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Modeling the Neural Representation of Interaural Level Differences for Linked and Unlinked Bilateral Hearing Aids Stephanie T. Cheung and Ian C. Bruce Department of Electrical & Computer Engineering McMaster University Hamilton, ON, Canada [email protected] AL AUDITORY ENGINEERING LABORATORY ABSTRACT Interaural level difference (ILD) cues facilitate horizontal localization of sound frequencies above 1.5 kHz and are processed by the lateral superior olive (LSO) of the brainstem. The LSO is excited by sounds that are louder in the ipsilateral ear than in the contralateral ear. Bilateral hearing aids with independent wide dynamic range compression (WDRC) will tend to reduce the ILD cues. Hearing aids with wirelessly-linked WDRC algorithms are reported to have varying levels of success in restoring or maintaining an individual’s ability to understand and localize sounds. In our study, we model the neural processing of the LSO in response to linked and unlinked WDRC schemes. Two computer models are used to predict LSO responses. The first is a phenomenological model that takes inputs directly from ipsilateral and contralateral outputs of an auditory nerve (AN) model [12, 13]. The LSO response is estimated from the difference in AN spike rates and a sigmoidal nonlinearity to account for the threshold and saturation in LSO spike rates. The second is a Hodgkin–Huxley-type biophysical model, which details the ion channel behavior of LSO cells. Inputs to this model are excitatory and inhibitory synaptic currents derived from the outputs of the ipsilateral and contralateral AN models, respectively. Both normal-hearing and hearing-impaired responses are generated. AUDIS [2] head-related transfer functions are used to simulate sound sources ranging from -80° to +80° on the horizontal plane. Unlinked WDRC degrades the coding of sound source azimuth in the LSO by reducing ILDs. Wirelessly linked hearing aids maintain ILD cues and have the potential to enable the LSO response to indicate azimuth, provided sufficient gain and appropriate compression time constants. These results will contribute to our understanding of the azimuthal perception of sounds at the neuronal level. This has implications for the design of hearing aids to provide optimal spatial awareness to the listener. I. INTRODUCTION A. Interaural Level Difference • Auditory localization is important for safe functioning in everyday environments and is a factor in speech intelligibility. • Azimuthal localization is facilitated by interaural timing difference (ITD) cues, which are most perceptually salient for low-frequency sounds (below 1500 Hz), while interaural level difference (ILD) cues are predominant for high-frequency sounds [1]. • ILDs are a product of the acoustic head shadow; sounds are attenuated at the farther, shadowed ear relative to the closer, unshadowed ear (Fig. 1(L)). Figure 1: [Left]: ILD cues arise from the head shadow effect. [Right]: Neural processing of ILD cues occurs in the lateral superior olive (LSO). • Sensorineural hearing impairment reduces dynamic range: Binaural cues for localization may be less salient, such that no benefit can be derived. B. Wireless Wide Dynamic Range Compression • Wide dynamic range compression (WDRC) hearing aids typically skew ILD cues by applying differing gains to the two bilateral devices. • Wireless technologies allow communication between bilateral devices so that the gain applied at each ear is dependent on the levels at both ears. • In one scheme under investigation, linked amplification applies the lowest gain required by either of the two [11]. • Performance with these devices is varied. Table 1: Reports on wireless WDRC benefit are varied. Author Measure and Test Conclusions Ibrahim et al., 2012 [4] Sound localization Speech intelligibility Improved localization. No benefit to speech-in-noise scores. Sockalingham, 2009 [8] Sound localization Quality Improved localization. Improved naturalness only in certain environments. Wiggins and Seeber, 2013 [11] Speech intelligibility Improved performance is due to improved SNR at worse ear, not wireless link. Schwartz and Shinn-Cunningham, 2013 [6] Spatial thresholds Improved localization for fast attack and release times. No trend in slow WDRC. Smith, 2008 [7] Preference 65% prefer wirelessly-linked WDRC. C. The Neural Pathway for ILD Cues • ILD cues are analyzed in the lateral superior olive (LSO) of the brainstem. • The LSO receives inhibitory inputs from the contralateral anteroventral