Quick Estimation of Minimum Hearing Levels Using a Binaural Multifrequency Stimulus Paradigm: Proof of Concept.

OBJECTIVES: Objective estimation of minimum hearing levels using auditory brainstem responses (ABRs) elicited by single frequency tone-bursts presented monaurally is currently considered the gold standard. However, the data acquisition time to estimate thresholds (for both ears across four audiometric frequencies) using this method usually exceeds the sleep time (ranging between 35 and 49 minutes) in infants below 4 months, thus providing incomplete information of hearing status which in turn delays timely clinical intervention. Alternate approaches using faster rate, or tone-burst trains have not been readily accepted due to additional hardware and software requirements. We propose here a novel binaural multifrequency stimulation paradigm wherein several stimuli of different frequencies are presented binaurally in an interleaved manner. The rationale here is that the proposed paradigm will increase acquisition efficiency, significantly reduce test time, and improve accuracy by incorporating an automatic wave V detection algorithm. It is important to note that this paradigm can be easily implemented in most commercial ABR systems currently used by most clinicians. DESIGN: Using this binaural multifrequency paradigm, ear specific ABRs were recorded in 30 normal-hearing young adults to both tone-bursts, and narrow-band (NB) iChirps at 500, 1000, 2000, and 4000 Hz. Comparison of ABRs elicited by tone-bursts and narrow-band chirps allowed us to determine if NB iChirps elicited a more robust wave V component compared with the tone-bursts. ABR data were characterized by measures of minimum hearing levels; wave V amplitude; and response detectability for two electrode configurations (high forehead-C7; and high forehead-linked mastoids). RESULTS: Consistent with the research literature, wave V response amplitudes were relatively more robust for NB iChirp stimuli compared with tone-burst stimuli. The easier identification and better detectability of wave V for the NB iChirps at lower stimulus levels contributed to their better thresholds compared with tone-burst elicited responses. It is important to note that binaural multifrequency hearing levels close to minimum hearing levels were determined in approximately 22 minutes using this paradigm-appreciably quicker than the 45 to 60 minutes or longer time required for threshold determination using the conventional single frequency method. CONCLUSIONS: Our novel and simple paradigm using either NB iChirps or tone-bursts provides a reliable method to rapidly estimate the minimum hearing levels across audiometric frequencies for both ears. Incorporation of an automatic wave V detection algorithm increases objectivity and further reduce test time and facilitate early hearing identification and intervention.

Frequency-Following Response to Steady-State Vowel in Quiet and Background Noise Among Marching Band Participants With Normal Hearing.

OBJECTIVE: Human studies enrolling individuals at high risk for cochlear synaptopathy (CS) have reported difficulties in speech perception in adverse listening conditions. The aim of this study is to determine if these individuals show a degradation in the neural encoding of speech in quiet and in the presence of background noise as reflected in neural phase-locking to both envelope periodicity and temporal fine structure (TFS). To our knowledge, there are no published reports that have specifically examined the neural encoding of both envelope periodicity and TFS of speech stimuli (in quiet and in adverse listening conditions) among a sample with loud-sound exposure history who are at risk for CS. METHOD: Using scalp-recorded frequency-following response (FFR), the authors evaluated the neural encoding of envelope periodicity (FFRENV) and TFS (FFRTFS) for a steady-state vowel (English back vowel /u/) in quiet and in the presence of speech-shaped noise presented at +5- and 0 dB SNR. Participants were young individuals with normal hearing who participated in the marching band for at least 5 years (high-risk group) and non-marching band group with low-noise exposure history (low-risk group). RESULTS: The results showed no group differences in the neural encoding of either the FFRENV or the first formant (F1) in the FFRTFS in quiet and in noise. Paradoxically, the high-risk group demonstrated enhanced representation of F2 harmonics across all stimulus conditions. CONCLUSIONS: These results appear to be in line with a music experience-dependent enhancement of F2 harmonics. However, due to sound overexposure in the high-risk group, the role of homeostatic central compensation cannot be ruled out. A larger scale data set with different noise exposure background, longitudinal measurements with an array of behavioral and electrophysiological tests is needed to disentangle the nature of the complex interaction between the effects of central compensatory gain and experience-dependent enhancement.

Cortical hemisphere preference and brainstem ear asymmetry reflect experience-dependent functional modulation of pitch.

