Modeling the nonlinear active cochlea: Mathematics and analog VLSI

MODELING THE NONLINEAR ACTIVE COCHLEA: MATHEMATICS AND ANALOG VLSI Bo Wen Supervisor: Kwabena A. Boahen The human auditory system vastly outperforms any machine in efficiency and robustness in perceiving sound. Complex and delicate, its front end, namely the cochlea, senses and processes sound in a nonlinear active fashion, exhibiting remarkable sensitivity and extraordinary frequency discrimination. The mechanism through which the mammalian cochlea achieves its incredible capability is postulated as the cochlear amplifier, which is generally believed to originate from outer hair cells electromotility. However, the detail of the cochlear amplifier remains unclear. Using both analytic and synthetic approaches, the present work provided a plausible basis for realization of the cochlear amplifier, thus advancing our understanding of cochlear mechanics. We proposed a novel cochlear amplifier mechanism based on the cochlea’s microanatomy, as well as outer hair cell (OHC) motility, to account for the cochlea’s characteristic behavior. This mechanism, active bidirectional coupling (ABC), considers both the basal tilt of OHCs and the apical tilt of phalangeal processes as critical for feeding OHC motile forces (with saturating property) forward and backward onto the basilar membrane, thereby enhancing the cochlea’s functioning. The ABC-based mathematical cochlear model produces responses that are comparable to physiological measurements. Theoretical model analysis reveals that ABC leads to negative damping basal to the response peak over a small longitudinal cochlear region. The tilted structure works as a v spatial filter, amplifying the cochlear traveling wave only when its wavelength becomes comparable to the tilt distance. Inspired by the biology, we proceeded to build a nonlinear active silicon cochlea—a very large scale integration (VLSI) physical cochlear model—for achieving real-time lowpower cochlear processing. Designed in current mode and operating in Class AB, this microchip implements the ABC mechanism, together with a silicon auditory nerve, in 0.25 μm complementary metal-oxide-semiconductor (CMOS) technology. The resultant new architecture addresses the shortcomings of existing silicon cochleae, filter banks in cascade and in parallel. Analog current representing the basilar membrane’s velocity drives the silicon auditory nerve, which encodes sound in digital pulses, mimicking neuronal spikes, as the final output of the cochlea.

[1]  Wai-Kai Chen,et al.  The VLSI Handbook , 2000 .

[2]  Carver A. Mead,et al.  Scanners for visualizing activity of analog VLSI circuitry , 1991 .

[3]  A. E. Hubbard,et al.  Rapid force production in the cochlea , 1989, Hearing Research.

[4]  W. Wong,et al.  A model cochlear partition involving longitudinal elasticity. , 2002, The Journal of the Acoustical Society of America.

[5]  G. K. Yates,et al.  Basilar membrane measurements and the travelling wave , 1986, Hearing Research.

[6]  Peter Dallos,et al.  Overview: Cochlear Neurobiology , 1996 .

[7]  L. Rabiner,et al.  CAN AUTOMATIC SPEECH RECOGNITION LEARN MORE FROM HUMAN SPEECH PERCEPTION ? , 2005 .

[8]  H. Engström,et al.  Supporting elements in the organ of Corti. I. Fibrillar structures in the supporting cells of the organ of Corti of mammals. , 1972, Acta oto-laryngologica. Supplementum.

[9]  S T Neely,et al.  A model of cochlear mechanics with outer hair cell motility. , 1993, The Journal of the Acoustical Society of America.

[10]  André van Schaik,et al.  Improved Silicon Cochlea using Compatible Lateral Bipolar Transistors , 1995, NIPS.

[11]  I. Whitfield Discharge Patterns of Single Fibers in the Cat's Auditory Nerve , 1966 .

[12]  Yichuang Sun,et al.  Continuous-Time Active Filter Design , 1998 .

[13]  Kwabena Boahen,et al.  A burst-mode word-serial address-event link-I: transmitter design , 2004, IEEE Transactions on Circuits and Systems I: Regular Papers.

[14]  P. Dallos The active cochlea , 1992, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[15]  W. S. Rhode,et al.  Mechanical responses to two-tone distortion products in the apical and basal turns of the mammalian cochlea. , 1997, Journal of neurophysiology.

[16]  J. Pierce,et al.  The cochlear compromise. , 1976, The Journal of the Acoustical Society of America.

[17]  Charles R. Steele,et al.  An improved WKB calculation for a two-dimensional cochlear model. , 1980, The Journal of the Acoustical Society of America.

