WHAT FIRE IS IN MINE EARS:

PROGRESS IN AUDITORY

BIOMECHANICS

Proceedings of the 11th International Mechanics

of Hearing Workshop

Williomstown, Massachusetts 16-22 July 2011

EDITORS

Christopher A. Shera

Harvard University Cambridge, MA

Elizabeth S. OlsonColumbia University, New York, NY

All papers have been peer reviewed.

Melville, New York, 2011

AIP I CONFERENCE PROCEEDINGS 1403

AIP Conference Proceedings, Volume 1403

What Fire is in Mine Ears: Progress in Auditory Biomechanics

Proceedings of the 11th International Mechanics of Hearing

Workshop

Table of Contents

Preface: What Fire is in Mine Ears: Progress in Auditory Biomechanics

Christopher A. Shera and Elizabeth S. Olson

Acknowledgments

Previous Mechanics of Hearing Workshops

MECHANICS OF HEARING 101

Moll 101: Basic concepts in the mechanics of hearing

Christopher Bergevin, Bastian Epp, and Sebastiaan W. F. Meenderink

HAIR CELLS: BUNDLES, TUNING, TRANSDUCTION

Damping properties of the hair bundle

Johannes Baumgart, Andrei S. Kozlev, Thomas Risler, and A. J.

Hudspeth

New devices for investigating hair cell mechanical propertiesJoseph C. Doll, Anthony Peng, Anthony Ricci, and Beth L. Pruitt

Hair bundle dynamics under steady-state deflection

Lea Fredrickson-Hemsing, Seung Ji, Robjin Bruinsma, and Dolores

Bozovic

Sound-evoked length changes of the outer hair cell stereocilia bundle are

modulated by endocochlear currents

Pierre Hakizimana, William E. Brownell, Stefan Jacob, and Anders

Fridberger

Effects of electrical and mechanical overstimulus on spontaneous

oscillations in hair bundles

Albert Kao, C. Elliott Strimbu, and Dolores Bozovic

Horizontal top connectors mediate a sliding adhesion to hair cell

stereocilia

K. D. Karavitaki and D. P. Corey 44

Elastostatic analysis of the membrane tenting deformation of inner-ear

stereocilia

Jichul Kim, Peter M. Pinsky, Anthony J. Ricci, Sunil Puria, and Charles

R. Steele 50

Dynamic aspects of cochlear microphonic potentialsSebastiaan W. F. Meenderink and Marcel van der Heijden 53

High-pass filtering at vestibular frequencies by transducer adaptation in

mammalian saccular hair cells

Jocelyn E. Songer and Ruth Anne Eatock 59

Molecular mechanics of tip-link cadhcrins

Marcos Sotomayor, Wilhehn A. Weihofen, Rachelle Gaudet, and David

P. Corey 64

Exploring the role of mcchanotransduction activation and adaptationkinetics in hair cell filtering using a Hodgkin-Huxlcy approach

Gregg B. Wells and Anthony J. Ricci 70

MECHANICAL COUPLING AND EXCITATION OF HAIR CELLS

Time domina one-dimensional cochlear model with integrated tectorial

membrane and outer hair cells

Oded Barzelay and Miriam Furst 79

Tectorial membrane traveling waves underlie impaired hearing in Tectb

mutant mice

Roozbeh Ghaff'ari, Shirin Farrahi, Alexander J. Aranyosi, Guy P.

Richardson, and Dennis M. Freeman 85

Mechanical excitation of 1HC stereocilia: An attempt to fit togetherdiverse evidence

John J. Guinan, Jr. 90

Unraveling traveling waves using WKB modelingJessica S. Lamb and Richard S. Chadwick 97

Coupling the subtectorial fluid with the tectorial membrane and hair

bundles of the cochlea

Yizeng Li, Julien Meaud, and Karl Grosh

The effect of superior semicircular canal dehiscence on intracochlear

sound pressuresHideko Heidi Nakajima, Dominic V. Pisano, Saumil N. Merchant, and

John J. Rosowski

Biophysical mechanisms underlying hearing loss associated with a

shortened tectorial membrane

John S. Oghalai, Anping Xia, Christopher C. Liu, Simon S. Gao, Brian E.