Temporal attributes of pitch processing at cortical and subcortical levels are differentially weighted and well-coordinated. The question is whether language experience induces functional modulation of hemispheric preference complemented by brainstem ear symmetry for pitch processing. Brainstem frequency-following and cortical pitch responses were recorded concurrently from Mandarin and English participants. A Mandarin syllable with a rising pitch contour was presented to both ears with monaural stimulation. At the cortical level, left ear stimulation in the Chinese group revealed an experience-dependent response for pitch processing in the right hemisphere, consistent with a functionalaccount. The English group revealed a contralateral hemisphere preference consistent with a structuralaccount. At the brainstem level, Chinese participants showed a functional leftward ear asymmetry, whereas English were consistent with a structural account. Overall, language experience modulates both cortical hemispheric preference and brainstem ear asymmetry in a complementary manner to optimize processing of temporal attributes of pitch.

Search for Electrophysiological Indices of Hidden Hearing Loss in Humans: Click Auditory Brainstem Response Across Sound Levels and in Background Noise.

OBJECTIVES: Recent studies in animals indicate that even moderate levels of exposure to noise can damage synaptic ribbons between the inner hair cells and auditory nerve fibers without affecting audiometric thresholds, giving rise to the use of the term "hidden hearing loss" (HHL). Despite evidence across several animal species, there is little consistent evidence for HHL in humans. The aim of the study is to evaluate potential electrophysiological changes specific to individuals at risk for HHL. DESIGN: Participants forming the high-risk experimental group consisted of 28 young normal-hearing adults who participated in marching band for at least 5 years. Twenty-eight age-matched normal-hearing adults who were not part of the marching band and had little or no history of recreational or occupational exposure to loud sounds formed the low-risk control group. Measurements included pure tone audiometry of conventional and high frequencies, distortion product otoacoustic emissions, and electrophysiological measures of auditory nerve and brainstem function as reflected in the click-evoked auditory brainstem response (ABR). In experiment 1, ABRs were recorded in a quiet background across stimulus levels (30-90 dB nHL) presented in 10 dB steps. In experiment 2, the ABR was elicited by a 70 dB nHL click stimulus presented in a quiet background, and in the presence of simultaneous ipsilateral continuous broadband noise presented at 50, 60, and 70 dB SPL using an insert earphone (Etymotic, ER2). RESULTS: There were no differences between the low- and high-risk groups in audiometric thresholds or distortion product otoacoustic emission amplitude. Experiment 1 demonstrated smaller wave-I amplitudes at moderate and high sound levels for high-risk compared to low-risk group with similar wave III and wave V amplitude. Enhanced amplitude ratio V/I, particularly at moderate sound level (60 dB nHL), suggesting central compensation for reduced input from the periphery for high-risk group. The results of experiment 2 show that the decrease in wave I amplitude with increasing background noise level was relatively smaller for the high-risk compared to the low-risk group. However, wave V amplitude reduction was essentially similar for both groups. These results suggest that masking induced wave I amplitude reduction is smaller in individuals at high risk for cochlear synaptopathy. Unlike previous studies, we did not observe a difference in the noise-induced wave V latency shift between low- and high-risk groups. CONCLUSIONS: Results of experiment 1 are consistent with findings in both animal studies (that suggest cochlear synaptopathy involving selective damage of low-spontaneous rate and medium-spontaneous rate fibers), and in several human studies that show changes in a range of ABR metrics that suggest the presence of cochlear synaptopathy. However, without postmortem examination by harvesting human temporal bone (the gold standard for identifying synaptopathy) with different noise exposure background, no direct inferences can be derived for the presence/extent of cochlear synaptopathy in high-risk group with high sound over-exposure history. Results of experiment 2 demonstrate that to the extent response amplitude reflects both the number of neural elements responding and the neural synchrony of the responding elements, the relatively smaller change in response amplitude for the high-risk group would suggest a reduced susceptibility to masking. One plausible mechanism would be that suppressive effects that kick in at moderate to high levels are different in these two groups, particularly at moderate levels of the masking noise. Altogether, a larger scale dataset with different noise exposure background, longitudinal measurements (changes due to recreational over-exposure by studying middle-school to high-school students enrolled in marching band) with an array of behavioral and electrophysiological tests are needed to understand the complex pathogenesis of sound over-exposure damage in normal-hearing individuals.

