell Beatty Cant Department of Neurobiology, Duke University Medical Center, Durham, NC 27710, USA Michael F. Huerta Department of Anatomy, The University of Connecticut Health Center, Farmington, CT 06032, USA Douglas L. Oliver Department of Anatomy, The University of Connecticut Health Center, Farmington, CT 06032, USA David K. Ryugo Center for Hearing Sciences, Departments of Otolaryngology-Head and Neck Surgery and Neuroscience, Johns Hopkins University School of Medicine, Baltimore, MD 21205, USA Ilsa R. Schwartz Department of Surgery/Otolaryngology, Yale University School of Medicine, New Haven, CT 06510, USA W. Bruce WaIT Boys Town National Research Hospital, Omaha, NE 68131, USA Douglas B. Webster Kresge Hearing Research Laboratory of the South, Department of Otorhinolaryngology, Louisiana State University, New Orleans, LA , USA Jeffery A. Winer Division of Neurobiology, Department of Molecular and Cell Biology, University of California, Berkeley, CA , USA xi