Low-frequency pitch coding: relationships with speech-in-noise and music perception by pediatric populations with typical hearing and cochlear implants.

PURPOSE: This study aimed to investigate the effects of low frequency (LF) pitch perception on speech-in-noise and music perception performance by children with cochlear implants (CIC) and typical hearing (THC). Moreover, the relationships between speech-in-noise and music perception as well as the effects of demographic and audiological factors on present research outcomes were studied. METHODS: The sample consisted of 22 CIC and 20 THC (7-10 years). Harmonic intonation (HI) and disharmonic intonation (DI) tests were used to assess LF pitch perception. Speech perception in quiet (WRSq)/noise (WRSn + 10) were tested with the Italian bisyllabic words for pediatric populations. The Gordon test was used to evaluate music perception (rhythm, melody, harmony, and overall). RESULTS: CIC/THC performance comparisons for LF pitch, speech-in-noise, and all music measures except harmony revealed statistically significant differences with large effect sizes. For the CI group, HI showed statistically significant correlations with melody discrimination. Melody/total Gordon scores were significantly correlated with WRSn + 10. For the overall group, HI/DI showed significant correlations with all music perception measures and WRSn + 10. Hearing thresholds showed significant effects on HI/DI scores. Hearing thresholds and WRSn + 10 scores were significantly correlated; both revealed significant effects on all music perception scores. CI age had significant effects on WRSn + 10, harmony, and total Gordon scores (p < 0.05). CONCLUSION: Such findings confirmed the significant effects of LF pitch perception on complex listening performance. Significant speech-in-noise and music perception correlations were as promising as results from recent studies indicating significant positive effects of music training on speech-in-noise recognition in CIC.

Altered binaural hearing in pre-ataxic and ataxic mutation carriers of spinocerebellar ataxia type 3.

Brainstem degeneration is a prominent feature of spinocerebellar ataxia type 3 (SCA3), involving structures that execute binaural synchronization with microsecond precision. As a consequence, auditory processing may deteriorate during the course of disease. We tested whether the binaural "Huggins pitch" effect is suitable to study the temporal precision of brainstem functioning in SCA3 mutation carriers. We expected that they would have difficulties perceiving Huggins pitch at high frequencies, and that they would show attenuated neuromagnetic responses to Huggins pitch. The upper limit of Huggins pitch perception was psychoacoustically determined in 18 pre-ataxic and ataxic SCA3 mutation carriers and in 18 age-matched healthy controls. Moreover, the cortical N100 response following Huggins pitch onset was acquired by means of magnetoencephalography (MEG). MEG recordings were analyzed using dipole source modeling and comprised a monaural pitch condition and a no-pitch condition with simple binaural correlation changes. Compared with age-matched controls, ataxic but not pre-ataxic SCA3 mutation carriers had significantly lower frequency limits up to which Huggins pitch could be heard. Listeners with lower frequency limits also showed diminished MEG responses to Huggins pitch, but not in the two control conditions. Huggins pitch is a promising tool to assess brainstem functioning in ataxic SCA3 patients. Future studies should refine the psychophysiological setup to capture possible performance decrements also in pre-ataxic mutation carriers. Longitudinal observations will be needed to prove the potential of the assessment of Huggins pitch as a biomarker to track brainstem functioning during the disease course in SCA3.

Correlation Between Objective and Subjective High-Pitched Voice Impairment in Patients After Thyroid Surgery.