Human Frequency Following Responses to Vocoded Speech: Amplitude Modulation Versus Amplitude Plus Frequency Modulation.

OBJECTIVES: The most commonly employed speech processing strategies in cochlear implants (CIs) only extract and encode amplitude modulation (AM) in a limited number of frequency channels. proposed a novel speech processing strategy that encodes both frequency modulation (FM) and AM to improve CI performance. Using behavioral tests, they reported better speech, speaker, and tone recognition with this novel strategy than with the AM-alone strategy. Here, we used the scalp-recorded human frequency following responses (FFRs) to examine the differences in the neural representation of vocoded speech sounds with AM alone and AM + FM as the spectral and temporal cues were varied. Specifically, we were interested in determining whether the addition of FM to AM improved the neural representation of envelope periodicity (FFRENV) and temporal fine structure (FFRTFS), as reflected in the temporal pattern of the phase-locked neural activity generating the FFR. DESIGN: FFRs were recorded from 13 normal-hearing, adult listeners in response to the original unprocessed stimulus (a synthetic diphthong /au/ with a 110-Hz fundamental frequency or F0 and a 250-msec duration) and the 2-, 4-, 8- and 16-channel sine vocoded versions of /au/ with AM alone and AM + FM. Temporal waveforms, autocorrelation analyses, fast Fourier Transform, and stimulus-response spectral correlations were used to analyze both the strength and fidelity of the neural representation of envelope periodicity (F0) and TFS (formant structure). RESULTS: The periodicity strength in the FFRENV decreased more for the AM stimuli than for the relatively resilient AM + FM stimuli as the number of channels was increased. Regardless of the number of channels, a clear spectral peak of FFRENV was consistently observed at the stimulus F0 for all the AM + FM stimuli but not for the AM stimuli. Neural representation as revealed by the spectral correlation of FFRTFS was better for the AM + FM stimuli when compared to the AM stimuli. Neural representation of the time-varying formant-related harmonics as revealed by the spectral correlation was also better for the AM + FM stimuli as compared to the AM stimuli. CONCLUSIONS: These results are consistent with previously reported behavioral results and suggest that the AM + FM processing strategy elicited brainstem neural activity that better preserved periodicity, temporal fine structure, and time-varying spectral information than the AM processing strategy. The relatively more robust neural representation of AM + FM stimuli observed here likely contributes to the superior performance on speech, speaker, and tone recognition with the AM + FM processing strategy. Taken together, these results suggest that neural information preserved in the FFR may be used to evaluate signal processing strategies considered for CIs.

Tone language experience-dependent advantage in pitch representation in brainstem and auditory cortex is maintained under reverberation.

Long-term language and music experience enhances neural representation of temporal attributes of pitch in the brainstem and auditory cortex in favorable listening conditions. Herein we examine whether brainstem and cortical pitch mechanisms-shaped by long-term language experience-maintain this advantage in the presence of reverberation-induced degradation in pitch representation. Brainstem frequency following responses (FFR) and cortical pitch responses (CPR) were recorded concurrently from Chinese and English-speaking natives, using a Mandarin word exhibiting a high rising pitch (/yi2/). Stimuli were presented diotically in quiet (Dry), and in the presence of Slight, Mild, and Moderate reverberation conditions. Regardless of language group, the amplitude of both brainstem FFR (F0) and cortical CPR (NaPb) responses decreased with increases in reverberation. Response amplitude for Chinese, however, was larger than English in all reverberant conditions. The Chinese group also exhibited a robust rightward asymmetry at temporal electrode sites (T8 > T7) across stimulus conditions. Regardless of language group, direct comparison of brainstem and cortical responses revealed similar magnitude of change in response amplitude with increasing reverberation. These findings suggest that experience-dependent brainstem and cortical pitch mechanisms provide an enhanced and stable neural representation of pitch-relevant information that is maintained even in the presence of reverberation. Relatively greater degradative effects of reverberation on brainstem (FFR) compared to cortical (NaPb) responses suggest relatively stronger top-down influences on CPRs.

Human frequency following responses to iterated rippled noise with positive and negative gain: Differential sensitivity to waveform envelope and temporal fine-structure.

