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283 Pages·1998·9.115 MB·English
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Central Auditory Processing and Neural Modeling Central Auditory Processing and Neural Modeling Edited by PaulW.F.Poon National Cheng Kung University Tainan, Taiwan and John F. Brugge University of Wisconsin Madison, Wisconsin Springer Science+Business Media, LLC Library of Congress Cataloging-in-Publication Data On file Proceedings of an International Workshop on Central Auditory Processing and Neural Modeling, held January 26-29,1997, in Kaohsiung, Taiwan ISBN 978-1-4613-7441-1 ISBN 978-1-4615-5351-9 (eBook) DOI 10.1007/978-1-4615-5351-9 © 1998 Springer Science+Business Media New York Originally published by PleDIUll Press, New York in 1998 Softcover reprint of the hardcover 1s t edition 1998 http://www.plenum.com 10987654321 AII rights reserved No part ofthis book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilm ing, recording, pr otherwise, without written permission from the Publisher ORGANIZERS * *- m-) Department of Physiology, Medical College, NCKU (~ iL.!&. 7-lJ 1-JX *~ ~ Ministry of Education (~ 11 .g~ ) National Science Council (H JltPit ~ ~H* t-~ -t) CO-ORGANIZERS NCKU Alumni Foundation for Culture and Education, Kaohsiung (~.$!&.7-lJ *-*~ -t ~~*1t-t) Tainan City Government (f; ~ -;p JliJit) NCKU Alumni Association, Malaysia (,~ *-iffIli!&'7-lJ *-*;ft~ xtt*:t--t) Kaohsiung City Government (it;.$ -;p JltJit) Kaohsiung Hsien Government ( iii.$ ~ JltJit ) Cheng Shing Medical Foundation (!&. ~ X tt.* :t--t) Bureau of Health, Kaohsiung City Government (it;.$ -;p JltJit ftf 1-1l] ) Council of Labor Affairs (H JlUit of t--t) Department of Health (H JlUitftj 1-:i) People to People International Tainan Chapter R.O,C ('f"* ~ III ~ ~;~ 3t W, -t f; ~ ~J".) Office of Research and Developement, N*CK U (~iL.!&.7-lJ *-o:f~1t1t~*1t-t) Kaohsiung Polytechnic Institute (~tft.x.. f;t) Wen Tzao Ursuline Junior College of Modern Languages (x~"t! xt-fto!t;ft) The Chinese Physiological Society ('f ~ 1-JX o!t -t) Taiwan Neurological Association (f; J,hff4'.Ho!to!t-t) Kaohsiung Medical College (~t1t"''' Pit) SPONSORS Lin Trading Co. Ltd. (If,j 1+ 1f ~ n~ 17)-t F!t ~ .;J) Eva Air (-k ~Ait 1: ~.;J) Gin Yuan Hotel (1t III *-1&.~) ENDORSED BY * Association for Research in Otolaryngology (~ I'!f.It iJ -t) v LOCAL ADVISORY COMMITTEE (II rIi iiilt-It ) * -·Y'-, ~1f~%~-k if1+ )11 ~*-.M ;l;-ifi;£ ~~iJ.¥,iJ.¥:-k *ft.l ,tit rfi rfi -k ~ ~~iJ a}j i>~rfirfi-k -t ;tlJo ~ iL!&."J-}] :k'd\'.~ti-k .l.7J-=- ~ iL!&."J-}] :k * -I * Fit Fit -k -tILt ~ iL!&."J-}] :k * -I * Fit I!ft -tt -I Fit Fit -k I&.-!,t ~ iL!&."J-}] :k*~*FitFit-k 1-fmH~ ~~~*F;tF;t-k ~a}j~ ~ iL'fIJ~~~ -*#*titi-k 7r .~ o#iIi>j iA iL X. ~ *~ t i X. -*#* titi -k .i. j. 5~K~ . ~ ~ iL!&."J-}] :k Iff -i-ktkt -k -tlAt ~iL!&.~:k*-I*Fit.~~a~~*~~,~1f~.M~.M .:J.f,7j(.f ~ iL!&."J-}] :k*1hfL~lIIff~PJf~~:lPJf-k ~~1/; ~iL!&'~:k*'*~ttIff~~~~:l~-k f~ a}j 1t ~~A~k*.*~#~#~~:li~ FtJl~ ~ iLlWi a}j :k*~f~£#* 'f '~i1:1 **-;~.tt ~~IWia}j:k*~~*Iff~~~~:l~-k 'llt;*1~ ~~*-~!.i.t-l F;t-l*Iff~ .g~i 1:1 1t~*- ~~A"J-}]k*ti~ 'f'~i1:1 -t.£~ A+A.f~~!&':kti~x.~£~~~-k ~~ik A+~.f~~!&.:kti~X.~£~~~-k .:J.f,.£~~ A+A.f~~!&':kti~~~-k *x.~r A+~.f~~!&':ktii.~~-k .l.i~ ~ 'f ~ ~~**~~-'-k,~ iLlWia}j :k*~~*Iff~~~~ 4-,t*- ~ ,tit rfi rfi ifiJit #\ -* ~jb -t 10[ ~iJt ~ ,tit rfi rfi ifiJit .tIJ ~jb -t -k .l.fJJx. ~ ~ rfi rfi ifiJit j:--. ~ifnii 'f*Iff~FitFit± .I..~H~ ~ ~A"J-}] k*;i~fL~;flIff~PJf~~ Itt J;f. ~~A~k*~#\~tt~~ffl~~~ Ft~i! ~~!&'~:k*.*~tt~~ffl~~~ ~i-t~ ~ iL!&."J-}] :k * -I * F;t# ftHt';Ui~ vi PREFACE The full power of combining experiment and theory has yet to be unleashed on studies of the neural mechanisms in the brain involved in acoustic information processing. In recent years, enormous amounts of physiological data have been generated in many laboratories around the world, characterizing electrical responses of neurons to a wide array of acoustic stimuli at all levels of the auditory neuroaxis. Modern approaches of cellular and molecular biology are leading to new understandings of synaptic transmission of acoustic information, while application of modern neuro-anatomical methods