Communication and Cybemetics 8 Editors: W D. Keidel and H. Wolter Facts and Models in Hearing Proceedings of the Symposium on Psychophysical Models and Physiological Facts in Hearing held at Tutzing, Oberbayem, Federa! Republic of Germany, April 22-26, 1974 Edited by E. Zwicker and E. Terhardt Springer-Verlag Berlin Heidelberg NewYork 1974 Volumes I to 7 appeared when the series was called Kommunikation und Kybemetik in Einzeldarstel/ungen Professor Dr.-Ing. Eberhard Zwicker Dr.-Ing. Ernst Terhardt Institut fdr Elektroakustik, Technisehe Universität MÜflchen Symposium was sponsored by the Deutsche Forsehungsgemeinsehqft and the Bayerisehes Staatsministerium for Unterrleht und Kultus With 176 figures ISBN-13: 978-3-642-65904-1 e-ISBN-13: 978-3-642-65902-7 001: 10.1007/978-3-642-65902-7 Library of Congress Cataloging in Publication Data Symposium on Psychophysical Models and Physiological Facts in Hearing, Tutzing, Ger., 1974. Facts and models in hearing. (Communication and cybemetics, v. 8) "Sponsored by the Deutsche Forschungs gemeinschaft and the Bayerisches Staatsministerium fm Unterricht und Kultus." Bibliography: p. 1. Hearing-Congresses. 2. Ear-Congresses. I. Zwicker, Eberhard, ed. 11. Terhardt, E., 1934-ed. III. Deutsche Forschungsgemeinschaft (Founded 1949) IV. Bavaria. Staatsministerium für Unterricht und Kultus. V. Title. [DNLM: 1. Ear-Physiology-Congresses. 2. Hearing-Congresses. 3. Models, Psycho physical-Congresses. WV272 S9895f 19741 QP460.S95 1974 612'.85 74-11221 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is coneemed, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under § 54 of the German Copyright Law, where copies are made for other than private use, a fee is payable to the publisher, the amount of the fee to be determined by agreement with the publisher. © by Springer-Verlag Berlin Heidelberg 1974. Softeover reprint of the hardeover 1st edition 1974 PREFACE During recent years auditory research has advanced quite rapidly in the area of experimental psychology as well as in that of physiology. Scientists working in both areas have in cornrnon the study of the process in HEARING, yet different scientific areas always tend to diverge. A SYMPOSIUM ON PSY CHOPHYSICAL MODELS AND PHYSIOLOGICAL FACTS IN HEARING was or ganized for the exchange of information and to stimulate dis cussion between research workers in psychoacoustics, neurophy siology, anatomy, morphology and hydromechanics. The basic aim of holding this syrnposium was to halt the divergence and to initiate the kind of multi-disciplinary research that will be need ed to elucidate the hearing process as a whole. The present proceedings comprise the papers, which were circulated to the participants two months before the syrnposium and discussed during the syrnposium, together with some cornrnents and additional re marks. These cornrnents and rernarks do not, however, represent the full discussions but only the parts available in written form. We have arranged the material in five sections: I. Structure and Neurobiology of the Inner Ear II. Cochlear Mechanisms III. Auditory Frequency Analysis IV. Auditory Time Analysis V. Nonlinear Effects Within the limits of a syrnposium, none of these topics could be treated comprehensively; moreover, most of the papers concerned problems having several aspects. The volume further contains three papers which were not presented at the syrnposium by their authors: Keidel's paper was read by Kallert; the papers of Engström and of M~ller were not presented orally at all. Our main endeavor was to meet the dernand for rapid publication of up-to-date information. In order to realize this goal, we pre scribed certain specifications and restrictions which, of course, did not always please the authors; we.apologize for this and thank thern for their