Objectives: High-pitched voice impairment (HPVI) is not uncommon in patients without recurrent laryngeal nerve (RLN) or external branch of superior laryngeal nerve (EBSLN) injury after thyroidectomy. This study evaluated the correlation between subjective and objective HPVI in patients after thyroid surgery. Methods: This study analyzed 775 patients without preoperative subjective HPVI and underwent neuromonitored thyroidectomy with normal RLN/EBSLN function. Multi-dimensional voice program, voice range profile and Index of voice and swallowing handicap of thyroidectomy (IVST) were performed during the preoperative(I) period and the immediate(II), short-term(III) and long-term(IV) postoperative periods. The severity of objective HPVI was categorized into four groups according to the decrease in maximum frequency (Fmax): <20%, 20-40%, 40-60%, and >60%. Subjective HPVI was evaluated according to the patient's answers on the IVST. Results: As the severity of objective HPVI increased, patients were significantly more to receive bilateral surgery (p=0.002) and have subjective HPVI (p<0.001), and there was no correlation with IVST scores. Among 211(27.2%) patients with subjective HPVI, patients were significantly more to receive bilateral surgery (p=0.003) and central neck dissection(p<0.001). These patients had very similar trends for Fmax, pitch range, and mean fundamental frequency as patients with 20-40% Fmax decrease (p>0.05) and had higher Jitter, Shimmer, and IVST scores than patients in any of the objective HPVI groups; subjective HPVI lasted until period-IV. Conclusion: The factors that affect a patient's subjective HPVI are complex, and voice stability (Jitter and Shimmer) is no less important than the Fmax level. When patients have subjective HPVI without a significant Fmax decrease after thyroid surgery, abnormal voice stability should be considered and managed. Fmax and IVST scores should be interpreted comprehensively, and surgeons and speech-language pathologists should work together to identify patients with HPVI early and arrange speech therapy for them. Regarding the process of fibrosis formation, anti-adhesive material application and postoperative intervention for HPVI require more future research.

Early cortical processing of pitch height and the role of adaptation and musicality.

Pitch is an important perceptual feature; however, it is poorly understood how its cortical correlates are shaped by absolute vs relative fundamental frequency (f0), and by neural adaptation. In this study, we assessed transient and sustained auditory evoked fields (AEFs) at the onset, progression, and offset of short pitch height sequences, taking into account the listener's musicality. We show that neuromagnetic activity reflects absolute f0 at pitch onset and offset, and relative f0 at transitions within pitch sequences; further, sequences with fixed f0 lead to larger response suppression than sequences with variable f0 contour, and to enhanced offset activity. Musical listeners exhibit stronger f0-related AEFs and larger differences between their responses to fixed vs variable sequences, both within sequences and at pitch offset. The results resemble prominent psychoacoustic phenomena in the perception of pitch contours; moreover, they suggest a strong influence of adaptive mechanisms on cortical pitch processing which, in turn, might be modulated by a listener's musical expertise.

Forward masking with frequency-following response analyses

ABSTRACT Purpose: to analyze forward masking in normally hearing young people, by using frequency-following responses. Methods: the synthetic syllable /da/ was used for the recordings of ten individuals, in the following conditions: /da/ with no masking, and /da/ after 4, 16, 32, and 64 milliseconds of masking. F-test (ANOVA) was applied for repeated measures with the Greenhouse-Geisser correction to compare testing conditions. For significant differences, multiple comparisons (between pairs of conditions) and Bonferroni correction were used. Data normality was verified by applying the Shapiro-Wilk test, and statistical significance was used at 5%. Results: wave latencies of all masking conditions were compared with those of no masking. A latency delay was observed in the transient region of the response (PV and A) in all masking conditions, except for 64 milliseconds. Latency delay also occurred for waves PW, PX, and PY, which corresponded to the sustained region of the response. Conclusion: forward masking was observed, by using frequency-following responses with /da/ syllable in four intervals (4, 16, 32, and 64 ms) of preceding masking. Forward masking was more evident in the transient region of the response than in the sustained one. This study highlights the importance of electrophysiological testing in temporal processing assessment.

Evaluating the effects of smoking on the voice and subjective voice problems using a meta-analysis approach.

