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Neural Coding and Perception of Pitch in the Normal and Impaired Human Auditory System

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Neural Coding and Perception of Pitch in the Normal and Impaired Human Auditory System Downloaded from orbit.dtu.dk on: Oct 05, 2021 Neural coding and perception of pitch in the normal and impaired human auditory system Santurette, Sébastien Publication date: 2011 Document Version Publisher's PDF, also known as Version of record Link back to DTU Orbit Citation (APA): Santurette, S. (2011). Neural coding and perception of pitch in the normal and impaired human auditory system. Contributions to hearing research No. 10 General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. Users may download and print one copy of any publication from the public portal for the purpose of private study or research. You may not further distribute the material or use it for any profit-making activity or commercial gain You may freely distribute the URL identifying the publication in the public portal If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. i “phd_thesis_book” — 2011/10/11 — 13:22 — page -1 — #1 i i i CONTRIBUTIONS TO HEARING RESEARCH Volume 10 Sébastien Santurette Neural coding and perception of pitch in the normal and impaired human auditory system i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page 0 — #2 i i i i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page i — #3 i i i Neural coding and perception of pitch in the normal and impaired human auditory system PhD thesis by Sébastien Santurette Technical University of Denmark 2011 i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page ii — #4 i i i © Sébastien Santurette, 2011 Cover illustration: Earamin by Emily Allen. Printed in Denmark by Rosendahls - Schultz Grafisk A/S. The defense was held on June 17, 2011. i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page iii — #5 i i i This PhD-dissertation is the result of a research project at the Centre for Applied Hearing Research, Department of Electrical Engineering, Technical University of Denmark (Kgs. Lyngby, Denmark). Parts of the project were carried out at the Divi- sion of Experimental Otorhinolaryngology, Department of Neurosciences, Katholieke Universiteit Leuven (Leuven, Belgium), and at the Auditory Perception and Cognition Lab, Department of Psychology, University of Minnesota (Minneapolis, USA). The project was financed by a DTU stipend. The external subprojects were supported by grants from the Fund for Scientific Research Flanders (K.U. Leuven, G.0331.08), the Research Council of Katholieke Universiteit Leuven (K.U. Leuven, OT/07/034), and the U.S. National Institute of Health (U of M, R01 DC 05216). The external stay at the University of Minnesota was further supported by travel grants from the Otto Mønsted Fund and the Oticon Foundation. Supervisors Main supervisor Co-supervisor Prof. Torsten Dau Ass. Prof. Jörg M. Buchholz Centre for Applied Hearing Research Centre for Applied Hearing Research Department of Electrical Engineering Department of Electrical Engineering Technical University of Denmark Technical University of Denmark Kgs. Lyngby, Denmark Kgs. Lyngby, Denmark External advisors Prof. Jan Wouters Prof. Andrew J. Oxenham Division of Experimental ORL Auditory Perception and Cognition Lab Department of Neurosciences Department of Psychology Katholieke Universiteit Leuven University of Minnesota Leuven, Belgium Minneapolis, MN, USA i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page iv — #6 i i i iv i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page v — #7 i i i Abstract Pitch is an important attribute of hearing that allows us to perceive the musical quality of sounds. Besides music perception, pitch contributes to speech communication, auditory grouping, and perceptual segregation of sound sources. In this work, several aspects of pitch perception in humans were investigated using psychophysical methods. First, hearing loss was found to affect the perception of binaural pitch, a pitch sensation created by the binaural interaction of noise stimuli. Specifically, listeners without binaural pitch sensation showed signs of retrocochlear disorders. Despite adverse effects of reduced frequency selectivity on binaural pitch perception, the ability to accurately process the temporal fine structure (TFS) of sounds at the output of the cochlear filters was found to be essential for perceiving binaural pitch. Monaural TFS processing also played a major and independent role for a variety of basic auditory tasks, indicating that it may be a crucial measure to consider for hearing-loss characterization. In contrast to hearing-impaired listeners, adults with dyslexia showed no deficits