is giving us a fairly comprehensive view ofthe bewildering complexity of neural circuitry within and between the major nuclei of the central auditory pathways. Although there is still the need to gather more data at all levels of organization, a ma jor challenge in auditory neuroscience is to develop new frameworks within which existing and future data can be incorporated and unified, and which will guide future laboratory ex perimentation. Here the field can benefit greatly from neural modeling, which in the central auditory system is still in its infancy. Indeed, such an approach is essential if we are to address questions related to perception of complex sounds including human speech, to the many di mensions of spatial hearing, and to the mechanisms that underlie complex acoustico-motor behaviors. The approach of combining experiment and theory has borne much fruit in other systems of the brain, and it was felt that auditory researchers had much to learn from those us ing it. It was also recognized that research on the central auditory system has a great deal to of fer those seeking to understand fundamental mechanisms of sensory processing in the brain. Thus, it seemed that this was the right time to bring together, in the form of a work shop, experimentalists and theoreticians studying the central auditory system and other sen sory systems with the hope and expectation that such a gathering would serve to nurture deeper insight into the fundamental problems before us and to generate new ideas on how fu ture research might accelerate the understanding of auditory cerebral function. We were not disappointed, though only time will tell the degree to which the workshop was a success. This volume is one result of that workshop. We are grateful to Moshe Abeles, Tsutomu Kamada, Gerald Langner, Petr Lansky, and JosefSyka for serving as an advisory group in putting together the workshop program, to Pi-Hsueh Shirley Li, Mei-Ling Tsai, Shun-Sheng Chen, Ling-Ru Lee, Jih-Ing Chuang, and Bu-Miin Huang for serving as the local organizing committee, and to Patrick Heinritz for his valuable editorial assistance in preparing this book. Paul Poon John 8rugge vii CONTENTS Processing of Vocalization Signals in Neurons of the Inferior Colliculus and Medial Geniculate Body ................................................... . Josef Syka. Jiri Popelat. Eugen K vasiHik. and Daniel Suta Coding of FM and AM Sounds at the Auditory Midbrain. . . . . . . . . . . . . . . . . . . . . . .. 13 Paul w.F. Poon. T.w. Chiu. and Xinde Sun Inhibition and Inhibitory Plasticity in the Mammalian Auditory Midbrain .......... 23 Mike B. Cal ford and Yuri Saalmann Neuronal Periodicity Coding and Pitch Effects ... . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 31 Gerald Langner Specializations of the Auditory System for the Analysis of Natural Sounds ......... 43 Israel Nelken. Varon Rotman. and Orner Bar Yosef The Processing of Species-Specific Complex Sounds by the Ascending and Descending Auditory Systems ......................................... 55 Nobuo Suga. Jun Van. and Yunfeng Zhang Speech Recognition System Using Dynamic Programming of Bayesian Neural Networks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 71 Chaug-Ching Huang. Jhing-Fa Wang, Chung-Hsien Wu, and Jau-Yien Lee A Computational Model of Birdsong Learning by Auditory Experience and Auditory Feedback .................................................. 77 Kenji Doya and Terrence 1. Sejnowski On Recent Results in Modeling of Sensory Neurons. . . . . . . . . . . . . . . . . . . . . . . . . . .. 89 Petr Lansky Interneurons Which Shape Response Properties in Dorsal Cochlear Nucleus ........ 101 Eric D. Young and Israel Nelken Behavioral and Physiological Studies of Sound Localization in the Cat. . . . . . . . . . . .. 117 Tom c.T. Yin and Luis C. Populin The Processing of Auditory Stimuli for Eye Movements in the Posterior Parietal Cortex of Monkeys ..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 129 Richard A. Andersen, Alexander Grunewald, and Jennifer F. Linden IX Location Coding by Auditory Cortical Neurons ............................... 