[18]  I. Russell,et al.  The spatial and temporal representation of a tone on the guinea pig basilar membrane. , 2000, Proceedings of the National Academy of Sciences of the United States of America.

[19]  E. de Boer,et al.  Solving cochlear mechanics problems with higher-order differential equations. , 1982, The Journal of the Acoustical Society of America.

[20]  Norma B. Slepecky,et al.  Structure of the Mammalian Cochlea , 1996 .

[21]  Christofer Toumazou,et al.  Micropower log-domain filter for electronic cochlea , 1994 .

[22]  Craig C. Bader,et al.  Evoked mechanical responses of isolated cochlear outer hair cells. , 1985, Science.

[23]  C. Steele,et al.  Behavior of the basilar membrane with pure-tone excitation. , 1974, The Journal of the Acoustical Society of America.

[24]  S. Khanna,et al.  Mechanical response characteristics of the hearing organ in the low-frequency regions of the cochlea. , 1996, Journal of neurophysiology.

[25]  P Dallos,et al.  Nonlinearities in cochlear receptor potentials and their origins. , 1989, The Journal of the Acoustical Society of America.

[26]  L. Robles,et al.  Middle-ear response in the chinchilla and its relationship to mechanics at the base of the cochlea. , 1990, The Journal of the Acoustical Society of America.

[27]  K. G. Hill Basilar membrane motion in relation to two-tone suppression , 1998, Hearing Research.

[28]  Rahul Sarpeshkar,et al.  A Low-Power Wide-Linear-Range Transconductance Amplifier , 1997 .

[29]  Donald Lloyd Watts,et al.  Cochlear Mechanics: Analysis and Analog VLSI , 1993 .

[30]  L. Robles,et al.  Two-tone suppression in the basilar membrane of the cochlea: mechanical basis of auditory-nerve rate suppression. , 1992, Journal of neurophysiology.

[31]  C. Daniel Geisler A cochlear model using feedback from motile outer hair cells , 1991, Hearing Research.

[32]  C. Toumazou,et al.  Design of a micropower current-mode log-domain analog cochlear implant , 2000 .

[33]  Kwabena Boahen,et al.  Learning in Silicon: Timing is Everything , 2005, NIPS.

[34]  S. Espejo,et al.  Mismatch-Induced Trade-Offs and Scalability of Analog Preprocessing Visual Microprocessor Chips , 2003 .

[35]  Richard F. Lyon,et al.  An analog electronic cochlea , 1988, IEEE Trans. Acoust. Speech Signal Process..

[36]  J B Allen,et al.  Two-dimensional cochlear fluid model: new results. , 1977, The Journal of the Acoustical Society of America.

[37]  Kwabena Boahen,et al.  A linear cochlear model with active bi-directional coupling , 2003, Proceedings of the 25th Annual International Conference of the IEEE Engineering in Medicine and Biology Society (IEEE Cat. No.03CH37439).

[38]  E. Piette,et al.  A Textbook of Histology , 1936, The Indian Medical Gazette.

[39]  S. Neely Finite difference solution of a two-dimensional mathematical model of the cochlea. , 1981, The Journal of the Acoustical Society of America.

[40]  L. Voldřich Experimental and Topographic Morphology in Cochlear Mechanics , 1983 .

[41]  Charles R. Steele,et al.  A three-dimensional nonlinear active cochlear model analyzed by the WKB-numeric method , 2002, Hearing Research.

[42]  P.R. Kinget Device mismatch and tradeoffs in the design of analog circuits , 2005, IEEE Journal of Solid-State Circuits.

[43]  S. Neely Mathematical modeling of cochlear mechanics. , 1985, The Journal of the Acoustical Society of America.

[44]  G. K. Yates,et al.  Rate-versus-level functions of primary auditory nerve fibres: Evidence for square law behaviour of all fibre categories in the guinea pig , 1991, Hearing Research.

[45]  C. Daniel Geisler,et al.  Two-tone suppression by a saturating feedback model of the cochlear partition , 1992, Hearing Research.

[46]  M. Liberman The cochlear frequency map for the cat: labeling auditory-nerve fibers of known characteristic frequency. , 1982, The Journal of the Acoustical Society of America.

[47]  Y. Raphael,et al.  The sensory epithelium and its innervation in the mole rat cochlea , 1991, The Journal of comparative neurology.