The perceived pitch of iterated rippled noise (IRN) with negative gain (IRNn) is an octave lower than that of IRN with positive gain (IRNp). IRNp and IRNn have identical waveform envelopes (ENV), but differing stimulus waveform fine structure (TFS), which likely accounts for this perceived pitch difference. Here, we examine whether differences in the temporal pattern of phase-locked activity reflected in the human brainstem Frequency Following Response (FFR) elicited by IRNp and IRNn can account for the differences in perceived pitch for the two stimuli. FFRs using a single onset polarity were measured in 13 normal-hearing, adult listeners in response to IRNp and IRNn stimuli with 2 ms, and 4 ms delay. Autocorrelation functions (ACFs) and Fast Fourier Transforms (FFTs) were used to evaluate the dominant periodicity and spectral pattern (harmonic spacing) in the phase-locked FFR neural activity. For both delays, the harmonic spacing in the spectra corresponded more strongly with the perceived lowering of pitch from IRNp to IRNn, compared to the ACFs. These results suggest that the FFR elicited by a single polarity stimulus reflects phase-locking to both stimulus ENV and TFS. A post-hoc experiment evaluating the FFR phase-locked activity to ENV (FFRENV), and TFS (FFRTFS) elicited by IRNp and IRNn confirmed that only the phase-locked activity to the TFS, reflected in FFRTFS, showed differences in both spectra and ACF that closely matched the pitch difference between the two stimuli. The results of the post-hoc experiment suggests that pitch-relevant information is preserved in the temporal pattern of phase-locked activity and suggests that the differences in stimulus ENV and TFS driving the pitch percept of IRNp and IRNn are preserved in the brainstem neural response. The scalp recorded FFR may provide for a noninvasive analytic tool to evaluate the relative contributions of envelope and temporal fine-structure in the neural representation of complex sounds in humans.

Language experience-dependent advantage in pitch representation in the auditory cortex is limited to favorable signal-to-noise ratios.

Long-term experience enhances neural representation of temporal attributes of pitch in the brainstem and auditory cortex in favorable listening conditions. Herein we examine whether cortical pitch mechanisms shaped by language experience are more resilient to degradation in background noise, and exhibit greater binaural release from masking (BRM). Cortical pitch responses (CPR) were recorded from Mandarin- and English-speaking natives using a Mandarin word exhibiting a high rising pitch (/yi2/). Stimuli were presented diotically in Quiet, and in noise at +5, and 0 dB SNR. CPRs were also recorded in binaural conditions, SONO (where signal and noise were in phase at both ears); or S0Nπ (where signal was in phase and noise 180° out of phase at each ear), using 0 dB SNR. At Fz, both groups showed increase in CPR peak latency and decrease in amplitude with increasing noise level. A language-dependent enhancement of Na-Pb amplitude (Chinese > English) was restricted to Quiet and +5 dB SNR conditions. At T7/T8 electrode sites, Chinese natives exhibited a rightward asymmetry for both CPR components. A language-dependent effect (Chinese > English) was restricted to T8. Regarding BRM, both CPR components showed greater response amplitude for the S0Nπ condition compared to S0N0 across groups. Rightward asymmetry for BRM in the Chinese group indicates experience-dependent recruitment of right auditory cortex. Restriction of the advantage in pitch representation to the quiet and +5 SNR conditions, and the absence of group differences in the binaural release from masking, suggest that language experience affords limited advantage in the neural representation of pitch-relevant information in the auditory cortex under adverse listening conditions.

Language-dependent changes in pitch-relevant neural activity in the auditory cortex reflect differential weighting of temporal attributes of pitch contours.

There remains a gap in our knowledge base about neural representation of pitch attributes that occur between onset and offset of dynamic, curvilinear pitch contours. The aim is to evaluate how language experience shapes processing of pitch contours as reflected in the amplitude of cortical pitch-specific response components. Responses were elicited from three nonspeech, bidirectional (falling-rising) pitch contours representative of Mandarin Tone 2 varying in location of the turning point with fixed onset and offset. At the frontocentral Fz electrode site, Na-Pb and Pb-Nb amplitude of the Chinese group was larger than the English group for pitch contours exhibiting later location of the turning point relative to the one with the earliest location. Chinese listeners' amplitude was also greater than that of English in response to those same pitch contours with later turning points. At lateral temporal sites (T7/T8), Na-Pb amplitude was larger in Chinese listeners relative to English over the right temporal site. In addition, Pb-Nb amplitude of the Chinese group showed a rightward asymmetry. The pitch contour with its turning point located about halfway of total duration evoked a rightward asymmetry regardless of group. These findings suggest that neural mechanisms processing pitch in the right auditory cortex reflect experience-dependent modulation of sensitivity to weighted integration of changes in acceleration rates of rising and falling sections and the location of the turning point.