139 John C. Middlebrooks Spatial Receptive Field Properties of Primary Auditory Cortical Neurons ........... 149 Richard A. Reale, John F. Brugge, and Joseph E. Hind Models of Direction Estimation with Spherical-Function Approximated Cortical Receptive Fields .................................................... 161 Rick L. Jenison Medical Image Data Compression Using Cubic Convolution Spline Interpolation .... 175 T. K. Truong and Lung-Jen Wang Independent Component Analysis of Electroencephalographic and Event-Related Potential Data ...................................................... 189 Tzyy-Ping Jung, Scott Makeig, Anthony 1. Bell, and Terrence 1. Sejnowski Time Structure of Cortical Activity 199 Yifat Prut and Moshe Abeles Organization and Perturbation of Neuronal Assemblies ......................... 211 George L. Gerstein, Marc J. Bloom, and Pedro E. Maldonado Neural Principles of Visual Cortical Processing: Scene Segmentation Investigated with Microelectrodes and Models ...................................... 225 Reinhard Eckhom Dynamic Mechanisms of Perception Exhibited by Bat Biosonar .................. 247 James A. Simmons, Michael 1. Ferragamo, Tim Haresign, Steven P. Dear, and Mark 1. Sanderson Poster Abstracts of the Workshop on Central Auditory Processing and Neural Modeling .............................................................. 261 Participants' Photographs ................................................. 276 Index ................................................................. 279 x PROCESSING OF VOCALIZATION SIGNALS IN NEURONS OF THE INFERIOR COLLICULUS AND MEDIAL GENICULATE BODY Josef Syka, Jiri PopehU', Eugen Kvasfuik and Daniel Suta Institute of Experimental Medicine Academy of Sciences of the Czech Republic Prague 4, Czech Republic INTRODUCTION Animals vocalize for different reasons: to express their feeling and mood, to emit warning signals, to communicate with individuals of the same species. Vocalization plays a specific role in bats and other echo-locating animals where it serves for the scanning of space. Of the vertebrates, birds in particular are known for their repertoire of vocalizations or singing. Vocalization also plays an important role in the behavior of primates. Undoubtedly an unprecedented and yet unknown process of the perfection and refinement of vocalization in primates resulted in the emergence of a unique feature of the human brain -speech. Although a great deal of knowledge has been accumulated about the production of animal calls and their acoustical features, an understanding of the mechanisms which subserve the processing of these signals in the brain of animals still remains to be elucidated. In the first studies, which were mostly performed in the auditory cortex of awake monkeys in the seventies (Wollberg and Newman, 1972; Winter and Funkenstein, 1973; Newman and Wollberg, 1973; Manley and Muller-Preuss, 1978; Newman, 1978; Newman and Symmes, 1974), the authors expected that they would find neurons utilizing a feature extraction like the neurons in the visual cortex. The existence of specific cells which function as "call detectors" was not confirmed in these studies, and the authors later came to the conclusion that the pattern discrimination of a complex sound can be accomplished by a functional ensemble of neurons (Pelleg-Toiba and Wollberg, 1991). Muller-Preuss and Ploog (1981) demonstrated that many cells in the auditory cortex of the squirrel monkey were inhibited during self-produced vocalizations. New interest in the investigation of the perception of species-specific vocalizations has recently emerged. Rauschecker et al. (1995) studied the responses of neurons in the superior temporal gyrus of anaesthetized rhesus monkeys to complex acoustic stimuli. They found a preference in these neurons for increasingly complex stimuli and suggested that the lateral areas of the monkey auditory cortex may form an important stage in the preprocessing of communication sounds. Wang et al. (1995) found a representation of behaviorally important and spectrotemporally Central Auditory Processing and Neural Modeling Edited by Poon and Brugge, Plenum Press, New York, 1998 complex species-specific