cochlear nucleus (AVCN) and excitatory inputs from the ipsilateral AVCN (Fig. 1(R)). • LSO cells are tonotopically mapped and are excited by greater intensity in the ipsilateral ear. • The relationship between ILD and excitation (discharge rate in spikes/s) is sigmoidal. • Some suggest that the sigmoidal tuning function shifts with respect to overall levels of sound presentation, although other studies find ILD thresholds to be level-invariant [9]. II. METHODS • LSO representations of stimuli were modelled for an ideal normal-hearing listener and an example hearing-impaired listener. A. Hearing Aid Model • A model of a simple, unilateral WDRC hearing aid was programmed in M ATLAB. • A separate script allowed for simulated wireless linking between two hearing aids, assuming instantaneous communication. • The model uses overlap-and-add, applying amplification in the frequency domain. B. Auditory Periphery Model • An existing model of the auditory periphery [12, 13] was used to generate ipsilateral and contralateral AN spike trains. • Inner and outer hair cell control parameters are adjusted to reflect the listener’s audiogram. C. Phenomenological Model of the Lateral Superior Olive • A phenomenological model of the lateral superior olive was programmed to simulate the sigmoidal relationship between ILD and spike rate. • The sigmoidal tuning function is defined as: y = a + b 1 + exp c-x d (1) y is LSO spike rate and x is the difference between ipsilateral and contralateral AN inputs. • Parameters a, b, c, and d are chosen to give spike rates consistent with Tsai et al. [9] D. Biophysical Model of the Lateral Superior Olive • A Hodgkin–Huxley-type biophysical model of the LSO was also used, guided by the ventral cochlear nucleus Hodgkin–Huxley models of Rothman and Manis [5]. • Ion channel parameters were adapted from Wang and Colburn [10]. C m d V d t = I A + I LT + I HT + I Na + I h + I lk + I Exc + I Inh (2) • Currents are as follows: Table 2: Currents of the Hodgkin–Huxley-type biophysical LSO model. Notation Current Type I A fast-inactivating K + current I LT low-threshold K + current I HT high-threshold K + current I Na Na + current I h hyperpolarization-activated cation current I lk leakage current I Exc excitatory synaptic current (i.e. ipsilateral AN input) I Inh inhibitory synaptic current (i.e. contralateral AN input) E. Listener Models • The Wiggins and Seeber [11] hearing-impaired audiogram and CAMEQ fit were used for the hearing-impaired model. • The normal-hearing model assumed completely undamaged inner and outer hair cells. Table 3: Hearing-impaired audiogram from Wiggins and Seeber [11]. Audiogram Frequency (Hz) 250 375 500 750 1000 1500 2000 3000 4000 6000 dB HL 35 35 35 35 40 45 50 55 60 65 Table 4: Hearing aid prescription as set by Wiggins and Seeber [11]. Channel Lower Edge Frequency Upper Edge Frequency Compression Threshold Compression Ratio IG50 IG80 1 125 250 46 1.53 10.2 0.0 2 250 750 47 1.11 10.3 7.4 3 750 1250 40 1.99 20.8 5.9 4 1250 1750 30 1.89 20.5 6.4 5 1750 2250 27 1.98 22.9 8.0 6 2250 2750 28 2.01 21.7 6.6 7 2750 3250 27 1.99 20.7 5.8 8 3250 3750 24 1.99 21.4 6.5 9 3750 4250 20 2.01 22.2 7.1 10 4250 4750 22 2.22 21.6 5.1 11 4750 5250 21 2.51 20.8 2.8 12 5250 5750 22 2.80 20.2 1.0 F. Stimuli • A 4 kHz puretone stimulus was used with a duration of 2 s and 5-ms on and off ramps. • A sentence of Northern American-accented speech from the TIMIT corpus was also used: “She had your dark suit in greasy wash water all year”. • Stimuli were presented at azimuths ranging from -80° to +80°. • AUDIS HRTFs were used to generate realistic ILD cues [2]. G. Test Conditions • For the normal-hearing model, only unaided listening was tested. • For the hearing-impaired model, simulations included: unaided hearing; linked, bilateral WDRC; and unlinked, bilateral WDRC. • In the aided conditions, three sets of compression times were used: Wiggins and Seeber “WS” (5 ms attack, 60 ms release) for all stimuli [11]. Very Fast (1 ms attack, 10 ms release) for the TIMIT sentence. Slow (100 ms attack, 400 ms release) for the TIMIT centence. • Attack and release time choices were guided by Dillon [3]. Table 5: Test Conditions. P denotes use of the phenomenological model and B of the biophysical model. Hearing-Impaired Unlinked WDRC Linked WDRC Stimulus Normal-Hearing No WDRC WS Very Fast Slow WS Very Fast Slow Puretone 35 dB SPL P/B P P P 45 dB SPL P P P P 55 dB SPL P P P P 65 dB SPL P/B P P P 75 dB SPL P P P P 85 dB SPL P/B P P P TIMIT Speech P P P P P P P P H. Measure: Mean Discharge Rate