The objectives of this study were to identify the effects of smoking on the voice of smokers and present the baseline data for establishing the basis for preventing voice disorders. This study was evaluated using a meta-analysis from studies published between Jan 1, 2000, and Nov 15, 2018. As a result, the final meta-analysis was conducted using nine papers. The standard mean difference was analyzed after dividing the effects of smoking on voice into the pitch (F0), sound quality (jitter, shimmer, and noise to harmonic ratio; NHR), Maximum Phonation Time (MPT), and subjective voice problem. The results showed that there was a significant difference in F0 and MPT. On the other hand, the jitter, shimmer, NHR, and Voice Handicap Index (VHI) had different mean effect size but they were not significantly different. The analysis by sub-function of VHI results showed that the mean effect size was significantly different only in VHI-P (Physical). This study evaluated the effects of smoking on voice using meta-analysis. It was confirmed that smoking had significant and moderate effects on the F0 of voice, MPT, VHI, and physical functions. It is necessary for future meta-analysis studies to conduct randomized controlled experiments or longitudinal studies to confirm the effect sizes of variables.

Transient and sustained processing of musical consonance in auditory cortex and the effect of musicality.

In recent years, electroencephalography and magnetoencephalography (MEG) have both been used to investigate the response in human auditory cortex to musical sounds that are perceived as consonant or dissonant. These studies have typically focused on the transient components of the physiological activity at sound onset, specifically, the N1 wave of the auditory evoked potential and the auditory evoked field, respectively. Unfortunately, the morphology of the N1 wave is confounded by the prominent neural response to energy onset at stimulus onset. It is also the case that the perception of pitch is not limited to sound onset; the perception lasts as long as the note producing it. This suggests that consonance studies should also consider the sustained activity that appears after the transient components die away. The current MEG study shows how energy-balanced sounds can focus the response waves on the consonance-dissonance distinction rather than energy changes and how source modeling techniques can be used to measure the sustained field associated with extended consonant and dissonant sounds. The study shows that musical dyads evoke distinct transient and sustained neuromagnetic responses in auditory cortex. The form of the response depends on both whether the dyads are consonant or dissonant and whether the listeners are musical or nonmusical. The results also show that auditory cortex requires more time for the early transient processing of dissonant dyads than it does for consonant dyads and that the continuous representation of temporal regularity in auditory cortex might be modulated by processes beyond auditory cortex.NEW & NOTEWORTHY We report a magnetoencephalography (MEG) study on transient and sustained cortical consonance processing. Stimuli were long-duration, energy-balanced, musical dyads that were either consonant or dissonant. Spatiotemporal source analysis revealed specific transient and sustained neuromagnetic activity in response to the dyads; in particular, the morphology of the responses was shaped by the dyad's consonance and the listener's musicality. Our results also suggest that the sustained representation of stimulus regularity might be modulated by processes beyond auditory cortex.

Elaboration of an instrument to evaluate the recognition of Brazilian melodies in children / Criação de um instrumento para avaliar o reconhecimento de melodias brasileiras em crianças

Abstract Introduction: There is evidence pointing to the importance of the evaluation of musical perception through objective and subjective instruments. In Brazil, there is a shortage of instruments that evaluates musical perception. Objective: To develop an instrument to evaluate the recognition of traditional Brazilian melodies and investigate the performance of children with typical hearing. Methods: The study was carried out after approval of the research ethics committee (1.198.607). The instrument was developed in software format with website access, using the languages PHP 5.5.12, Javascript, Cascade style sheets and "HTML5"; database "MYSQL 5.6.17" on the "Apache 2.4.9" server. Fifteen melodies of Brazilian folk songs were recorded in piano synthesized timbre, with 12 seconds per melody reproduction and four second intervals between them. A total of 155 schooled children, aged eight to 11 years, of both sexes, with typical hearing participated in the study. The test was performed in a silent room with sound stimuli amplified by a sound box at 65 dBNA, positioned at 0 azimuth, and at one meter from the participant, the notebook was used for children to play with on the screen on the title and illustration of the melody they recognized they were listening to. The responses were recorded on their own database. Results: The instrument titled "Evaluation of recognition of traditional melodies in children" can be run on various devices (computers, notebooks, tablets, mobile phones) and operating systems (Windows, Macintosh, Android, Linux). Access: http://192.185.216.17/ivan/home/login.php by login and password. The most easily recognized melody was "Cai, cai balão" (89%) and the least recognized was "Capelinha de melão" (25.2%). The average time to perform the test was 3′15″. Conclusion: The development and application of the software proved effective for the studied population. This instrument may contribute to the improvement of protocols for the evaluation of musical perception in children with hearing aid and/or cochlear implants users.