Human Frequency Following Responses to Vocoded Speech.

OBJECTIVES: Vocoders offer an effective platform to simulate the effects of cochlear implant speech processing strategies in normal-hearing listeners. Several behavioral studies have examined the effects of varying spectral and temporal cues on vocoded speech perception; however, little is known about the neural indices of vocoded speech perception. Here, the scalp-recorded frequency following response (FFR) was used to study the effects of varying spectral and temporal cues on brainstem neural representation of specific acoustic cues, the temporal envelope periodicity related to fundamental frequency (F0) and temporal fine structure (TFS) related to formant and formant-related frequencies, as reflected in the phase-locked neural activity in response to vocoded speech. DESIGN: In experiment 1, FFRs were measured in 12 normal-hearing, adult listeners in response to a steady state English back vowel /u/ presented in an unaltered, unprocessed condition and six sine-vocoder conditions with varying numbers of channels (1, 2, 4, 8, 16, and 32), while the temporal envelope cutoff frequency was fixed at 500 Hz. In experiment 2, FFRs were obtained from 14 normal-hearing, adult listeners in response to the same English vowel /u/, presented in an unprocessed condition and four vocoded conditions where both the temporal envelope cutoff frequency (50 versus 500 Hz) and carrier type (sine wave versus noise band) were varied separately with the number of channels fixed at 8. Fast Fourier Transform was applied to the time waveforms of FFR to analyze the strength of brainstem neural representation of temporal envelope periodicity (F0) and TFS-related peaks (formant structure). RESULTS: Brainstem neural representation of both temporal envelope and TFS cues improved when the number of channels increased from 1 to 4, followed by a plateau with 8 and 16 channels, and a reduction in phase-locking strength with 32 channels. For the sine vocoders, peaks in the FFRTFS spectra corresponded with the low-frequency sine-wave carriers and side band frequencies in the stimulus spectra. When the temporal envelope cutoff frequency increased from 50 to 500 Hz, an improvement was observed in brainstem F0 representation with no change in brainstem representation of spectral peaks proximal to the first formant frequency (F1). There was no significant effect of carrier type (sine- versus noise-vocoder) on brainstem neural representation of F0 cues when the temporal envelope cutoff frequency was 500 Hz. CONCLUSIONS: While the improvement in neural representation of temporal envelope and TFS cues with up to 4 vocoder channels is consistent with the behavioral literature, the reduced neural phase-locking strength noted with even more channels may be because of the narrow bandwidth of each channel as the number of channels increases. Stronger neural representation of temporal envelope cues with higher temporal envelope cutoff frequencies is likely a reflection of brainstem neural phase-locking to F0-related periodicity fluctuations preserved in the 500-Hz temporal envelopes, which are unavailable in the 50-Hz temporal envelopes. No effect of temporal envelope cutoff frequency was seen for neural representation of TFS cues, suggesting that spectral side band frequencies created by the 500-Hz temporal envelopes did not improve neural representation of F1 cues over the 50-Hz temporal envelopes. Finally, brainstem F0 representation was not significantly affected by carrier type with a temporal envelope cutoff frequency of 500 Hz, which is inconsistent with previous results of behavioral studies examining pitch perception of vocoded stimuli.

Differential sensitivity to changes in pitch acceleration in the auditory brainstem and cortex.

The cortical pitch-specific response (CPR) is differentially sensitive to pitch contours varying in rate of acceleration-time-variant Mandarin Tone2 (T2) versus constant, linear rising ramp (Linear)-as a function of language experience (Krishnan, Gandour, & Suresh, 2014). CPR and brainstem frequency following response (FFR) data were recorded concurrently from native Mandarin listeners using the same stimuli. Results showed that T2 elicited larger responses than Linear at both cortical and brainstem levels (CPR: Na-Pb, Pb-Nb; FFR). However, Pb-Nb exhibited a larger difference in magnitude between T2 and Linear than either Na-Pb or FFR. This finding highlights differential weighting of brain responses elicited by a specific temporal attribute of pitch. Consistent with the notion of a distributed, integrated hierarchical pitch processing network, temporal attributes of pitch are differentially weighted by subcortical and cortical level processing.