Difference • The mean discharge rate difference was calculated to quantify the degree of similarity to which the hearing-impaired model performed, relative to the normal-hearing model. CFs (Hz) Azimuth Normal−Hearing, Left LSO −50 0 50 2000 4000 6000 0 50 100 150 CFs (Hz) Azimuth Hearing−Impaired, Linked Compression, Left LSO −50 0 50 2000 4000 6000 0 50 100 150 −80 −60 −40 −20 0 20 40 60 80 −20 −10 0 10 20 65 dB SPL Azimuth Mean Discharge Rate Difference (spikes/s) ) = mean ( Figure 2: Calculation of the mean discharge rate difference over CF. III. RESULTS A. ILD Cue Distortion • Linked WDRC retains ILD cues (Fig. 3 A). • Unlinked compression reduces ILD cues, and the reduction varies with presentation level (Fig. 3 B). −80 −60 −40 −20 0 20 40 60 80 −15 −10 −5 0 5 10 15 ILD (Left Level − Right Level) in dB SPL Azimuth −80 −60 −40 −20 0 20 40 60 80 −6 −4 −2 0 2 4 6 ILD (Left Level − Right Level) in dB SPL Azimuth Figure 3: [Left]: Linked WDRC retains ILD cues. Dots denote natural ILDs. Open circles denote linked ILDs. [Right]: Unlinked WDRC distorts ILD cues to an extent variable with presentation level. Black: 35 dB SPL; Blue: 45 dB SPL; Red: 55 dB SPL; Green: 65 dB SPL; Magenta: 75 dB SPL; Cyan: 85 dB SPL B. Puretone Stimuli Results • In the hearing-impaired model, linked compression performs most similarly to the normal-hearing model for all puretone stimuli. • The impact of WDRC is greatest near 55 – 65 dB SPL and least at low frequencies. • Unaided listening results in the most different neural representations from normal hearings at all levels but 85 dB SPL. • At 85 dB SPL, unlinked WDRC is worse than unaided listening. • Normal-hearing discharge rates exceed all discharge rates for impaired hearing (Fig. 5). −80 −60 −40 −20 0 20 40 60 80 −20 −10 0 10 20 35 dB SPL −80 −60 −40 −20 0 20 40 60 80 −20 −10 0 10 20 45 dB SPL −80 −60 −40 −20 0 20 40 60 80 −20 −10 0 10 20 55 dB SPL −80 −60 −40 −20 0 20 40 60 80 −20 −10 0 10 20 65 dB SPL Mean Discharge Rate Difference (spikes/s) −80 −60 −40 −20 0 20 40 60 80 −20 −10 0 10 20 75 dB SPL Azimuth −80 −60 −40 −20 0 20 40 60 80 −20 −10 0 10 20 85 dB SPL Figure 4: LSO mean differences under the phenomenological model in spikes/s. The following hearing-impaired neural representations are subtracted from the normal-hearing response, and denoted as follows: Black: unaided; Blue: unlinked WS WDRC; Red: linked WS WDRC. CFs (Hz) Normal-Hearing, Left LSO -80 -60 -40 -20 0 20 40 60 80 2000 4000 6000 0 50 100 150 CFs (Hz) Azimuth Normal-Hearing, Right LSO -80 -60 -40 -20 0 20 40 60 80 2000 4000 6000 0 50 100 150 CFs (Hz) Hearing-Impaired, Unaided, Left LSO -80 -60 -40 -20 0 20 40 60 80 2000 4000 6000 0 50 100 150 CFs (Hz) Azimuth Hearing-Impaired, Unaided, Right LSO -80 -60 -40 -20 0 20 40 60 80 2000 4000 6000 0 50 100 150 CFs (Hz) Hearing-Impaired, Unlinked Compression, Left LSO -80 -60 -40 -20 0 20 40 60 80 2000 4000 6000 0 50 100 150 CFs (Hz) Azimuth Hearing-Impaired, Unlinked Compression, Right LSO -80 -60 -40 -20 0 20 40 60 80 2000 4000 6000 0 50 100 150 CFs (Hz) Hearing-Impaired, Linked Compression, Left LSO -80 -60 -40 -20 0 20 40 60 80 2000 4000 6000 0 50 100 150 CFs (Hz) Azimuth Hearing-Impaired, Linked Compression, Right LSO -80 -60 -40 -20 0 20 40 60 80 2000 4000 6000 0 50 100 150 Figure 5: An example of patterns of neural response to ILD cues (65 dB SPL puretone stimulus). Normal-hearing spike rates are higher than those in the hearing-impaired model for all compression conditions. C. TIMIT Sentence Results • No compression condition consistently results in the lowest mean difference scores (Fig. 6). • Not all phones contain enough energy to trigger WDRC (Fig. 7). • WDRC benefit varies within phone class; frequency band energy is not the only indicator of performance. • Phone placement is important: In all stop closures but ‘gcl’, which immediately followed a nasal, spike rates for the hearing-impaired model exceeded those of the normal-hearing model (8). no WDRC unlinked WS linked WS unlinked Very Fast linked Very Fast unlinked Slow linked Slow 0 0.5 1 Fricative no WDRC unlinked WS linked WS unlinked Very Fast linked Very Fast unlinked Slow linked Slow 0 0.5 1 Affricatives no WDRC unlinked WS linked WS unlinked Very Fast linked Very Fast unlinked Slow linked Slow 0 0.2 0.4 Normalized Number of Occurances Across All Azimuths Lowest Mean Discharge Rate Difference Semivowels/Glides no WDRC unlinked WS linked WS unlinked Very Fast linked Very Fast unlinked Slow linked Slow 0 0.5 1 Vowels Figure 6: Neither linked nor unlinked WDRC is consistently