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Auditory Computation PDF

527 Pages·1996·26.552 MB·English
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SPRINGER HANDBOOK OF AUDITORY RESEARCH Series Editors: Richard R. Fay and Arthur N. Popper Springer New York Berlin Heidelberg Barcelona Budapest Hong Kong London Milan Paris Santa Clara Singapore Tokyo SPRINGER HANDBOOK OF AUDITORY RESEARCH Volume I: The Mammalian Auditory Pathway: Neuroanatomy Edited by Douglas B. Webster, Arthur N. Popper, and Richard R. Fay Volume 2: The Mammalian Auditory Pathway: Neurophysiology Edited by Arthur N. Popper and Richard R. Fay Volume 3: Human Psychophysics Edited by William Yost, Arthur N. Popper, and Richard R. Fay Volume 4: Comparative Hearing: Mammals Edited by Richard R. Fay and Arthur N. Popper Volume 5: Hearing by Bats Edited by Arthur N. Popper and Richard R. Fay Volume 6: Auditory Computation Edited by Harold L. Hawkins, Teresa A. McMullen, Arthur N. Popper, and Richard R. Fay Volume 7: Clinical Aspects of Hearing Edited by Thomas R. Van De Water, Arthur N. Popper, and Richard R. Fay Volume 8: The Cochlea Edited by Peter Dallas, Arthur N. Popper, and Richard R. Fay Volume 9: Development of the Auditory System Edited by Edwin W Rubel, Arthur N. Popper, and Richard R. Fay Forthcoming Volumes (partial list) Comparative Hearing: Insects Edited by Ronald Hoy, Arthur N. Popper, and Richard R. Fay Speech Processing in the Auditory System Edited by Steven Greenberg, William Ainsworth, Arthur N. Popper, and Richard R. Fay Comparative Hearing: Fish and Amphibians Edited by Arthur N. Popper and Richard R. Fay Comparative Hearing: Birds and Reptiles Edited by Robert Dooling, Arthur N. Popper, and Richard R. Fay Harold L. Hawkins Teresa A. McMullen Arthur N. Popper Richard R. Fay Editors Auditory Computation With 138 Illustrations i Springer Harold L. Hawkins Teresa A. McMullen Office of Naval Research Office of Naval Research 800 N. Quincy Street 800 N. Quincy Street Arlington, VA 22217, USA Arlington, VA 22217, USA Arthur N. Popper Richard R. Fay Department of Zoology Parmly Hearing Institute University of Maryland Loyola University of Chicago College Park, MD 20742-9566, USA Chicago, I~ 60626, USA Series Editors: Richard R. Fay and Arthur N. Popper Cover illustration: Equivalent electrical circuit model for a cochlear hair cell. This figure appears on p. 125 of the text. Library of Congress Cataloging in Publication Data Auditory computation I editors, Harold L. Hawkins ... let al.]. p. cm.-(Springer handbook of auditory research: v. 6) Includes bibliographical references and index. ISBN-13: 978-1-4612-8487-1 e-ISBN-13: 978-1-4612-4070-9 DOl: 10.1007/978-1-4612-4070-9 1. Auditory perception-Mathematical models. I. Hawkins. Harold L. II. Series. QP461.A916 1995 612.8'5'01l-dc20 95-12906 Printed on acid-free paper. © 1996 Springer-Verlag New York, Inc. Softcover reprint oft he hardcover 1s t edition 1996 Copyright is not claimed for works by U.S. Government employees. 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 WOW, 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 namcs, 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 Best-set Typesetter, Ltd., Chaiwan, Hong Kong. 9 8 7 6 5 4 3 2 Series Preface The Springer Handbook of Auditory Research presents a series of com prehensive and synthetic reviews of the fundamental topics in modern 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 focuses on topics that have developed a solid data and con ceptual 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 that 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. Richard R. Fay Arthur N. Popper Preface The purpose of this volume is to provide an overview and understanding of computational analyses of auditory system function. The approach expressed in the chapters of this volume embodies the ideas that complex information processing must be understood at multiple levels of analysis, that it must be addressed from a multidisciplinary perspective that incor porates the constraints imposed by neurobiology, psychophysics, and computational analysis, and that the end product of computational analysis should be the development of formal models. These ideas and approaches arc dealt with in the ten chapters of this volume. In the first chapter, Hawkins and McMullen provide an overview of the volume as well as a brief introduction to the computational ap proaches that are used to increase understanding of auditory information processing. The next several chapters arc organized with respect to levels of auditory analysis and processing. In Chapter 2, Rosowski discusses models of the external and middle ears, while in Chapter 3 Hubbard and Mountain consider computational issues related to inner car function, primarily cochlear mechanics. In Chapter 4, these same authors (Mountain and Hubbard) discuss computational analysis of the hair cells and eighth nerve fibers that transduce and transmit information to the brain. In Chapter 5 Delgutte treats extraction