Resumo Introdução: Há evidências que apontam para a importância da avaliação da percepção musical através de instrumentos objetivos e subjetivos. No Brasil, há escassez de instrumentos que avaliem a percepção musical. Objetivo: Desenvolver um instrumento para avaliar o reconhecimento de melodias tradicionais brasileiras e investigar o desempenho de crianças com audição típica. Método: O estudo foi desenvolvido após aprovação do comitê de ética em pesquisa (1.198.607). O instrumento foi desenvolvido em formato de software com acesso ao site, com as linguagens de programação PHP 5.5.12, Javascript, Cascade Style Sheets e HTML5; banco de dados MYSQL 5.6.17 no servidor Apache 2.4.9. Quinze melodias de canções folclóricas brasileiras foram gravadas com timbre sintetizado em piano, com 12 segundos de reprodução da melodia e quatro segundos de intervalo entre elas. Participaram do estudo 155 crianças, alfabetizadas, entre oito e 11 anos, de ambos os sexos, com audição típica. O teste foi feito em sala silenciosa com estímulo sonoro amplificado por uma caixa sonora a 65 dB NA, posicionada a 0 azimute e a um metro do participante e o notebook foi usado para que as crianças brincassem na tela com o título e a ilustração da melodia que eles reconheceram que ouviam. As respostas foram registradas em seu próprio banco de dados. Resultados: O instrumento intitulado "Avaliação do reconhecimento de melodias tradicionais em crianças" pode ser executado em vários dispositivos (computadores, notebooks, tablets, telefones celulares) e sistemas operacionais (Windows, Macintosh, Android, Linux). Acesse: http://192.185.216.17/ivan/home/login.php através de login e senha. A melodia mais facilmente reconhecida foi "Cai cai balão" (89%) e a menos reconhecida foi "Capelinha de melão" (25,2%). O tempo médio para fazer o teste foi de 3′15″. Conclusão: O desenvolvimento e a aplicação do software se mostraram eficazes para a população estudada. Esse instrumento pode contribuir para o aprimoramento de protocolos de avaliação da percepção musical em crianças usuárias de próteses auditivas e/ou usuárias de implante coclear.

Perception of lexical stress cued by low-frequency pitch and insights into speech perception in noise for cochlear implant users and normal hearing adults.

PURPOSE: Cochlear implant (CI) users show great difficulty for understanding speech in noise and this fact may partly stem from their poor low-frequency (LF) pitch perception and temporal fine structure (TFS) processing. Clinical assessment of pitch perception is usually based on non-speech tasks. However, linguistically relevant contexts such as lexical stress may better reflect the role of pitch in speech perception, especially for everyday speech where background noise is inevitable. Hence, the study aimed to assess perception of lexical stress cued by LF pitch and TFS cues for CI and normal hearing (NH) listeners, and to investigate relationships with speech perception in noise. METHODS: The low-pass-filtered Word Stress Pattern (WSP-LPF) test was used to evaluate perception of lexical stress cued by LF pitch. Speech perception was assessed with the sentence test with adaptive randomized roving level (STARR) test which presented everyday sentences at low, medium, and high levels in a fluctuating noise to estimate a Speech Reception Threshold. This new test intended to give a realistic estimate for real-world listening. RESULTS: Median WSP-LPF scores in NH (N = 18) and CI listeners (N = 18) were 12.0 Hz and 67.0 Hz, respectively. The corresponding STARR scores were - 9.1 dB and 17.3 dB. Group differences were statistically significant (p < 0.001). Analysis showed significant positive correlations for NH (rs = 0.50) and CI listeners (rs = 0.60). CONCLUSIONS: Present findings reveal stronger correlations than previous studies using non-speech materials, supporting that CI listeners' poor speech perception in noise might be strongly associated with their inability for LF pitch perception and TFS processing.