cooperation. The organization of the symposium and the preparation of this volume would not have been possible without the help of A. Schumann, F. Eberding, H. Fleischer, A. Frei, J. Oelmann, H. SchUtte, D. Schultz, W. Suchowerskyj, and several others. The symposium as well as the pUblication of the proceedings were sponsored by the Deutsche Fo~schungsgemeinschaft and by the Bay~isches Staatsministe~ium fü~ Unte~~icht und Kultus. Finally, we acknowledge the good cooperation with Springer Verlago Mai 1974 Eberhard Zwicker Ernst Terhardt CONTENTS PARTICIPANTS VIII I. STRUCTURE AND NEUROBIOLOGY OF THE INNER EAR 1 H. ENGSTRÖM Morphology of the walls of the cochlear duct 3 C. ANGELBORG H. SPOENDLIN Neuroanatomy of the cochlea 18 R.R. PFEIFFER Comments 33 C.E. MOLNAR J.R. COX, JR. A. FLOCK Neurobiology of ha ir cells and their synapses 37 II. COCHLEAR MECHANISMS 43 V. NEDZELNITSKY Measurements of sound pressure in the cochleae of 45 anesthetized cats P. DALLOS Comments 54 J.P. WILSON Basilar membrane data and their relation to theories of frequency analysis 56 L. ROBLES Comments 64 J. TONNDORF The significance of shearing displacements for the mechanieal stimulation of cochlear hair cells 65 P. DALLOS Comments 75 R. HELLE Enlarged hydromechanical cochlea model with basilar membrane and tectorial membrane 77 M.R. SCHROEDER A model for mechanieal to neural transduction in J.L. HALL the auditory receptor 86 H. DUIFHUIS Comments 94 E. ZWICKER A "second filter" established with in the scala media (General Comment) 95 H. DUIFHUIS An alternative approach to the second filter (General Comment) 100 III. AUDITORY FREQUENCY ANALYSIS 105 J. J. ZWISLOCKI Neuro-mechanical frequency analysis in the cochlea 107 W.G. SOKOLICH E.F. EVANS Auditory frequency selectivity and the cochlear nerve 118 J. SCHWARTZKOPFF Comments 130 E. ZWICKER on a psychoacoustical equivalent of tuning curves 132 VIII L.L.M. VOGTEN Pure-tone masking; a new re sult from a new method 142 R.J. RITSMA Frequeney seleetivity and the tonal residue 156 A. HOEKSTRA B.L.CARDOZO Frequeney diserimination at the threshold 164 G.VAN DEN BRINK Monotie and diehotie piteh matehings with eomplex sounds 178 E. TERHARDT Comments 189 IV. AUDITORY TIME ANALYSIS 191 L.U.E. KOHLLÖFFEL Reeordings from spiral ganglion neurons 193 E.F. EVANS Comments 204 G. BOERGER Coding of repetition noise in the eoehlear nueleus in eat 206 J.P. WILSON Comments 213 F.A. BILSEN Comments 214 W.D. KEIDEL Information proeessing in the higher parts of the auditory pathway 216 A.R. M0LLER Dynamie properties of eoehlear nueleus units in response to exeitory and inhibitory tones 227 A. VOGEL Roughness and its relation to the time-pattern of psyehoaeoustieal exeitation 241 H. FASTL Transient masking pattern of narrow band maskers 251 T. HOUTGAST Masking patterns and lateral inhibition 258 F.A. BILSEN Comments 266 T. HOUTGAST The slopes of masking patterns (General Comments) 269 H. FASTL Comments 273 H. DUIFHUIS A erude quantitative theory of baekward masking 275 V. NONLINEAR EFFECTS L. ROBLES Nonlinear effeets in the transient response of W.S. RHODE the basilar membrane 287 J. P. LEGOUIX Nonlinear meehanisms and eoehlear seleetivity 299 M.C. REMOND P. DALLOS Coehlear mierophonie eorrelates of eubie MARY ANN CHEATHAM differenee tones 312 IX R.R. PFEIFFER The representation of tones and combination C.E. MOLNAR tones in spike discharge pattems of single J.R. COX, JR. cochlear nerve fibers 323 G.F. SMOORENBURG on the mechanisms of combination tone generation and lateral inhibition in hearing 332 R. HELLE Comments 343 H. FASTL E. ZWICKER T.J.F. BUUNEN Subjective phase effects and combination tones 344 F.A. BILSEN E. TERHARDT Pitch of pure tones: its relation to intensity 353 PARTlCIPANTS F.A. Bilsen, Teehnisehe Hogesehool Delft, Lab. voor Natuurkunde, Lorentz weg 1, Delft - 8, Netherland J. Blauert, Institut für Elektrisehe Naehriehtenteehnik der RWTH, 51 Aaehen, Älte Mastriehterstr. 