Changes in pitch height elicit both language-universal and language-dependent changes in neural representation of pitch in the brainstem and auditory cortex.

Language experience shapes encoding of pitch-relevant information at both brainstem and cortical levels of processing. Pitch height is a salient dimension that orders pitch from low to high. Herein we investigate the effects of language experience (Chinese, English) in the brainstem and cortex on (i) neural responses to variations in pitch height, (ii) presence of asymmetry in cortical pitch representation, and (iii) patterns of relative changes in magnitude of pitch height between these two levels of brain structure. Stimuli were three nonspeech homologs of Mandarin Tone 2 varying in pitch height only. The frequency-following response (FFR) and the cortical pitch-specific response (CPR) were recorded concurrently. At the Fz-linked T7/T8 site, peak latency of Na, Pb, and Nb decreased with increasing pitch height for both groups. Peak-to-peak amplitude of Na-Pb and Pb-Nb increased with increasing pitch height across groups. A language-dependent effect was restricted to Na-Pb; the Chinese had larger amplitude than the English group. At temporal sites (T7/T8), the Chinese group had larger amplitude, as compared to English, across stimuli, but also limited to the Na-Pb component and right temporal site. In the brainstem, F0 magnitude decreased with increasing pitch height; Chinese had larger magnitude across stimuli. A comparison of CPR and FFR responses revealed distinct patterns of relative changes in magnitude common to both groups. CPR amplitude increased and FFR amplitude decreased with increasing pitch height. Experience-dependent effects on CPR components vary as a function of neural sensitivity to pitch height within a particular temporal window (Na-Pb). Differences between the auditory brainstem and cortex imply distinct neural mechanisms for pitch extraction at both levels of brain structure.

Language-experience plasticity in neural representation of changes in pitch salience.

Neural representation of pitch-relevant information at the brainstem and cortical levels of processing is influenced by language experience. A well-known attribute of pitch is its salience. Brainstem frequency following responses and cortical pitch specific responses, recorded concurrently, were elicited by a pitch salience continuum spanning weak to strong pitch of a dynamic, iterated rippled noise pitch contour-homolog of a Mandarin tone. Our aims were to assess how language experience (Chinese, English) affects i) enhancement of neural activity associated with pitch salience at brainstem and cortical levels, ii) the presence of asymmetry in cortical pitch representation, and iii) patterns of relative changes in magnitude along the pitch salience continuum. Peak latency (Fz: Na, Pb, and Nb) was shorter in the Chinese than the English group across the continuum. Peak-to-peak amplitude (Fz: Na-Pb, Pb-Nb) of the Chinese group grew larger with increasing pitch salience, but an experience-dependent advantage was limited to the Na-Pb component. At temporal sites (T7/T8), the larger amplitude of the Chinese group across the continuum was both limited to the Na-Pb component and the right temporal site. At the brainstem level, F0 magnitude gets larger as you increase pitch salience, and it too reveals Chinese superiority. A direct comparison of cortical and brainstem responses for the Chinese group reveals different patterns of relative changes in magnitude along the pitch salience continuum. Such differences may point to a transformation in pitch processing at the cortical level presumably mediated by local sensory and/or extrasensory influence overlaid on the brainstem output.

Human Frequency Following Response: Neural Representation of Envelope and Temporal Fine Structure in Listeners with Normal Hearing and Sensorineural Hearing Loss.