in binaural pitch perception, suggesting intact low-level auditory mechanisms. The second part of this work investigated the role of temporal and spectral information for complex pitch perception. In particular, it was shown that the low pitch evoked by high-frequency complex tones was not conveyed by temporal- envelope cues as such. Moreover, the fact that the individual frequency components could not be heard out separately by the listeners suggested that the low pitch relied on TFS information, even in high-frequency regions where phase-locking in auditory- nerve cells is believed to be weak. A second set of experiments could however not validate the assumption of a temporally-coded pitch and indicated that the use of spectral cues remained plausible. Simulations of auditory-nerve representations of the complex tones further suggested that a spectrotemporal mechanism combining precise timing information across auditory channels might best account for the behavioral data. Overall, this work provides insights into the fundamental auditory mechanisms underlying pitch perception, and may have implications for future pitch-perception models, as well as strategies for auditory-profile characterization and restoration of accurate pitch perception in impaired hearing. v i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page vi — #8 i i i i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page vii — #9 i i i Resumé Tonehøjde er en vigtig egenskab ved hørelsen, der giver os mulighed for at opfatte musikalsk kvalitet af lyde. Udover opfattelse af musik, bidrager tonehøjde til talekommunikation, auditiv gruppering, og perceptuel adskillelse af lydkilder. I denne afhandling blev adskillige aspekter af tonehøjde i det menneskelige auditive system undersøgt ved hjælp af psykofysiske forsøg. Først sås det at høretab påvirker opfattelsen af binaural pitch, en opfattelse af tonehøjde skabt af den binaurale interaktion af støjstimuli. Især viste individer uden binaural-pitch opfattelse tegn på retrocochlear lidelser. Evnen til præcist at bearbejde lydes temporale finstruktur (TFS) ved udgangen af det indre øres filtre sås at være af afgørende betydning også selv om de negative effekter ved reduceret frekvensselektivitet tages i betragtning. Bearbejdning af den monaurale TFS sås også at spille en vigtig og selvstændig rolle for en række grundlæggende auditive funktioner, hvilket tyder på, at denne evne kunne være en vigtig størrelse at måle for karakterisering af høretab. I modsætning til hørehæmmede viste personer med ordblindhed intet besvær med at opfatte binaural pitch, hvilket antyder intakte lavniveau auditive mekanismer. Den anden del af dette arbejde undersøgte rollen af temporal og spektral information for komplekse toners tonehøjde. Mest bemærkelsesværdigt blev det påvist, at tonehøjden for tonekomplekser med høje frekvenser ikke blev bestemt ud fra deres temporale indhylningskurve. Desuden kunne de enkelte frekvenskomponenter ikke adskilles, hvilket antyder at tonehøjden bestemmes fra TFS oplysninger, selv i højfrekvens regioner, hvor synkronien i auditive nerveceller menes at være svag. Andre forsøg kunne dog ikke bekræfte antagelsen om en temporalt kodet tonehøjde og viste, at muligheden for spektral kodning foreligger. Simuleringer af stimuliene i den auditive nerve tydede endvidere på, at en spektrotemporal mekanisme, der kombinerer den temporale information på tværs af auditive kanaler, muligvis bedst stemmer overens med de adfærdsmæssige data. Samlet set giver dette arbejde indsigt i de grundlæggende auditive mekanismer bag opfattelsen af tonehøjde, og kan have betydning for fremtidige auditive modeller, samt strategier for karakterisering af den auditive profil og retablering af tonehøjde opfattelse hos personer med nedsat hørelse. vii i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page viii — #10 i i i i i i i i “phd_thesis_book” — 2011/10/11 — 13:22 — page ix — #11 i i i Résumé La hauteur tonale est un élément important de l’audition nous permettant de percevoir la qualité musicale des sons. Outre la perception musicale, la hauteur facilite la communication orale, le groupement auditif, et la ségrégation de sources sonores. Par le biais d’expériences psychophysiques, cette thèse étudie plusieurs aspects de la perception de la hauteur dans le système auditif humain. Tout d’abord, il est démontré que la perte d’audition affecte la perception du pitch binaural, une sensation de hauteur tonale créée par l’interaction binaurale de deux bruits. Plus précisément, des signes de troubles rétrocochléaires sont trouvés chez les sujets sans sensation de pitch binaural. Bien que la perception du pitch binaural soit affectée par une sélectivité en fréquence réduite, la
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