Pitch: Neural Coding and Perception PDF
Preview Pitch: Neural Coding and Perception
Springer Handbook of Auditory Research Series Editors: Richard R. Fay and Arthur N. Popper Christopher J. Plack Andrew J. Oxenham Richard R. Fay Arthur N. Popper Editors Pitch Neural Coding and Perception With 74 illustrations and 5 color illustrations Christopher J. Plack AndrewJ. Oxenham Departmentof Psychology ResearchLaboratoryof Electronics University of Essex MassachusettsInstitute of Technology ColchesterCO4 3SQ Cambridge, MA 02139, USA United Kingdon [email protected] [email protected] Richard R. Fay Arthur N. Popper Parmly HearingInstitute and Departmentof Biology Departmentof Psychology University of Maryland Loyola Universityof Chicago College Park, MD 20742, USA Chicago, IL 60626, USA [email protected] [email protected] Cover illustration: The image includes parts of Figures 4.6 and 6.4 appearingin the text. LibraryofCongressCataloging-in-PublicationData Pitch:neuralcodingandperception/[editedby]ChristopherJ.Plack,AndrewJ.Oxenham, RichardR.Fay,ArthurN.Popper. p. cm.—(Springerhandbookofauditoryresearch;v.24) Includesbibliographicalreferencesandindex. ISBN10:0-387-23472-1(alk.paper) 1. Auditoryperception. 2. Musicalpitch. I. Plack,ChristopherJ. II. Oxenham, AndrewJ. III. Fay,RichardR. IV. Series. QP465.P545 2005 152.1'52—dc22 2004057843 ISBN10:0-387-23472-1 Printedonacid-freepaper ISBN13:978-0387-23472-4 (cid:1)2005SpringerScience(cid:1)BusinessMedia,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 Science(cid:1)Business Media, Inc., 233 Spring Street, NewYork, NY 10013, 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 developedisforbidden. Theuseinthispublicationoftradenames,trademarks,servicemarks,and similarterms,eveniftheyarenotidentifiedassuch,isnottobetakenasanexpressionofopinion astowhetherornottheyaresubjecttoproprietaryrights. PrintedintheUnitedStatesofAmerica. (EB) 9 8 7 6 5 4 3 1 springeronline.com Each of the editors takes pleasure in dedicating this volume to his parents in gratitude for their support and guidance: Audrey and Jim Plack Margaret and John Oxenham Ingrid and Charles Fay Evelyn and Martin Popper Series Preface TheSpringerHandbookofAuditoryResearchpresentsaseriesofcomprehensive and synthetic reviews of the fundamental topics in modern auditory research. The volumes are aimed at all individuals with interests in hearing research in- cluding advanced graduate students, postdoctoral researchers, and clinical in- vestigators. The volumes are intended to introduce new investigators to important aspects of hearing science and to help established investigators to betterunderstandthefundamentaltheoriesanddatainfieldsofhearingthatthey may not normally follow closely. Each volume presents a particular topic comprehensively, and each servesas 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 volumes focus 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. Eachvolumeintheseriesconsistsofafewsubstantialchaptersonaparticular topic. In some cases, the topics will be ones of traditional interest for which there is a substantial body of data and theory, such as auditory neuroanatomy (Vol. 1) and neurophysiology (Vol. 2). Other volumes in the series deal with topics that have begun to mature more recently, suchasdevelopment,plasticity, and computational models of neural processing. In many cases, the series ed- itorsarejoinedbyaco-editorhavingspecialexpertiseinthetopicofthevolume. Richard R. Fay, Chicago, Illinois Arthur N. Popper, College Park, Maryland vii Volume Preface TheseedsforthisvolumeonpitchweresowninOctober2001,whenWolfgang Stenzel, Andrew Oxenham, and Chris Plack met for dinner in a Spanishrestau- rant in Bremen, Germany. They discussed the possibility of organizing a con- ference on pitch perception to be hosted by the Hanse Wissenschaftskolleg (Hanse Institute for Advanced Study) in Delmenhorst (Wolfgang Stenzel ad- ministers the Neurosciences and Cognitive Sciences Program at the Institute). The proposal to the Institute began as follows: “Although pitch has been con- sideredanimportantareaofauditoryresearchsincethenineteenthcentury,some of the most significant developments in our understanding of this phenomenon haveoccurredcomparativelyrecently. Thetimeisripeforameetingthatbrings togetherexpertsfromseveraldifferentdisciplinestoshareideasandgaininsights into the fundamental (and still largely unsolved) problem of how the brainpro- cesses the pitch of acoustic stimuli.” The conference took place August 2002, bringing together scientists in the fields of neuroscience, computationalmodel- ing, cognitive science, and music psychology. Rather than publish a standard conference proceedings, Plack and Oxenham approachedArthurPopperandRichardFayaboutproducingthisvolume,which is a “stand-alone” review of the current state of pitch research, inspiredby(but not limited to) the presentations and discussions at the conference. All the chapter authors attended the conference, and, like the conference, the volume brings together researchers from a range of different disciplines. It is hoped thatthereadermayobtainabroadviewofthetopicfrombasicneurophysiology to more cognitive processes. Chapter 1, by Plack and Oxenham, provides a definition of pitch and an overview of the field. A description of the basic