OBJECTIVE: Listeners with sensorineural hearing loss (SNHL) typically experience reduced speech perception, which is not completely restored with amplification. This likely occurs because cochlear damage, in addition to elevating audiometric thresholds, alters the neural representation of speech transmitted to higher centers along the auditory neuroaxis. While the deleterious effects of SNHL on speech perception in humans have been well-documented using behavioral paradigms, our understanding of the neural correlates underlying these perceptual deficits remains limited. Using the scalp-recorded frequency following response (FFR), the authors examine the effects of SNHL and aging on subcortical neural representation of acoustic features important for pitch and speech perception, namely the periodicity envelope (F0) and temporal fine structure (TFS; formant structure), as reflected in the phase-locked neural activity generating the FFR. DESIGN: FFRs were obtained from 10 listeners with normal hearing (NH) and 9 listeners with mild-moderate SNHL in response to a steady-state English back vowel /u/ presented at multiple intensity levels. Use of multiple presentation levels facilitated comparisons at equal sound pressure level (SPL) and equal sensation level. In a second follow-up experiment to address the effect of age on envelope and TFS representation, FFRs were obtained from 25 NH and 19 listeners with mild to moderately severe SNHL to the same vowel stimulus presented at 80 dB SPL. Temporal waveforms, Fast Fourier Transform and spectrograms were used to evaluate the magnitude of the phase-locked activity at F0 (periodicity envelope) and F1 (TFS). RESULTS: Neural representation of both envelope (F0) and TFS (F1) at equal SPLs was stronger in NH listeners compared with listeners with SNHL. Also, comparison of neural representation of F0 and F1 across stimulus levels expressed in SPL and sensation level (accounting for audibility) revealed that level-related changes in F0 and F1 magnitude were different for listeners with SNHL compared with listeners with NH. Furthermore, the degradation in subcortical neural representation was observed to persist in listeners with SNHL even when the effects of age were controlled for. CONCLUSIONS: Overall, our results suggest a relatively greater degradation in the neural representation of TFS compared with periodicity envelope in individuals with SNHL. This degraded neural representation of TFS in SNHL, as reflected in the brainstem FFR, may reflect a disruption in the temporal pattern of phase-locked neural activity arising from altered tonotopic maps and/or wider filters causing poor frequency selectivity in these listeners. Finally, while preliminary results indicate that the deleterious effects of SNHL may be greater than age-related degradation in subcortical neural representation, the lack of a balanced age-matched control group in this study does not permit us to completely rule out the effects of age on subcortical neural representation.

Pitch processing of dynamic lexical tones in the auditory cortex is influenced by sensory and extrasensory processes.

The aim is to evaluate how language experience (Chinese, English) shapes processing of pitch contours as reflected in the amplitude of cortical pitch response components. Responses were elicited from three dynamic curvilinear nonspeech stimuli varying in pitch direction and location of peak acceleration: Mandarin lexical Tone 2 (rising) and Tone 4 (falling), and a flipped variant of Tone 2, Tone 2' (nonnative). At temporal sites (T7/T8), Chinese listeners' Na-Pb response amplitudes to Tones 2 and 4 were greater than those of English listeners in the right hemisphere only; a rightward asymmetry for Tones 2 and 4 was restricted to the Chinese group. In common to both Fz-to-linked T7/T8 and T7/T8 electrode sites, the stimulus pattern (Tones 2 and 4 > Tone 2') was found in the Chinese group only. As reflected by Pb-Nb at Fz, Chinese subjects' amplitudes were larger than those of English subjects in response to Tones 2 and 4, and Tones 2 and 4 were larger than Tone 2', whereas for English subjects, Tone 2 was larger than Tone 2' and Tone 4. At frontal electrode sites (F3/F4), regardless of component or hemisphere, Chinese subjects' responses were larger in amplitude than those of English subjects across stimuli. For either group, responses to Tones 2 and 4 were larger than Tone 2'. No hemispheric asymmetry was observed at the frontal electrode sites. These findings demonstrate that cortical pitch response components are differentially modulated by experience-dependent, temporally distinct but functionally overlapping, weighting of sensory and extrasensory effects on pitch processing of lexical tones in the right temporal lobe and, more broadly, are consistent with a distributed hierarchical predictive coding process.

Language experience enhances early cortical pitch-dependent responses.