preferable for speech sounds, although linked is often better. No particular set of compression speeds is consistently best. 0 1000 2000 3000 4000 5000 6000 7000 8000 −20 0 20 40 60 Fricatives 0 1000 2000 3000 4000 5000 6000 7000 8000 −20 0 20 40 60 Vowels 0 1000 2000 3000 4000 5000 6000 7000 8000 −20 0 20 40 60 Semivowels/Glides Frequency (Hz) 0 1000 2000 3000 4000 5000 6000 7000 8000 −20 0 20 40 60 Affricatives Sound Level (dB SPL) Figure 7: Energy in some phones of the TIMIT sentence. Solid blue lines indicate the range of AN fiber CFs modelled, while the dashed blue line indicates the 1500 Hz boundary between low and high frequencies. The red curve denotes the compression threshold. −50 0 50 −10 −5 0 5 10 "dcl", 0.4775s to 0.5188s Azimuth Mean Discharge Rate Difference (spikes/s) −50 0 50 −10 −5 0 5 10 "dcl", 0.6525s to 0.7288s Azimuth Mean Discharge Rate Difference (spikes/s) −50 0 50 −10 −5 0 5 10 "kcl", 0.8975s to 0.9619s Azimuth Mean Discharge Rate Difference (spikes/s) −50 0 50 −10 −5 0 5 10 "gcl", 1.4800s to 1.5378s Azimuth Mean Discharge Rate Difference (spikes/s) Figure 8: Phone placement seems to impact WDRC performance. The following hearing-impaired neural representations are subtracted from the normal-hearing response, and denoted as follows:- Black: unaided; Blue: unlinked WDRC; Red: linked WDRC. Compression speeds are denoted as follows:- Solid lines: WS compression; Dashed lines: Very Fast compression; Dotted lines: Slow compression. IV. CONCLUSIONS • The benefit of linked WDRC over unlinked WDRC is clear for puretone stimuli. • In speech stimuli, the benefit is dependent on frequency content, presentation level, phone class, and phone placement within a sentence. • Adaptive attack and release times may show the most promise as this technology continues to develop. V. ACKNOWLEDGMENTS This work is supported by NSERC Discovery Grant 261736. REFERENCES [1] M. A. Akeroyd, “The psychoacoustics of binaural hearing,” International Journal of Audiology. Volume 45, Issue S1, pgs. 25-33, 2006. [2] J. Blauert et al., “The AUDIS catalog of human HRTFs,” The Journal of the Acoustical Society of America. Volume 103, Issue 5, pgs. 3082-3082, May 1998. [3] H. Dillon, Hearing Aids (Second Edition). Turramurra, Australia: Boomerang Press, 2012. [4] I. Ibrahim et al., “Evaluation of speech intelligibility and sound localization abilities with hearing aids using binaural wireless technology,” Audiology Research. Volume 3, Issue 1, pgs. 1-9, December 2012. [5] J. S. Rothman And P. B. Manis, “The Roles Potassium Currents Play in Regulating the Electrical Activity of Ventral Cochlear Nucleus Neurons,” Journal of Neurophysiology. Volume 89, Issue 6, pgs. 3097-3113, February 2003. [6] A. H. Schwartz and B. Shinn-Cunningham, “Effects of dynamic range compression on spatial selective auditory attention in normal-hearing listeners,” The Hearing Journal. Volume 133, Issue 4, pgs. 2329-2339, 2013. [7]P. Smith et al., “Real-world preferences for linked bilateral processing,” The Hearing Journal. Volume 61, Issue 7, pgs. 33-34, July 2008. [8] R. Sockalingham et al., “Binaural hearing aid communication shown to improve sound quality and localization,” The Hearing Journal. Volume 62, Issue 10, pgs. 46-47, October 2009. [9]J. J. Tsai et al., “Varying Overall Sound Intensity to the Two Ears Impacts Interaural Level Difference Discrimination Thresholds by Single Neurons in the Lateral Superior Olive,” Journal of Neurophysiology. Volume 103, Issue 2, pgs. 875-886, February 2010. [10] L. Wang et S. Colburn, “A Modeling Study of the Responses of the Lateral Superior Olive to Ipsilateral Sinusoidally Amplitude-Modulated Tones,” Journal of the Association for Research in Otolaryngology. Volume 13, Issue 2, pgs. 249-267, April 2012. [11] I. M. Wiggins and B. U. Seeber, “Linking dynamic-range compression across the ears can improve speech intelligibility in spatially separated noise,” The Journal of the Acoustical Society of America. Volume 133, Issue 2, pgs. 1004-1016, 2013. [12] M. S. A. Zilany et al., “Improved parameters and expanded simulation options for a model of the auditory periphery,” Abstracts of the 36 th ARO Midwinter Research Meeting. Association for Research in Otolaryngology. Baltimore, MD, pgs. 440-441, 2013. [13] M. S. A. Zilany et al., “A phenomenologicial model of the synapse between the inner hair cell and auditory nerve: Long-term adaptation with power-law dynamics,” Journal of the Acoustical Society of America. Volume 126, Issue 5, pgs. 2390-2412, 2009. IHCON 2014, Lake Tahoe - CA, August 13-17