Modeling and MEG evidence of early consonance processing in auditory cortex.

Pitch is a fundamental attribute of auditory perception. The interaction of concurrent pitches gives rise to a sensation that can be characterized by its degree of consonance or dissonance. In this work, we propose that human auditory cortex (AC) processes pitch and consonance through a common neural network mechanism operating at early cortical levels. First, we developed a new model of neural ensembles incorporating realistic neuronal and synaptic parameters to assess pitch processing mechanisms at early stages of AC. Next, we designed a magnetoencephalography (MEG) experiment to measure the neuromagnetic activity evoked by dyads with varying degrees of consonance or dissonance. MEG results show that dissonant dyads evoke a pitch onset response (POR) with a latency up to 36 ms longer than consonant dyads. Additionally, we used the model to predict the processing time of concurrent pitches; here, consonant pitch combinations were decoded faster than dissonant combinations, in line with the experimental observations. Specifically, we found a striking match between the predicted and the observed latency of the POR as elicited by the dyads. These novel results suggest that consonance processing starts early in human auditory cortex and may share the network mechanisms that are responsible for (single) pitch processing.

Tripolar Stimulation Improves Polyphonic Pitch Detection in Cochlear Implant Users.

OBJECTIVE: Cochlear implant (CI) users struggle with pitch perception, particularly for polyphonic stimuli. Tripolar (TP) stimulation has been proposed as a way to mitigate the broad spread of neural excitation observed in traditional monopolar (MP) stimulation, thereby potentially improving perception of polyphony. STUDY DESIGN: Prospective cohort study. SETTING: Tertiary academic center. PATIENTS: Eleven postlingually deafened adults with Advanced Bionics HiRes 90K CIs. INTERVENTION(S): We performed pitch ranking and polyphonic pitch detection testing under MP and TP configurations. To assess pitch ranking, users were asked to identify the higher pitch between two notes. In polyphonic pitch detection, users were asked to distinguish between single-pitch tones and two-pitch tones. Two-pitch stimuli consisted of one pitch of three base frequencies (392, 523, 740 Hz) and a second pitch between 1 and 12 semitones above the base frequency. MAIN OUTCOME MEASURE: Pitch performance was analyzed as a function of current delivery mode (tripolar vs. monopolar), with smaller semitone interval pitch resolution indicating better performance. RESULTS: In pitch ranking tasks, TP configuration did not confer an advantage over MP stimulation. In polyphonic perception, however, tripolar stimulation improved performance in lower frequencies and resulted in statistically significant (p < 0.05) improvement at the highest base frequency, 740 Hz. CONCLUSIONS: These data suggest that TP configuration may confer an advantage in the perception of polyphonic pitch, which may not be observed in monophonic pitch ranking tasks. Since music is typically polyphonic, such data offer approaches toward improving perception of real-world musical stimuli.

Does passive sound attenuation affect responses to pitch-shifted auditory feedback?