vocalizations in the primary auditory cortex of anaesthetized common marmoset and suggested that the representation is carried by dispersed and synchronized cortical cell assemblies that correspond to each individual's vocalization in a specific and abstracted way. In contrast to the large amount of studies which have investigated the role of individual cortical areas in the processing of animal vocalizations, the subcortical nuclei have attracted less attention from this point of view. Creutzfeldt et aI. (1980) studied in unanaesthetized guinea pigs the thalamocortical transformation of responses to complex auditory stimuli. The results of their experiments demonstrated that the responses of medial geniculate body (MGB) cells represent more components of a call than cortical cells, even if the two cells are synaptically connected. Responses ofMGB neurons to species-specific vocalizations were also the subject of a study by Tanaka and Taniguchi (1991), performed in guinea pigs. These authors observed low responsiveness ofMGB neurons to vocalizations; however, the responses to vocalizations displayed discharge patterns which were not possible to predict from the properties of their responses to pure tones. In contrast to this, Buchwald et al. (1988) found that approximately 30% of the units in the caudal part of the cat MGB responded only to species-specific calls, while relatively few responded to a tone or click. Only a few studies were designed to investigate the responses of neurons of the inferior colliculus to vocal stimuli (Aitkin et aI., 1994; Poon and Chiu, 1997). The aim of the study by Aitkin et aI. (1994), performed in anaesthetized cats, was to gain information about the differential coding properties of neurons in three major subdivisions of the inferior colliculus: the central (CNIC) and external (EN) nuclei and dorsal cortex (DC). Feline vocal stimuli were found to be more effective in terms of higher firing rates than white noise or pure tone stimuli at the characteristic frequency (CF) in 27% of the units in the CNIC, 82% in EN and 72% in DC. There were no units that responded exclusively to one vocal stimulus, but a high proportion of units in EN responded strongly to broad band stimuli, and some of these showed clear preferences for one vocal stimulus over another. The aim of the present study was to find out how single cells in the inferior colliculus and in the medial geniculate body of the guinea pig respond to four main types of calls of this species. In addition, we asked the question to what extent could the responses to animal calls be predicted from the responses of the same unit to pure tones at the characteristic frequency (CF) and to white noise. In several animals, we also investigated the responses of two or more neurons recorded in the medial geniculate body with one rnicroelectrode and their interaction with the aid of crosscorrelation technique. METHODS The extracellular responses of IC and MGB neurons to pure tones, white noise and four typical species-specific vocalizations (purr, chutter, chirp and whistle) were recorded in guinea pigs. These calls were selected from a large repertoire of guinea pig calls as the most frequently used. Spontaneous vocalizations were recorded in female pigmented guinea pigs (age 2-24 months) placed in a sound-attenuated room. Recorded calls were analyzed with a high resolution signal analyser B&K 2033 and with aCED 1401plus interface connected to a PC 486 (CED program for frequency analysis Waterfall). Purr is a special call which consists of a bout of regular acoustical impulses with a very small variability of frequency and time parameters. The fundamental frequency of the purr is around 300 Hz. Animals express purr in conjunction with mating behavior and when they seek contact. General exploratory activity is accompanied by a frequently occurring call chutt or chutter, which lasts for 200-300 ms and may appear in a series with variable intercall intervals. The chutter or its parts can sometimes be aperiodic; its fundamental frequency is between 800 and 1500 Hz. The calming of an animal in 2

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