Applegate, Sunil Puria, Hay Rousso, and Charles Steele

Numerical study of the complex temporal pattern of spontaneousoscillation in bullfrog saccular hair cells

Yuttana Roongthumskul, Lea Fredrickson-Hemsing, Albert Kao, and

Dolores Bozovic

Magnetic bead actuation of saccular hair cells

David Rowland, Damien Ramunno-Johnson, Jae-Hyun Lee, Jinwoo

Cheon, and Dolores Bozovic

Active motion of hair bundles coupled to the otolithic membrane in the

frog sacculus

C. Elliott Strimbu, Lea Fredrickson-Hemsing, and Dolores Bozovic

Otoancorin knockout mice reveal inertia is the force for hearingThomas Weddell, P. Kevin Legan, Victoria A. Lukashkina, Richard J.

Goodyear, Lindsy Welstead, Chisline Petit, lan J. Russell, Andrei N.

Lukashkin, and Guy P. Richardson

OUTER HAIR CELLS AND PRESTIN

Evaluating prestin's changing biophysical attributes in development

using a tet-induced cell line

Shumin Bian, Bon W. Koo, Stephen Kelleher, Joseph Santos-Sacchi, and

Dhasakumar Navaratnam

Decreasing outer hair cell membrane cholesterol increases cochlear

electromechanics

William E. Brownell, Stefan Jacob, Pierre Hakizimana, Mats Ulfendahl,

and Anders Fridberger

Homo- and hetero-oligomerization in the Slc26a protein family

Benjamin Currall, Heather Jensen-Smith, and Richard Hallworth 154

Continuum mechanical model of the outer hair cell

Mario Fleischer, Csaba Harasztosi, Manuela Nowotny, Thomas Zahnert,

and Anthony W. Glimmer 160

The evolution of prestin: Examination with membrane thickness

sensitivityChisako Izumi, Jonathan Bird, Torsten Schweninger, Dominik Oliver, and

Kuni H. Iwasa 166

Analysis of membrane topology of prestin expressing in CHO cells

Michio Murakoshi, Tomohiro Kawase, Shun Knmano, and Hiroshi VVada 168

A cochlear partition model incorporating realistic electrical and

mechanical parameters for outer hair cells

Jong-Hoon Nam and Robert Fetliplace 170

Outer hair cell electromotility /// vivo

Sripriya Ramamoorthy and Alfred L. Nuttall 176

Chloride dependent coupling of molecular to cellular mechanics in the

outer hair cell of Corti's organLei Song and Joseph Santos-Sacchi 179

High-resolution AFM imaging of prestin purified and reconstituted into

an artificial lipid bilayerHiroshi Wada, Shun Kumano, Michio Murakoshi, Koji lida, and Hiroshi

Hamana 185

Normal hearing sensitivity at low to middle frequencies with -25%

prestinJian Zuo, Tetsuji Yamashita, Jie Fang, Jiangang Gao, Yiling Yu, andMarcia Mellado-Lagarde 191

NONLINEAR1TY IN THE COCHLEA

Hopf-bifurcations and Van der Pol oscillator models of the mammalian

cochlea

Hendrikus Duifhuis 199

Cochlear fine structure—Implications for modulation processing at the

level of the cochlea

Bastian Epp, Manfred Mauennann, and Jesko L. Verhey

Modeling scala media as a pressure vessel

Eric LePage and Ake Olofsson

Quasi-linear cochlear responses to noise can result from instantaneous

nonlincarities

Yi-Wen Liu and Stephen T. Neely

A pole-zero filter cascade provides good fits to human masking data and

to basilar membrane and neural data

Richard F. Lyon

The generation of harmonic distortion and distortion products in a

computational model of the cochlea

Julien Meaud, Yizeng Li, and Karl Grosh

Medial cochlear efferent function: A theoretical analysisDavid C. Mountain

Estimation of psychophysical thresholds based on neural network

analysis of DPOAE input/output functions

Maryam Naghibolhosseini and Glenis Long

On modeling nonlinearity, longitudinal coupling, and spatialinhomogenity in the cochlea

Robert Szalai, Nigel P. Cooper, Alan R. Champneys, and Martin Homer

Can a static nonlinearity account for the dynamics of otoacoustic

emission suppression?Sarah Verhulst, Christopher A. Shera, James M. Harte, and Torsten Dau