Pitch processing at cortical and subcortical stages of processing is shaped by language experience. We recently demonstrated that specific components of the cortical pitch response (CPR) index the more rapidly-changing portions of the high rising Tone 2 of Mandarin Chinese, in addition to marking pitch onset and sound offset. In this study, we examine how language experience (Mandarin vs. English) shapes the processing of different temporal attributes of pitch reflected in the CPR components using stimuli representative of within-category variants of Tone 2. Results showed that the magnitude of CPR components (Na-Pb and Pb-Nb) and the correlation between these two components and pitch acceleration were stronger for the Chinese listeners compared to English listeners for stimuli that fell within the range of Tone 2 citation forms. Discriminant function analysis revealed that the Na-Pb component was more than twice as important as Pb-Nb in grouping listeners by language affiliation. In addition, a stronger stimulus-dependent, rightward asymmetry was observed for the Chinese group at the temporal, but not frontal, electrode sites. This finding may reflect selective recruitment of experience-dependent, pitch-specific mechanisms in right auditory cortex to extract more complex, time-varying pitch patterns. Taken together, these findings suggest that long-term language experience shapes early sensory level processing of pitch in the auditory cortex, and that the sensitivity of the CPR may vary depending on the relative linguistic importance of specific temporal attributes of dynamic pitch.

Cortical pitch response components show differential sensitivity to native and nonnative pitch contours.

The aim of this study is to evaluate how nonspeech pitch contours of varying shape influence latency and amplitude of cortical pitch-specific response (CPR) components differentially as a function of language experience. Stimuli included time-varying, high rising Mandarin Tone 2 (T2) and linear rising ramp (Linear), and steady-state (Flat). Both the latency and magnitude of CPR components were differentially modulated by (i) the overall trajectory of pitch contours (time-varying vs. steady-state), (ii) their pitch acceleration rates (changing vs. constant), and (iii) their linguistic status (lexical vs. non-lexical). T2 elicited larger amplitude than Linear in both language groups, but size of the effect was larger in Chinese than English. The magnitude of CPR components elicited by T2 were larger for Chinese than English at the right temporal electrode site. Using the CPR, we provide evidence in support of experience-dependent modulation of dynamic pitch contours at an early stage of sensory processing.

Cortical pitch response components index stimulus onset/offset and dynamic features of pitch contours.

Voice pitch is an important information-bearing component of language that is subject to experience dependent plasticity at both early cortical and subcortical stages of processing. We have already demonstrated that pitch onset component (Na) of the cortical pitch response (CPR) is sensitive to flat pitch and its salience … CPR responses from Chinese listeners were elicited by three citation forms varying in pitch acceleration and duration. Results showed that the pitch onset component (Na) was invariant to changes in acceleration. In contrast, Na­Pb and Pb­Nb showed a systematic decrease in the interpeak latency and decrease in amplitude with increase in pitch acceleration that followed the time course of pitch change across the three stimuli. A strong correlation with pitch acceleration was observed for these two components only ­ a putative index of pitch-relevant neural activity associated with the more rapidly-changing portions of the pitch contour. Pc­Nc marks unambiguously the stimulus offset … and their functional roles as related to sensory and cognitive properties of the stimulus. [Corrected]

Phase locked neural activity in the human brainstem predicts preference for musical consonance.

When musical notes are combined to make a chord, the closeness of fit of the combined spectrum to a single harmonic series (the 'harmonicity' of the chord) predicts the perceived consonance (how pleasant and stable the chord sounds; McDermott, Lehr, & Oxenham, 2010). The distinction between consonance and dissonance is central to Western musical form. Harmonicity is represented in the temporal firing patterns of populations of brainstem neurons. The current study investigates the role of brainstem temporal coding of harmonicity in the perception of consonance. Individual preference for consonant over dissonant chords was measured using a rating scale for pairs of simultaneous notes. In order to investigate the effects of cochlear interactions, notes were presented in two ways: both notes to both ears or each note to different ears. The electrophysiological frequency following response (FFR), reflecting sustained neural activity in the brainstem synchronised to the stimulus, was also measured. When both notes were presented to both ears the perceptual distinction between consonant and dissonant chords was stronger than when the notes were presented to different ears. In the condition in which both notes were presented to the both ears additional low-frequency components, corresponding to difference tones resulting from nonlinear cochlear processing, were observable in the FFR effectively enhancing the neural harmonicity of consonant chords but not dissonant chords. Suppressing the cochlear envelope component of the FFR also suppressed the additional frequency components. This suggests that, in the case of consonant chords, difference tones generated by interactions between notes in the cochlea enhance the perception of consonance. Furthermore, individuals with a greater distinction between consonant and dissonant chords in the FFR to individual harmonics had a stronger preference for consonant over dissonant chords. Overall, the results provide compelling evidence for the role of neural temporal coding in the perception of consonance, and suggest that the representation of harmonicity in phase locked neural firing drives the perception of consonance.