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Page 1: A L - McMaster Universityibruce/papers/cheung_IHCON2014.pdf · •A separate script allowed for simulated wireless linking between two hearing aids, assuming instantaneous communication

Modeling the Neural Representation of Interaural Level Differences for Linked and Unlinked Bilateral Hearing AidsStephanie T. Cheung and Ian C. Bruce

Department of Electrical & Computer Engineering • McMaster University • Hamilton, ON, [email protected]

A L

AUDITORYENGINEERINGLABORATORY

ABSTRACT

Interaural level difference (ILD) cues facilitate horizontal localization of soundfrequencies above ∼ 1.5 kHz and are processed by the lateral superior olive (LSO)of the brainstem. The LSO is excited by sounds that are louder in the ipsilateral earthan in the contralateral ear. Bilateral hearing aids with independent wide dynamicrange compression (WDRC) will tend to reduce the ILD cues. Hearing aids withwirelessly-linked WDRC algorithms are reported to have varying levels of success inrestoring or maintaining an individual’s ability to understand and localize sounds. In ourstudy, we model the neural processing of the LSO in response to linked and unlinkedWDRC schemes.

Two computer models are used to predict LSO responses. The first is aphenomenological model that takes inputs directly from ipsilateral and contralateraloutputs of an auditory nerve (AN) model [12, 13]. The LSO response is estimatedfrom the difference in AN spike rates and a sigmoidal nonlinearity to account for thethreshold and saturation in LSO spike rates. The second is a Hodgkin–Huxley-typebiophysical model, which details the ion channel behavior of LSO cells. Inputs tothis model are excitatory and inhibitory synaptic currents derived from the outputs ofthe ipsilateral and contralateral AN models, respectively. Both normal-hearing andhearing-impaired responses are generated. AUDIS [2] head-related transfer functionsare used to simulate sound sources ranging from −80° to +80° on the horizontal plane.