psychophysics of pitch is the focusofChapters2and3. PlackandOxenham(Chapter2)describehowhuman perceptions are related to the physical characteristicsof thestimulusandasim- ilar approach is taken in a discussion of psychophysical studies on nonhuman animals by Shofner in Chapter 3. In Chapter 4, Winter examines in detail the neural representation of periodicity information and describes how and where in the auditory system periodicity information may be processed and extracted. Animal experiments are required for a detailed investigation of neural mecha- ix x Volume Preface nisms. However, it is also possible to observe more general physiologicalpro- cesses in the human auditory system. In Chapter 5, Griffiths explains how modern brain-imaging techniques (PET, fMRI, EEG, and MEG) have enabled researchers to probe the regions responsible for pitch processing in the human brain. InChapter6,deCheveigne´ providesadetailedtaxonomyofpitchmodels using arich historicalandconceptualcontext. Hehighlightsthecommonalities between models and outlines the bases for selecting between them. Pitch per- ception for listeners with hearing impairment and with cochlear implants is discussed in Chapter 7 by Moore and Carlyon. In addition to the clinical ben- efits, such as the design of prostheses, readers of this chapter will be aware of just how much we can learn about “normal” pitch mechanisms by examining the consequences of disrupted auditory processing. In Chapter 8, Darwin considers one ofthe most importantusesofperiodicity information, the segregation of sounds from different sources and the grouping of frequency components from the same source. Finally, in Chapter 9, Bigand andTillmannconsiderwhatmayberegardedas“higher-level”ormorecognitive aspectsofpitch perception,withparticularreferencetotheperceptionofmusic. The chapter topics of this volumehavebeen discussedmorebrieflyandfrom other viewpoints in other volumes of the Springer Handbook of Auditory Re- search series. The psychoacoustics of spectral, temporal, and pitch processing have been presented earlier inVolume 3 (Human Psychophysics). Comparative studies of hearing at the anatomical, physiological, and behavioral levels have been extensively treated in Volumes 4 (Comparative Hearing: Mammals), 11 (Comparative Hearing: Fish and Amphibians), and 13 (Comparative Hearing: Birds and Reptiles). Neurophysiological studies of coding and auditory repre- sentations relevant to pitch perception have been discussed in Volumes 4, 11, and13andinVolumes2(TheMammalianAuditoryPathway:Neurophysiology) and 15 (Integrative Functions in the Mammalian Auditory Pathway). Models of auditory information processing, including pitch, were introduced inVolume 8 (The Cochlea) and more extensively developed in Volume 6 (Auditory Com- putation). More information on pitch perception and the hearing functions of persons with hearing impairments and cochlear implants can be found in Vol- umes7(ClinicalAspectsofHearing)and20(CochlearImplants:AuditoryPros- theses and Electric Hearing). We thank the authors of the chapters for giving so much of their time to the project and for enduring the nagging of the editors. We hope that you agree that the scholarship exhibited isof thehigheststandard. Thevolumewouldnot be what it is without our quality-control team of expert chapter reviewers:Josh Bernstein, John Culling, Alain de Cheveigne´, Steve McAdams, Christophe Micheyl,BrianMoore,AlanPalmer,DanielPressnitzer,andLutzWiegrebe. For no reward, these noble individuals made detailedandconstructivecommentson earlierdrafts,excisingthechaffandinvigoratingthewheat. Wewouldalsolike to express our appreciation to thestaffatSpringer,particularlyJanetSlobodien. Finally, we thank the Hanse Wissenschaftskolleg for facilitating the conference Volume Preface xi that led to this book and for providing financial support in covering the addi- tional cost of the color figures in this volume. Christopher J. Plack, Colchester, United Kingdom Andrew J. Oxenham, Cambridge, Massachusetts Richard R. Fay, Chicago, Illinois Arthur N. Popper, College Park, Maryland Contents Series Preface........................................... vii Volume Preface.......................................... ix Contributors ............................................ xv Chapter 1 Overview: The Present and Future of Pitch ........... 1 Christopher J. Plack and Andew J. Oxenham Chapter 2 The Psychophysics of Pitch....................... 7 Christopher J. Plack and Andrew J. Oxenham Chapter 3 Comparative Aspects of Pitch Perception............. 56 William P. Shofner Chapter 4 The Neurophysiology of Pitch..................... 99 Ian M. Winter Chapter 5 Functional Imaging of Pitch Processing .............. 147 Timothy D. Griffiths Chapter 6 Pitch Perception Models ......................... 169 Alain de Cheveigne´ Chapter 7 Perception of Pitch by People with Cochlear Hearing Loss and by Cochlear Implant Users ................ 234 Brian C.J. Moore and Robert P. Carlyon Chapter 8 Pitch and Auditory Grouping...................... 278 Christopher J. Darwin Chapter 9 Effect of Context on the Perception of Pitch Structures .. 306 Emmanuel Bigand and Barbara Tillmann Index ................................................. 353 xiii
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