The role of auditory feedback in vocal production has mainly been investigated by altered auditory feedback (AAF) in real time. In response, speakers compensate by shifting their speech output in the opposite direction. Current theory suggests this is caused by a mismatch between expected and observed feedback. A methodological issue is the difficulty to fully isolate the speaker's hearing so that only AAF is presented to their ears. As a result, participants may be presented with two simultaneous signals. If this is true, an alternative explanation is that responses to AAF depend on the contrast between the manipulated and the non-manipulated feedback. This hypothesis was tested by varying the passive sound attenuation (PSA). Participants vocalized while auditory feedback was unexpectedly pitch shifted. The feedback was played through three pairs of headphones with varying amounts of PSA. The participants' responses were not affected by the different levels of PSA. This suggests that across all three headphones, PSA is either good enough to make the manipulated feedback dominant, or differences in PSA are too small to affect the contribution of non-manipulated feedback. Overall, the results suggest that it is important to realize that non-manipulated auditory feedback could affect responses to AAF.

Music Appreciation after Cochlear Implantation in Adult Patients: A Systematic Review.

Objective The cochlear implant (CI) improves quality of life for people who are severely and profoundly deafened, allowing implantees to perceive speech at levels similar to those of individuals with normal hearing. However, patients with CIs generally report a reduced appreciation of music after implantation. We aimed to systematically review the English-language literature for studies evaluating music enjoyment and perception among adult patients with CIs. Data Sources A systematic review of PubMed/MEDLINE, Scopus, Embase, and the Cochrane Library. Review Methods The PRISMA statement was utilized to identify English-language studies reporting music appreciation among adults with CIs. Two independent reviewers performed searches through May 2017. Included studies investigated parameters related to music enjoyment and music perception, including (1) pitch and timbre perception, (2) noise-canceling algorithms, and (3) the presence of dissonant chords, lyrics, or visual cues. Results A total of 508 articles were screened for relevance. Forty-one full-text articles were evaluated, and 18 met final inclusion criteria. Studies used heterogeneous methods of outcome measurement for identifying music appreciation. The outcome measures suggest that rhythm and lyrics are important components of enjoyment. Patients with CIs had difficulty with pitch and timbre perception. Conclusion The heterogeneous outcome measures identified in this systematic review suggest that rhythm and lyrics are important components of enjoyment, while patients with CIs had difficulty with pitch and timbre perception. Because there is no standardized reporting metric for music appreciation among adult patients with CIs, a standardized validated outcome-measuring tool is warranted.

A comparison of speech intonation production and perception abilities of Farsi speaking cochlear implanted and normal hearing children.

INTRODUCTION: Cochlear implant prosthesis facilitates spoken language development and speech comprehension in children with severe-profound hearing loss. However, this prosthesis is limited in encoding information about fundamental frequency and pitch that are essentially for recognition of speech prosody. The purpose of the present study is to investigate the perception and production of intonation in cochlear implant children and comparison with normal hearing children. METHOD: This study carried out on 25 cochlear implanted children and 50 children with normal hearing. First, using 10 action pictures statements and questions sentences were extracted. Fundamental frequency and pitch changes were identified using Praat software. Then, these sentences were judged by 7 adult listeners. In second stage 20 sentences were played for child and he/she determined whether it was in a question form or statement one. RESULTS: Performance of cochlear implanted children in perception and production of intonation was significantly lower than children with normal hearing. The difference between fundamental frequency and pitch changes in cochlear implanted children and children with normal hearing was significant (P < 0/05). Cochlear implanted children performance in perception and production of intonation has significant correlation with child's age surgery and duration of prosthesis use (P < 0/05). DISCUSSION: The findings of the current study show that cochlear prostheses have limited application in facilitating the perception and production of intonation in cochlear implanted children. It should be noted that the child's age at the surgery and duration of prosthesis's use is important in reduction of this limitation. According to these findings, speech and language pathologists should consider intervention of intonation in treatment program of cochlear implanted children.

Perception of Iterated Rippled Noise Periodicity in Cochlear Implant Users.