Unlinked WDRC degrades the coding of sound source azimuth in the LSO by reducingILDs. Wirelessly linked hearing aids maintain ILD cues and have the potential toenable the LSO response to indicate azimuth, provided sufficient gain and appropriatecompression time constants. These results will contribute to our understanding of theazimuthal perception of sounds at the neuronal level. This has implications for thedesign of hearing aids to provide optimal spatial awareness to the listener.

I. INTRODUCTION

A. Interaural Level Difference

• Auditory localization is important for safe functioning in everyday environments andis a factor in speech intelligibility.

• Azimuthal localization is facilitated by interaural timing difference (ITD) cues, whichare most perceptually salient for low-frequency sounds (below 1500 Hz), whileinteraural level difference (ILD) cues are predominant for high-frequency sounds [1].

• ILDs are a product of the acoustic head shadow; sounds are attenuated at the farther,shadowed ear relative to the closer, unshadowed ear (Fig. 1(L)).

Figure 1: [Left]: ILD cues arise from the head shadow effect. [Right]: Neural processingof ILD cues occurs in the lateral superior olive (LSO).

• Sensorineural hearing impairment reduces dynamic range: Binaural cues forlocalization may be less salient, such that no benefit can be derived.

B. Wireless Wide Dynamic Range Compression

• Wide dynamic range compression (WDRC) hearing aids typically skew ILD cues byapplying differing gains to the two bilateral devices.

• Wireless technologies allow communication between bilateral devices so that the gainapplied at each ear is dependent on the levels at both ears.

• In one scheme under investigation, linked amplification applies the lowest gainrequired by either of the two [11].

• Performance with these devices is varied.

Table 1: Reports on wireless WDRC benefit are varied.

Author Measure and Test Conclusions

Ibrahim et al., 2012 [4] Sound localizationSpeech intelligibility

Improved localization.No benefit to speech-in-noise scores.

Sockalingham, 2009 [8] Sound localizationQuality

Improved localization.Improved naturalness only in certain environments.

Wiggins and Seeber, 2013 [11] Speech intelligibility Improved performance is due to improved SNR at worseear, not wireless link.

Schwartz andShinn-Cunningham, 2013 [6] Spatial thresholds Improved localization for fast attack and release times.

No trend in slow WDRC.Smith, 2008 [7] Preference 65% prefer wirelessly-linked WDRC.

C. The Neural Pathway for ILD Cues

• ILD cues are analyzed in the lateral superior olive (LSO) of the brainstem.• The LSO receives inhibitory inputs from the contralateral anteroventral cochlearnucleus (AVCN) and excitatory inputs from the ipsilateral AVCN (Fig. 1(R)).

• LSO cells are tonotopically mapped and are excited by greater intensity in theipsilateral ear.

• The relationship between ILD and excitation (discharge rate in spikes/s) is sigmoidal.• Some suggest that the sigmoidal tuning function shifts with respect to overalllevels of sound presentation, although other studies find ILD thresholds to belevel-invariant [9].

II. METHODS

• LSO representations of stimuli were modelled for an ideal normal-hearing listenerand an example hearing-impaired listener.

A. Hearing Aid Model

• A model of a simple, unilateral WDRC hearing aid was programmed in MATLAB.• A separate script allowed for simulated wireless linking between two hearing aids,assuming instantaneous communication.

• The model uses overlap-and-add, applying amplification in the frequency domain.

B. Auditory Periphery Model

• An existing model of the auditory periphery [12, 13] was used to generate ipsilateraland contralateral AN spike trains.

• Inner and outer hair cell control parameters are adjusted to reflect the listener’saudiogram.

C. Phenomenological Model of the Lateral Superior Olive

• A phenomenological model of the lateral superior olive was programmed to simulatethe sigmoidal relationship between ILD and spike rate.

• The sigmoidal tuning function is defined as:

y = a+b

1+ exp c−xd

(1)

• y is LSO spike rate and x is the difference between ipsilateral and contralateral ANinputs.