OTOACOUSTIC EMISSIONS

Accuracy of noninvasive estimation techniques for the state of the

cochlear amplifierErnst Dalhoff and Anthony W. dimmer

Generation of distortion product otoacoustic emissions in the gerbil

cochlea

Wei Dong and Elizabeth S. Olson 273

Delays and growth rates of multiple TEOAE componentsShawn S. Goodman, Ian B. Merles, and Rachel A. Scheperle 279

Otoacoustic estimates of cochlear tuning: Testing predictions in macaque

Christopher A. Shera, Christopher Bergevin, Radha Kalluri,

Myles Me Laughlin, Pascal Michelet, Marcel Van der Heijden,and Philip X. Joris 286

High frequency click-evoked otoacoustic emission measurements usingmaximum length sequences

B. Linelon, R. Sohal, E. Harvey, and A. R. D. Thornton 293

Modification of DPOAE fine structure stemming from changes in outer

and middle ear function

Glenis R. Long, Simon Henin, and Suzanne Thompson 299

Stimulus frequency otoacoustic emissions in wild-type and TECTA mice

(Carina N, Pikhart, Gerald R. Popelka, Arturo Moleti, Renata Sisto, John

S. Oghalai, Anping Xia, and Sunil Puria 305

Transient-and tone-evoked otoacoustic emissions in three speciesJ. H. Siegel, K. Charaziak, and M. A. Cheatham 307

Investigating the origin of upper-side-band distortion-productotoacoustic emissions within a micromechanical cochlear model

J. Young, S. J. Elliott, and B. Linelon 315

COCHLEAR ARCHITECTURE: MEASUREMENTS AND MODELING

A unified numerical method for fluid-structure interaction applied to

human cochlear mechanics

Frank Bohnke and Daniel Roster 323

Fluid coupling in a discrete cochlear model

S. J. Elliott, B. Linelon, and G. Ni 328

Mechanics of the unusual basilar membrane in gerbilSanlosh Kapuria, Charles R. Steele, and Sunil Puria 333

Bone conduction hearing: Three-dimensional finite element model of the

human middle and inner ear

Namkeun Kim, Kenji Homma, Sunil Puria, and Charles R. Steele 340

Determining the orthotropic properties of gerbil basilar membrane from

space constant measurements

Shuangqin Liu and Robert White 346

Finite element modelling of fluid coupling in the coiled cochlea

Guangjian Ni, S. J. Elliott, B. Linelon, and R. Saba 350

Three-dimensional imaging of the mouse organ of Corti cytoarchitecturefor mechanical modeling

Sunil Puria, Byron Hartman, Jichul Kim, John S. Oghalai, Anthony J.

Ricci, and M. Charles Liberman 356

A morphological theory of human hearingPaolo Pamieri 363

A physiological signal transmission model to be used for specificdiagnosis of cochlear impairments