Pitch perception is more challenging for individuals with cochlear implants (CIs) than normal-hearing subjects because the signal processing by CIs is restricted. Processing and perceiving the periodicity of signals may contribute to pitch perception. Whether individuals with CIs can discern pitch within an iterated rippled noise (IRN) signal is still unclear. In a prospective controlled psychoacoustic study with 34 CI users and 15 normal-hearing control subjects, the difference limen between IRN signals with different numbers of iterations was measured. In 7 CI users and 15 normal-hearing control listeners with single-sided deafness, pitch matching between IRN and harmonic complex tones was measured. The pitch onset response (POR) following signal changes from white noise to IRN was measured electrophysiologically. The CI users could discriminate different numbers of iteration in IRN signals, but worse than normal-hearing listeners. A POR was measured for both normal-hearing subjects and CI users increasing with the pitch salience of the IRN. This indicates that the POR could serve as an objective measure to monitor progress during audioverbal therapy after CI surgery.

Neuromagnetic correlates of voice pitch, vowel type, and speaker size in auditory cortex.

Vowel recognition is largely immune to differences in speaker size despite the waveform differences associated with variation in speaker size. This has led to the suggestion that voice pitch and mean formant frequency (MFF) are extracted early in the hierarchy of hearing/speech processing and used to normalize the internal representation of vowel sounds. This paper presents a magnetoencephalographic (MEG) experiment designed to locate and compare neuromagnetic activity associated with voice pitch, MFF and vowel type in human auditory cortex. Sequences of six sustained vowels were used to contrast changes in the three components of vowel perception, and MEG responses to the changes were recorded from 25 participants. A staged procedure was employed to fit the MEG data with a source model having one bilateral pair of dipoles for each component of vowel perception. This dipole model showed that the activity associated with the three perceptual changes was functionally separable; the pitch source was located in Heschl's gyrus (bilaterally), while the vowel-type and formant-frequency sources were located (bilaterally) just behind Heschl's gyrus in planum temporale. The results confirm that vowel normalization begins in auditory cortex at an early point in the hierarchy of speech processing.

Neural representation of a melodic motif: Effects of polyphonic contexts.

In music, a melodic motif is often played repeatedly in different pitch ranges and at different times. Event-related potential (ERP) studies have shown that the mismatch negativity (MMN) reflects memory trace processing that encodes two separate melodic lines ("voices") with different motifs. Here we investigated whether a single motif presented in two voices is encoded as a single entity or two separate entities, and whether motifs overlapping in time impede or enhance encoding strength. Electroencephalogram (EEG) from 11 musically-trained participants was recorded while they passively listened to sequences of 5-note motifs where the 5th note either descended (standard) or ascended (deviant) relative to the previous note (20% deviant rate). Motifs were presented either in one pitch range, or alternated between two pitch ranges, creating an "upper" and a "lower" voice. Further, motifs were either temporally isolated (silence in between), or temporally concurrent with two tones overlapping. When motifs were temporally isolated, MMN amplitude in the one-pitch-range condition was similar to that in the two-pitch-range upper voice. In contrast, no MMN, but P3a, was observed in the two-pitch-range lower voice. When motifs were temporally concurrent and presented in two pitch ranges, MMN exhibited a more posterior distribution in the upper voice, but again, was absent in the lower voice. These results suggest that motifs presented in two separate voices are not encoded entirely independently, but hierarchically, causing asymmetry between the upper and lower voice encoding even when no simultaneous pitches are presented.

High-Frequency Resonance in the Gerbil Medial Superior Olive.