• Parameters a, b, c, and d are chosen to give spike rates consistent with Tsai et al. [9]

D. Biophysical Model of the Lateral Superior Olive

• A Hodgkin–Huxley-type biophysical model of the LSO was also used, guided by theventral cochlear nucleus Hodgkin–Huxley models of Rothman and Manis [5].

• Ion channel parameters were adapted from Wang and Colburn [10].

CmdVdt

= IA+ ILT+ IHT+ INa+ Ih+ Ilk+ IExc+ IInh (2)

• Currents are as follows:

Table 2: Currents of the Hodgkin–Huxley-type biophysical LSO model.Notation Current TypeIA fast-inactivating K+ currentILT low-threshold K+ currentIHT high-threshold K+ currentINa Na+ currentIh hyperpolarization-activated cation currentIlk leakage currentIExc excitatory synaptic current (i.e. ipsilateral AN input)IInh inhibitory synaptic current (i.e. contralateral AN input)

E. Listener Models

• The Wiggins and Seeber [11] hearing-impaired audiogram and CAMEQ fit were usedfor the hearing-impaired model.

• The normal-hearing model assumed completely undamaged inner and outer haircells.

Table 3: Hearing-impaired audiogram from Wiggins and Seeber [11].

AudiogramFrequency (Hz) 250 375 500 750 1000 1500 2000 3000 4000 6000dB HL 35 35 35 35 40 45 50 55 60 65

Table 4: Hearing aid prescription as set by Wiggins and Seeber [11].

Channel Lower Edge Frequency Upper Edge Frequency Compression Threshold Compression Ratio IG50 IG801 125 250 46 1.53 10.2 0.02 250 750 47 1.11 10.3 7.43 750 1250 40 1.99 20.8 5.94 1250 1750 30 1.89 20.5 6.45 1750 2250 27 1.98 22.9 8.06 2250 2750 28 2.01 21.7 6.67 2750 3250 27 1.99 20.7 5.88 3250 3750 24 1.99 21.4 6.59 3750 4250 20 2.01 22.2 7.110 4250 4750 22 2.22 21.6 5.111 4750 5250 21 2.51 20.8 2.812 5250 5750 22 2.80 20.2 1.0

F. Stimuli

• A 4 kHz puretone stimulus was used with a duration of 2 s and 5-ms on and off ramps.• A sentence of Northern American-accented speech from the TIMIT corpus was alsoused: “She had your dark suit in greasy wash water all year”.

• Stimuli were presented at azimuths ranging from −80° to +80°.• AUDIS HRTFs were used to generate realistic ILD cues [2].

G. Test Conditions

• For the normal-hearing model, only unaided listening was tested.• For the hearing-impaired model, simulations included: unaided hearing; linked,bilateral WDRC; and unlinked, bilateral WDRC.

• In the aided conditions, three sets of compression times were used:– Wiggins and Seeber “WS” (5 ms attack, 60 ms release) for all stimuli [11].– Very Fast (1 ms attack, 10 ms release) for the TIMIT sentence.– Slow (100 ms attack, 400 ms release) for the TIMIT centence.

• Attack and release time choices were guided by Dillon [3].

Table 5: Test Conditions. P denotes use of the phenomenological model and B of thebiophysical model.

Hearing-ImpairedUnlinked WDRC Linked WDRCStimulus Normal-Hearing No WDRC WS Very Fast Slow WS Very Fast Slow

Puretone 35 dB SPL P/B P P P45 dB SPL P P P P55 dB SPL P P P P65 dB SPL P/B P P P75 dB SPL P P P P85 dB SPL P/B P P P

TIMIT Speech P P P P P P P P

H. Measure: Mean Discharge Rate Difference

• The mean discharge rate difference was calculated to quantify the degree of similarityto which the hearing-impaired model performed, relative to the normal-hearing model.

CFs

(Hz)

Azimuth

Normal−Hearing, Left LSO

−50 0 50

2000

4000

6000

0

50

100

150

CFs

(Hz)

Azimuth

Hearing−Impaired, Linked Compression, Left LSO

−50 0 50

2000

4000

6000

0

50

100

150

−80 −60 −40 −20 0 20 40 60 80

−20

−10

0

10

20

65 dB SPL

Azimuth

Mea

n D

isch

arge

Rat

e D

iffer

ence

(spi

kes/

s)

− ) =mean (

Figure 2: Calculation of the mean discharge rate difference over CF.