of acoustic features such as fre quency, intensity, and loudness at the eighth nerve. The remaining chapters are concerned with computations involving higher levels of the central auditory system, from the cochlear nucleus to the auditory cortex. In Chapter 6 Lyon and Shamma describe computations underlying the extraction of pitch and timbre, and Mellinger and Mont-Reynaud (Chapter 7) develop the computational basis of auditory scene analysis. In Chapter R Colburn summarizes the empirical findings on binaural hearing and describes a number of computational models based on these findings. In Chapter 9 Simmons outlines several computational models of the processes by which spectral and temporal information is combined in the bat auditory system to reconstruct target images. In the final chapter vii viii Preface (10), Lewis describes inferential computations that could underlie the analysis and retention of sequences of acoustic events. While this volume stands alone, it is important to point out that there are a number of other chapters and volumes in this series that are relevant to the issues discussed in this volume. For example, the psycho physical basis for much of the computational analysis described herein by Lyon and Shamma, Delgutte, and by Colburn is considered in detail in Volume 3 of this series (Human Psychophysics, 1993, Yost, Popper and Fay, eds.) and by Long in Volume 4 of the series (Comparative Hearing: Mammals, 1994, Fay and Popper, eds.). The structures of the middle and inner ears that form the bases for the chapters by Rosowski and by Hubbard and Mountain are also described in detail in by Rosowski and Echteler, Fay and Popper, respectively, in Volume 4. The discussions of Mountain and Hubbard on computation in the eighth nerve are comple mented by papers on eighth nerve anatomy by Ryugo in Volume 1 (The Mammalian Auditory Pathway: Neuroanatomy, 1992, Webster, Popper and Fay, eds.) and by Ruggero on physiology of the eighth nerve in Volume 2 (The Marnmalian Auditory Pathway: Neurophysiology, 1992, Popper and Fay, eds.). The chapter on binaural processing by Colburn is complemented by chapters on the neurophysiological basis of such pro cessing by Irvine in Volume 2, while sound localization is considered in detail in Volume 4 by Brown and by Wightman and Kistler in Volume 3. Finally, the chapter by Simmons et al. on biosonar is paralleled by a chapter on psychophysical aspects of this system in a chapter in Volume 5 (Hearing by Bats, 1995, Popper and Fay, eds.). Harold L. Hawkins Teresa A. McMullen Arthur N. Popper Richard R. Fay Contents Series Preface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v Preface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . VI Contributors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . XI Chapter 1 Auditory Computation: An Overview ............ . 1 HAROLD L. HAWKINS AND TERESA A. McMuLLEN Chapter 2 Models of External- and Middle-Ear Function ..... . 15 JOHN J. ROSOWSKI Chapter 3 Analysis and Synthesis of Cochlear Mechanical Function Using Models ......................... . 62 ALLYN E. HUBBARD AND DAVID C. MOUNTAIN Chapter 4 Computational Analysis of Hair Cell and Auditory Nerve Processes ............................... . 121 DAVID C. MOUNTAIN AND ALLYN E. HUBBARD Chapter 5 Physiological Models for Basic Auditory Percepts ...................................... . 157 BERTRAND DELGUTfE Chapter 6 Auditory Representations of Timbre and Pitch 221 RICHARD LYON AND SHIHAB SHAMMA Chapter 7 Scene Analysis ................................ . 271 DAVID K. MELLINGER AND BERNARD M. MONT-REYNAUD Chapter 8 Computational Models of Binaural Processing ..... . 332 H. STEVEN COLBURN ix x Contents Chapter 9 Auditory Computations for Biosonar Target Imaging in Bats ....................................... . 401 JAMES A. SIMMONS, PRESTOR A. SAILLANT, MICHAEL J. FERRAGAMO, TIM HARESIGN, STEVEN P. DEAR, JONATHAN FRITZ, AND TERESA A. McMuLLEN Chapter 10 Further Computations Involving Time. . . . . . . . . . . . . 469 EDWIN R. LEWIS Index ............ '" ..... , . . . . . . . . . . . . .. . . . .. . . . ... . . . . . . . 505 Contributors H. Steven Colborn Biomedical Engineering Department, Boston University, Boston, MA 02215, USA Steven P. Dear Department of Psychology, Brown University, Providence, RI 02912, USA Bertrand Delgutte Eaton Peabody Laboratory, Massachusetts Eye & Ear Infirmary, Boston, MA 02114, USA Michael J. Ferragamo Department of Psychology, Brown University, Providence, RI 02912, USA Jonathan Fritz Department of Psychology, Brown University, Providence, RI 02912, USA Tim Haresign Department of Psychology, Brown University, Providence, RI 02912, USA Harold L. Hawkins Office of Naval Research, Arlington , VA 22217, USA Allyn E. Hubbard Biomedical Engineering Department, Boston University, Boston, MA 02215, USA Edwin Lewis Department of Electrical Engineering and Computer Science, University of California, Berkeley, Berkeley, CA 94720, USA xi

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