[48]  Coding of sound intensity in the chick cochlear nerve. , 2002, Journal of neurophysiology.

[49]  J. Santos-Sacchi,et al.  On the frequency limit and phase of outer hair cell motility: effects of the membrane filter , 1992, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[50]  J. L. Hall Two-tone suppression in a nonlinear model of the basilar membrane. , 1977, The Journal of the Acoustical Society of America.

[51]  J.V. Arthur,et al.  Recurrently connected silicon neurons with active dendrites for one-shot learning , 2004 .

[52]  L. Robles,et al.  Basilar-membrane responses to tones at the base of the chinchilla cochlea. , 1997, The Journal of the Acoustical Society of America.

[53]  C. Daniel Geisler,et al.  A cochlear model using feed-forward outer-hair-cell forces , 1995, Hearing Research.

[54]  J. Flanagan Speech Analysis, Synthesis and Perception , 1971 .

[55]  Kwabena Boahen,et al.  The Retinomorphic Approach: Pixel-Parallel Adaptive Amplification, Filtering, and Quantization , 1997 .

[56]  Christian Enz,et al.  Low-voltage log-domain signal processing in CMOS and BiCMOS , 1999 .

[57]  Kwabena Boahen,et al.  An ON-OFF log domain circuit that recreates adaptive filtering in the retina , 2005, IEEE Transactions on Circuits and Systems I: Regular Papers.

[58]  Kian-Meng Lim Physical and mathematical cochlear models , 2000 .

[59]  E. de Boer,et al.  Validity of the Liouville-Green (or WKB) method for cochlear mechanics , 1982, Hearing Research.

[60]  Corné J. Kros,et al.  Physiology of Mammalian Cochlear Hair Cells , 1996 .

[61]  Stephen T. Neely,et al.  An active cochlear model showing sharp tuning and high sensitivity , 1983, Hearing Research.

[62]  J. Allen,et al.  A second cochlear-frequency map that correlates distortion product and neural tuning measurements. , 1993, The Journal of the Acoustical Society of America.

[63]  M. Sondhi,et al.  Method for computing motion in a two-dimensional cochlear model. , 1978, The Journal of the Acoustical Society of America.

[64]  A. Nuttall,et al.  Two-tone suppression of inner hair cell and basilar membrane responses in the guinea pig. , 1993, The Journal of the Acoustical Society of America.

[65]  Kwabena Boahen,et al.  A Recurrent Model of Orientation Maps with Simple and Complex Cells , 2003, NIPS.

[66]  Jonathan E. Gale,et al.  An intrinsic frequency limit to the cochlear amplifier , 1997, Nature.

[67]  Malcolm Slaney,et al.  Lyon's Cochlear Model , 1997 .

[68]  D. R. Frey,et al.  Log-domain filtering: An approach to current-mode fil-tering , 1993 .

[69]  L. Robles,et al.  Mechanics of the mammalian cochlea. , 2001, Physiological reviews.

[70]  L A Taber,et al.  Comparison of WKB and finite difference calculations for a two-dimensional cochlear model. , 1979, The Journal of the Acoustical Society of America.

[71]  M A Viergever,et al.  Numerical methods for solving one-dimensional cochlear models in the time domain. , 1987, The Journal of the Acoustical Society of America.

[72]  Kwabena Boahen,et al.  A retinomorphic vision system , 1996, IEEE Micro.

[73]  Manfred Kössl,et al.  A Targeted Deletion in α-Tectorin Reveals that the Tectorial Membrane Is Required for the Gain and Timing of Cochlear Feedback , 2000, Neuron.

[74]  A. Hubbard,et al.  A traveling-wave amplifier model of the cochlea. , 1993, Science.

[75]  M A Viergever,et al.  Nonlinear and active two-dimensional cochlear models: time-domain solution. , 1989, The Journal of the Acoustical Society of America.

[76]  H. Ohmori,et al.  Voltage‐gated and chemically gated ionic channels in the cultured cochlear ganglion neurone of the chick. , 1990, The Journal of physiology.

[77]  The Basilar Membrane Nonlinear Input-Output Function , 1990 .

[78]  C. Geisler,et al.  Suppression in auditory-nerve fibers of cats using low-side suppressors. I. Temporal aspects , 1996, Hearing Research.

[79]  P M Sellick,et al.  Low‐frequency characteristics of intracellularly recorded receptor potentials in guinea‐pig cochlear hair cells. , 1983, The Journal of physiology.