A high-frequency, subthreshold resonance in the guinea pig medial superior olive (MSO) was recently linked to the efficient extraction of spatial cues from the fine structure of acoustic stimuli. We report here that MSO neurons in gerbil also have resonant properties and, based on our whole-cell recordings and computational modeling, that a low-voltage-gated potassium current, IKLT, underlies the resonance. We show that resonance was lost following dynamic clamp replacement of IKLT with a leak conductance and in the model when voltage-gating of IKLT was suppressed. Resonance was characterized using small amplitude sinusoidal stimuli to generate impedance curves as typically done for linear systems analysis. Extending our study into the nonlinear, voltage-dependent regime, we increased stimulus amplitude and found, experimentally and in simulations, that the subthreshold resonant frequency (242Hz for weak stimuli) increased continuously to the resonant frequency for spiking (285Hz). The spike resonance of these phasic-firing (type III excitable) MSO neurons and of the model is of particular interest also because previous studies of resonance typically involved neurons/models (type II excitable, such as the standard Hodgkin-Huxley model) that can fire tonically for steady inputs. To probe more directly how these resonances relate to MSO neurons as slope-detectors, we presented periodic trains of brief, fast-rising excitatory post-synaptic potentials (EPSCs) to the model. While weak subthreshold EPSC trains were essentially low-pass filtered, resonance emerged as EPSC amplitude increased. Interestingly, for spike-evoking EPSC trains, the threshold amplitude at spike resonant frequency (317Hz) was lower than the single ESPC threshold. Our finding of a frequency-dependent threshold for repetitive brief EPSC stimuli and preferred frequency for spiking calls for further consideration of both subthreshold and suprathreshold resonance to fast and precise temporal processing in the MSO.

Place dependent stimulation rates improve pitch perception in cochlear implantees with single-sided deafness.

In normal hearing, the pitch of an acoustic tone can theoretically be encoded by either the place of stimulation in the cochlea or the corresponding rate of vibration. Thus spectral attributes and temporal fine structure of an acoustic signal are naturally correlated. Cochlear implants (CIs), neural prosthetic devices that restore hearing in the profoundly hearing impaired, currently disregard this mechanism; electrical stimulation is provided at fixed electrode positions with default place independent stimulation rate assignments. This does not account for individual cochlear encoding depending on electrode array placement, variations in insertion depth, and the proximity to nerve fibers. Encoding pitch in such manner delivers limited tonal information. Consequently, music appraisal in CI users is often rated cacophonic while speech perception in quiet is close to normal in top performers. We hypothesize that this limitation in electric stimulation is at least partially due to the mismatch between frequency and place encoding in CIs. In the present study, we determined individual electrode locations by analysis of cochlear radiographic images obtained after surgery and calculated place dependent stimulation rates according to models of the normal tonotopic function. Pitch matching in CI users with single-sided deafness shows that place dependent stimulation rates allow thus far unparalleled restoration of tonotopic pitch perception. Collapsed data of matched pitch frequencies as a function of calculated electrical stimulation rate were well fitted by linear regression (R(2) = 0.878). Sound processing strategies incorporating place dependent stimulation rates are expected to improve pitch perception in CI users.

Implicit and explicit statistical learning of tone sequences across spectral shifts.

We investigated how the statistical learning of auditory sequences is reflected in neuromagnetic responses in implicit and explicit learning conditions. Complex tones with fundamental frequencies (F0s) in a five-tone equal temperament were generated by a formant synthesizer. The tones were subsequently ordered with the constraint that the probability of the forthcoming tone was statistically defined (80% for one tone; 5% for the other four) by the latest two successive tones (second-order Markov chains). The tone sequence consisted of 500 tones and 250 successive tones with a relative shift of F0s based on the same Markov transitional matrix. In explicit and implicit learning conditions, neuromagnetic responses to the tone sequence were recorded from fourteen right-handed participants. The temporal profiles of the N1m responses to the tones with higher and lower transitional probabilities were compared. In the explicit learning condition, the N1m responses to tones with higher transitional probability were significantly decreased compared with responses to tones with lower transitional probability in the latter half of the 500-tone sequence. Furthermore, this difference was retained even after the F0s were relatively shifted. In the implicit learning condition, N1m responses to tones with higher transitional probability were significantly decreased only for the 250 tones following the relative shift of F0s. The delayed detection of learning effects across the sound-spectral shift in the implicit condition may imply that learning may progress earlier in explicit learning conditions than in implicit learning conditions. The finding that the learning effects were retained across spectral shifts regardless of the learning modality indicates that relative pitch processing may be an essential ability for humans.