III. RESULTS

A. ILD Cue Distortion

• Linked WDRC retains ILD cues (Fig. 3 A).• Unlinked compression reduces ILD cues, and the reduction varies with presentationlevel (Fig. 3 B).

−80 −60 −40 −20 0 20 40 60 80−15

−10

−5

0

5

10

15

ILD

(Lef

t Lev

el −

Rig

ht L

evel

) in

dB S

PL

Azimuth−80 −60 −40 −20 0 20 40 60 80

−6

−4

−2

0

2

4

6

ILD

(Lef

t Lev

el −

Rig

ht L

evel

) in

dB S

PL

Azimuth

Figure 3: [Left]: Linked WDRC retains ILD cues. Dots denote natural ILDs. Opencircles denote linked ILDs. [Right]: Unlinked WDRC distorts ILD cues to an extentvariable with presentation level. Black: 35 dB SPL; Blue: 45 dB SPL; Red: 55 dB SPL;Green: 65 dB SPL; Magenta: 75 dB SPL; Cyan: 85 dB SPL

B. Puretone Stimuli Results

• In the hearing-impaired model, linked compression performs most similarly to thenormal-hearing model for all puretone stimuli.

• The impact of WDRC is greatest near 55 – 65 dB SPL and least at low frequencies.• Unaided listening results in the most different neural representations from normalhearings at all levels but 85 dB SPL.

• At 85 dB SPL, unlinked WDRC is worse than unaided listening.• Normal-hearing discharge rates exceed all discharge rates for impaired hearing(Fig. 5).

−80 −60 −40 −20 0 20 40 60 80

−20

−10

0

10

20

35 dB SPL

−80 −60 −40 −20 0 20 40 60 80

−20

−10

0

10

20

45 dB SPL

−80 −60 −40 −20 0 20 40 60 80

−20

−10

0

10

20

55 dB SPL

−80 −60 −40 −20 0 20 40 60 80

−20

−10

0

10

20

65 dB SPL

Mea

n D

isch

arge

Rat

e D

iffer

ence

(spi

kes/

s)

−80 −60 −40 −20 0 20 40 60 80

−20

−10

0

10

20

75 dB SPL

Azimuth

−80 −60 −40 −20 0 20 40 60 80

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Figure 4: LSO mean differences under the phenomenological model in spikes/s.The following hearing-impaired neural representations are subtracted from thenormal-hearing response, and denoted as follows: Black: unaided; Blue: unlinked WSWDRC; Red: linked WS WDRC.

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Figure 5: An example of patterns of neural response to ILD cues (65 dB SPL puretonestimulus). Normal-hearing spike rates are higher than those in the hearing-impairedmodel for all compression conditions.

C. TIMIT Sentence Results

• No compression condition consistently results in the lowest mean difference scores(Fig. 6).

• Not all phones contain enough energy to trigger WDRC (Fig. 7).

• WDRC benefit varies within phone class; frequency band energy is not the onlyindicator of performance.

• Phone placement is important:– In all stop closures but ‘gcl’, which immediately followed a nasal, spike rates for the

hearing-impaired model exceeded those of the normal-hearing model (8).

no WDRC unlinked WS linked WS unlinked Very Fast linked Very Fast unlinked Slow linked Slow0

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Figure 6: Neither linked nor unlinked WDRC is consistently preferable for speechsounds, although linked is often better. No particular set of compression speeds isconsistently best.

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Figure 7: Energy in some phones of the TIMIT sentence. Solid blue lines indicatethe range of AN fiber CFs modelled, while the dashed blue line indicates the 1500 Hzboundary between low and high frequencies. The red curve denotes the compressionthreshold.

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Figure 8: Phone placement seems to impact WDRC performance. The followinghearing-impaired neural representations are subtracted from the normal-hearingresponse, and denoted as follows:- Black: unaided; Blue: unlinked WDRC; Red: linkedWDRC. Compression speeds are denoted as follows:- Solid lines: WS compression;Dashed lines: Very Fast compression; Dotted lines: Slow compression.

IV. CONCLUSIONS

• The benefit of linked WDRC over unlinked WDRC is clear for puretone stimuli.• In speech stimuli, the benefit is dependent on frequency content, presentation level,phone class, and phone placement within a sentence.

• Adaptive attack and release times may show the most promise as this technologycontinues to develop.

V. ACKNOWLEDGMENTS

This work is supported by NSERC Discovery Grant 261736.

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