23, West Germany G. Boerger, Heinrieh-Hertz-Institut rur Sehwingungsforschung, 1 Berlin 10, Elnstelnufer 37, West Germany G. van den Brink, Faculteit der Geneeskunde, Erasmus Universiteit Rotterdam, Lab.f. Blolog.a.Medicial Physics, P.O. Box 1738, Rotterdam, Netherland B.L. Cardozo, Instituut voor Perceptie Onderzoek, Insulindelaan 2, Elndhoven, Netherland P.J. Dallos, Auditory Research Laboratory, Northwestern University, 2299 Sherldan Road, Evanston, Illinois, 60201, U.S.A. H. Duifhuis, Instituut voor Perceptie Onderzoek, Insulindelaan 2, Eindhoven, Netherland E.F. Evans, Department of Communication, University of Keele, Keele, Staffordshire, ST5 5BG, Great Britain H. Fastl, Institut für Elektroakustik der Techn. Universität München, Ärclsstr. 21, 0-8 München 2, West Germany A. Flock, Konung Gustaf V forskningsinstitut, S-104 01 Stockholm 60, Sweden C.A.A.J. Greebe, Instituut voor Perceptie Onderzoek, Insulindelaan 2, Eindhoven, Netherland R. Helle, Institut für Elektroakustik der Techn. Universität München, Arcisstr. 21, 0-8 München 2, West Germany T. Houtgast, Institute for Perception TNO, Kampweg 5, Postbus 23, Soesterberg, Netherland Mrs. John, Deutsche Forschungsgemeinschaft, 53 Bonn-Bad Godesberg, Kenneayallee 40, West Germany H.R. de Jongh, Wilhelmina Gasthuis, Academisch Ziekenhuis bij de Universiteit van Amsterdam, ENT- Department, Amsterdam-Oud west, Netherland S. Kallert, I. Physiologisches Institut der Universität Erlangen-Nürnberg, 852 Erlangen, Universitätsstr. 17, West Germany L.U.E. Kohllöffel, I. Pfiysiologisches Instftut der Universität Erlangen Nurnberg, 852 Erlangen, Universitätsstr. 17, West Germany J.P. Legouix, Laboratoire de Neurophysiologie Generale, College de France, 11 Place Marcelin-Berthelot, Paris-50, France V. Nedzelnitsky, Eaton-Peabody Laboratory of Auditory Physiology, 42 Carlton Street, Cambridge, Massachusetts, 02142, U.S.A. R.R. Pfeiffer, Department of Electrical Engineering, Washington University, St. LOU1S, Mlssouri, 63130, U.S.A. R. Plomp, Institute for Perception TNO, Kampweg 5, Postbus 23, Soesterberg, Netherland R.J. Ritsma, Academisch Ziekenhuis Groningen Kliniek voor Keel-Neus-en öorheelkunde, Oostersingel 59, Groningen, Netherland L. Robles, Laboratory of Neurophysiology Medical School, University of Wlsconsln, 283 Medical Sciences Building, Madison, Wisconsin, 53706, U.S.A. J.F. Schouten, Parklaan 36, Eindhoven, Netherland M.R. Schröder, Direktor des Dritten Physik. Instituts der Universität Gottlngen, Bürgerstr. 42-44, West Germany J. Schwartzkopff, Institut für Allgemeine Zoologie der Ruhr-Universität Bochum, 463 Bochum-Querenburg, Postfach 2148, West Germany G.F. Smoorenburg, Institute for Perception TNO, Kampweg 5, Postbus 23, Soesterberg, Netherland H. Spoendlin, Kantonsspital Zürich, Rämistr. 100 CH-8006 Zürich, Switzerland E. Terhardt, Institut für Elektroakustik der Techn. Universität München, Ärcisstr. 21, 0-8 München 2, West Germany J. Tonndorf, Department of Otolaryngology, College of Physicians and Surgeons of Columbia University, 630 West 168th Street, New York 32, New York, U.S.A. A. Vogel, Institut für Elektroakustik der Techn. Universität München, Ärclsstr. 21, 0-8 München 2, West Germany Vogten, Instituut voor Perceptie Onderzoek, Insulindelaan 2, L.L.~ Eindhoven, Netherland J.P. Wilson, Department of Communication, University of Keele, Keele, Staffordshlre ST5 5BG, Great Britain E. Zwicker, Institut für Elektroakustik der Techn. Universität München, Ärclsstr. 21, 0-8 München 2, West Germany J.J. Zwislocki, Lab. of Sensory Communication, Syracuse University, 821 Unlverslty Ävenue, Syracuse, New York, 13210, U.S.A.
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