[80]  I. J. Russell,et al.  The responses of inner and outer hair cells in the basal turn of the guinea-pig cochlea and in the mouse cochlea grown in vitro , 1986, Hearing Research.

[81]  W. S. Rhode Observations of the vibration of the basilar membrane in squirrel monkeys using the Mössbauer technique. , 1971, The Journal of the Acoustical Society of America.

[82]  Guy Rebillard,et al.  Correlation Between the Length of Outer Hair Cells and the Frequency Coding of the Cochlea , 1992 .

[83]  T. Fukazawa,et al.  Spontaneous otoacoustic emissions in an active feed-forward model of the cochlea , 1996, Hearing Research.

[84]  Eric Fragnière,et al.  Analogue VLSI emulation of the cochlea , 1998 .

[85]  H. Spoendlin Innervation densities of the cochlea. , 1972, Acta oto-laryngologica.

[86]  Kwabena Boahen,et al.  Topographic Map Formation by Silicon Growth Cones , 2002, NIPS.

[87]  L. Voldřich,et al.  Mechanical properties of basilar membrane. , 1978, Acta oto-laryngologica.

[88]  Kwabena Boahen,et al.  Point-to-point connectivity between neuromorphic chips using address events , 2000 .

[89]  W. T. Peake,et al.  Experiments in Hearing , 1963 .

[90]  C D Geisler,et al.  A two-stage nonlinear cochlear model possesses automatic gain control. , 1986, The Journal of the Acoustical Society of America.

[91]  Misha Mahowald,et al.  A silicon model of early visual processing , 1993, Neural Networks.

[92]  A theoretical basis for the high-frequency performance of the outer hair cell’s receptor potential , 1997 .

[93]  R. Schaumann,et al.  Low-voltage high-speed current-mode continuous-time IC filters with orthogonal /spl omega//sub 0/-Q tuning , 1999 .

[94]  Longitudinal Coupling in the Basilar Membrane , 2001, Journal of the Association for Research in Otolaryngology.

[95]  R. S. Chadwick WHAT SHOULD BE THE GOALS OF COCHLEAR MODELING , 1997 .

[96]  Tatsuya Fukazawa,et al.  How can the cochlear amplifier be realized by the outer hair cells which have nothing to push against? , 2002, Hearing Research.

[97]  Fabio Mammano,et al.  Reverse transduction measured in the isolated cochlea by laser Michelson interferometry , 1993, Nature.

[98]  S. Neely From Sound to Synapse: Physiology of the Mammalian Ear , 1998 .

[99]  Kwabena Boahen,et al.  Translinear circuits in subthreshold MOS , 1996 .

[100]  C. Daniel Geisler A realizable cochlear model using feedback from motile outer hair cells , 1993, Hearing Research.

[101]  J. Ashmore A fast motile response in guinea‐pig outer hair cells: the cellular basis of the cochlear amplifier. , 1987, The Journal of physiology.

[102]  H. Schmid Why the terms 'current mode' and 'voltage mode' neither divide nor qualify circuits , 2002, 2002 IEEE International Symposium on Circuits and Systems. Proceedings (Cat. No.02CH37353).

[103]  E. Fragniere,et al.  A 100-channel analog CMOS auditory filter bank for speech recognition , 2005, ISSCC. 2005 IEEE International Digest of Technical Papers. Solid-State Circuits Conference, 2005..

[104]  M. Sondhi,et al.  Cochlear macromechanics: time domain solutions. , 1979, The Journal of the Acoustical Society of America.

[105]  S. Neely,et al.  A model for active elements in cochlear biomechanics. , 1986, The Journal of the Acoustical Society of America.

[106]  D O Kim,et al.  A system of nonlinear differential equations modeling basilar-membrane motion. , 1973, The Journal of the Acoustical Society of America.

[107]  C D Geisler,et al.  A hybrid-computer model of the cochlear partition. , 1972, The Journal of the Acoustical Society of America.

[108]  D. Mountain,et al.  Haircell forward and reverse transduction: Differential suppression and enhancement , 1990, Hearing Research.

[109]  D. D. Greenwood A cochlear frequency-position function for several species--29 years later. , 1990, The Journal of the Acoustical Society of America.

[110]  Carver Mead,et al.  Analog VLSI and neural systems , 1989 .

[111]  Bertram E. Shi,et al.  Neuromorphic implementation of orientation hypercolumns , 2005, IEEE Transactions on Circuits and Systems I: Regular Papers.