Amin Sareini and Stefan Stenfelt 369

Coiled hydromechanical scale model of the inner ear

Bonita Tam, Arielle Fakhraee, and Robert D. White 374

COCHLEAR MACRO- AND MICROMECHANICS

Using stimulus frequency emissions to characterize cochlear function in

mice

M. A. Cheatham, E. D. Katz, K. Charaziak, P. Dallos, and J. H. Siegel 383

Vibration measurement on reticular lamina and basilar membrane at

multiple longitudinal locations

Fangyi Chen, Dingjun Zha, Niloy Choudhury, Anders Fridberger, and

Alfred L. Nultall 389

Imaging organ of Corti vibration using Fourier-domain OCT

Niloy Choudhury, Fangyi Chen, Anders Fridberger, Dingjun Zha, Steven

L. Jacques, Ruikang K. Wang, and Alfred L. Nultall 391

Efferent insights into cochlear mechanics

Nigel P. Cooper and John J. Guinan, Jr. 396

Lowered pH alters decay but not speed of tectorial membrane waves

Shirin Farrahi, Roozbeh Ghaffari, and Dennis M. Freeman 403

Dissection of the mechanical impedance components of the outer hair

cell using a chloride-channel blocker

Csaba Harasztosi and Anthony W. Glimmer 405

Active outer hair cells affect the sound-evoked vibration of the reticular

lamina

Stefan Jacob and Anders Fridberger 411

Laser interferometer measurements of the viscoelastic properties of

tectorial membrane mutants

Gareth Jones, Ian Russell, and Andrei Lukashkin 419

Mechanisms of cochlear stimulation through the round window

Andrei N. Lukashkin, Thomas Weddell, and Ian J. Russell 421

Measurement of basilar membrane, reticular lamina, and tectorial

membrane vibrations in the intact mouse cochlea

Tianying Ren and Wenxuan He 423

Direct measurement of basilar membrane motion using pulsed-wave

doppler high-frequency ultrasound

Z. Torbatian, P. Garland, R. B. A. Adamson, M. Bance, and J. A. Brown 430

Basilar membrane measurements from wild type, prestin 499, and

prestin KO mice

Thomas Weddell, Marcia Mellado-Lagarde, Victoria Lukashkina, Andrei

Lukashkin, Jian Zuo, and Ian Russell 432

Acoustically evoked different vibration pattern across the width of the

cochlea partition

Dingjun Zha, Fangyi Chen, Anders Friderberg, Niloy Choudhury, and

Alfred Nuttall 434

Characterizing wave propagation in the organ of Corti with stroboscopicimaging

Aleks Zosuls, Laura C. Rupprecht, and David C, Mountain 438

COMPARATIVE AUDITORY MECHANICS

Modelling the active hearing process in mosquitoesDaniele Avitabile, Martin Homer, Joe Jackson, Daniel Robert, and Alan

Champneys

Coupled, active oscillators and lizard otoacoustic emissions

Christopher Bergevin, David S. Velenovsky, and Kevin E. Bonine

Sound transduction in the auditory system of bushcrickets

Manuela Nowotny, Arun Palghat Udayashankar, Melanie Weber, Jennifer

Hummel, and Manfred Kossl

Tonotopically ordered traveling waves in the hearing organs of

bushcrickets in-vivo

Arun Palghat Udayashankar, Manfred Kossl, and Manuela Nowotny

Biornechanical analysis of hearing in whales using nanoindentation and

the finite element method

Andrew A. Tubelli, Aleks Zosuls, Darlene R. Ketten, and David C.

Mountain

Mosquitoes on the wing "tune in" to acoustic distortion

Ben Warren and Ian Russell

HERE BE DRAGONS: MECHANICS IN THE APEX

Deviations from scaling symmetry in the apical half of the human

cochlea

Carolina Abdala, Sumitrajit Dhar, and Radha Kalluri

Suppression of auditory-nerve-fiber responses to off-CF tones

Maria A. Berezina and John J. Guinan, Jr.

The effect of the helicotrema on low-frequency cochlear mechanics and

hearingTorsten Marquardt and Carlos Jurado

Auditory-nerve responses to clicks at low levels, and the initial peak at

high levels, are suppressed at opposite bias-tone phasesHui Nam and John J. Guinan, Jr.

Unidirectional amplification as a mechanism for low-frequency hearingin mammals

Tobias Reichenbach and A. J. Hudspeth 507

EXTERNAL AND MIDDLE EARS

Numerical analysis of the influence of the auditory external canal

geometry on the human hearing response

Luis Caminos, Antonio Garcia-Gonzalez, and Antonio Gonzalez-Herrera 515

The tympanic membrane motion in forward and reverse middle-ear

sound transmission

Jeffrey Tao Cheng, Ellery Harrington, Rachelle Horwitz, Cosine

Furlong, and John J. Rosowski 521

Measurement of the three-dimensional vibration motion of the ossicular

chain in the living gerbilWillem F. Decraemer, Ombeline de La Rochefoucauld, and Elizabeth S.

Olson 528

Spatial motion in natural and reconstructed middle ears and the impacton sound transfer

Albrecht Eiber, Christoph Heckeler, Michael Lauxmann, Hannes Maier,

and Mohammed Saffarini 534

Modeling analysis of biomcchanical changes of middle ear and cochlea in

otitis media

Rong Z. Can, Xiangming Zhang, and Xiying Guan 539

Modeling ear-canal acoustics, incorporating visco-thermal effects and

the influence of the middle ear

Lauren E. Gowdy and Robert H. Withnell 545

Towards creation of a human-head auditory model for simulating bone-

conduction pathwaysKenji Homma, Namkeun Kim, and Sunil Puria 552

Comparison of ear-canal reflectance and umbo velocity in patients with

conductive hearing loss

Gabrielle R. Merchant, Hideko H. Nakajima, Dominic V. Pisano, Chrislof

Roosli, Mohamad A. Hamade, Lorice Mafoud, Christopher F. Halpin,Saumil N. Merchant, and John J. Rosowski 554

WAVE PROPAGATION IN THE COCHLEA

Tracing distortion product (DP) waves in a cochlear model

Egbert de Boer, Christopher A. Shera, and Alfred L. Nuttall 557

How many waves propagate in the cochlea?

S. J. Elliott, G. Ni, B. R. Mace, and B. Lineton 563

Harmonic response of the organ of Corti: Results for wave dispersionSimon Foucaud, Guilhem Michon, Joseph Morlier, and Yves Gourinat 569

Time-domain representation of active nonlinear cochlear waves

Florian Fruth, Kai Dierkes, Benjamin Lindner, and Frank Jiilicher 576

An active, reflectionless transmission-line model of the cochlea: Revisited

Tohru Kohda, Takao Une, and Kazuyuki Aihara 578

Forward- and reverse-traveling waves in DP phenomenology: Does

inverted direction of wave propagation occur in classical models?

Renala Sisto, Christopher A. Shera, Arluro Moleti, and Teresa Bolti 584

Retrograde propagation mechanisms of Oaes: Slow-wave interpretationof the Ren et al. experiments

Ales Vetesnik and Anthony W. Gummer 590

THE COCHLEAR AMPLIFIER

Stiffness vs damping in the cochlea: A negative conclusion?

Andrew J. Binder, A. J. Aranyosi, and Christopher Bergevin 595

Cochlear gain control estimated from distortion product otoacoustic

emissions evoked by amplitude modulated tones

Shixiong Chen and Lin Bian 601

Simulation of fluid flow and basilar-membrane motion in a two-

dimensional box model of the cochlea

Elisabeth Edom, Dominik Obrist, and Leonhard Kleiser 608

A mixed mode cochlear amplifier including neural feedback

Matthew R, Flax and W. Harvey Holmes 613

The cascaded cochlea

Marcel van tier Heijden 618

Comparing longitudinal coupling and temporal delay in a transmission-

line model of the cochlea

Martin Homer, Robert Szalai, Alan Champneys, and Baslian Epp 625

How much do somatic and hair bundle motility contribute to cochlear

amplification?Duk Joong Kim, David C. Mountain, and Allyn E. Hubbard 632

Simultaneous measurements of pressure and voltage at the basilar

membrane inform theories of amplificationElizabeth S. Olson, Wei Dong, and Stephen T. Neely 638

High-frequency power gain in the mammalian cochlea

Daibhid 6. Maoileidigh and A. j. Hudspeth 645

An electromechanical model for the cochlear microphonicPaul D. Teal, Ben Lineton, and Stephen J, Elliott 652

Can outer hair cells actively pump fluid into the tunnel of Corti?

Brissi Franck Zagadou and David C. Mountain 658

MODERATED DISCUSSION SESSIONS

Hair cells: Bundles, tuning, transduction—A moderated discussion

K. Domenica Karavitaki and Anthony J. Ricci 667

Mechanical coupling and excitation of hair cells—A moderated

discussion

Mary Ann Cheatham and Dennis M. Freeman 673

Outer hair cells and prestin—A moderated discussion

William E. Brownell and Anthony W. Gummer 678

Nonlinearity in the cochlea—Moderated discussions

David C. Mountain and John J. Guinan, Jr. 684

Otoacoustic emissions and cochlear architecture: Measurements and

modeling—A discussion

Stephen T. Neely and Sunil Puria 692

Cochlear macro- and micromechanics—A moderated discussion

Wei Dong and Mario A. Ruggero 695

Comparative auditory mechanics: From species to species and from base

to apex—A moderated discussion

Nigel P. Cooper and Geoffrey A. Manley 701

Wave propagation and amplification in the cochlea—A moderated

discussion

Richard S. Chadwick and Karl Grosh 707

Mechanics of hearing potpourri—A moderated discussion

John J. Guinan, Jr. 710

Workshop Participants 715

Author Index 719