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.

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.

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.

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.

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]

LANGUAGE EXPERIENCE SHAPES PROCESSING OF PITCH RELEVANT INFORMATION IN THE HUMAN BRAINSTEM AND AUDITORY CORTEX: ELECTROPHYSIOLOGICAL EVIDENCE.

Pitch is a robust perceptual attribute that plays an important role in speech, language, and music. As such, it provides an analytic window to evaluate how neural activity relevant to pitch undergo transformation from early sensory to later cognitive stages of processing in a well coordinated hierarchical network that is subject to experience-dependent plasticity. We review recent evidence of language experience-dependent effects in pitch processing based on comparisons of native vs. nonnative speakers of a tonal language from electrophysiological recordings in the auditory brainstem and auditory cortex. We present evidence that shows enhanced representation of linguistically-relevant pitch dimensions or features at both the brainstem and cortical levels with a stimulus-dependent preferential activation of the right hemisphere in native speakers of a tone language. We argue that neural representation of pitch-relevant information in the brainstem and early sensory level processing in the auditory cortex is shaped by the perceptual salience of domain-specific features. While both stages of processing are shaped by language experience, neural representations are transformed and fundamentally different at each biological level of abstraction. The representation of pitch relevant information in the brainstem is more fine-grained spectrotemporally as it reflects sustained neural phase-locking to pitch relevant periodicities contained in the stimulus. In contrast, the cortical pitch relevant neural activity reflects primarily a series of transient temporal neural events synchronized to certain temporal attributes of the pitch contour. We argue that experience-dependent enhancement of pitch representation for Chinese listeners most likely reflects an interaction between higher-level cognitive processes and early sensory-level processing to improve representations of behaviorally-relevant features that contribute optimally to perception. It is our view that long-term experience shapes this adaptive process wherein the top-down connections provide selective gating of inputs to both cortical and subcortical structures to enhance neural responses to specific behaviorally-relevant attributes of the stimulus. A theoretical framework for a neural network is proposed involving coordination between local, feedforward, and feedback components that can account for experience-dependent enhancement of pitch representations at multiple levels of the auditory pathway. The ability to record brainstem and cortical pitch relevant responses concurrently may provide a new window to evaluate the online interplay between feedback, feedforward, and local intrinsic components in the hierarchical processing of pitch relevant information.

A functional deficit in the sensorimotor interface component as revealed by oral reading in Thai conduction aphasia.

The contemporary view is that a disruption in phonological encoding underlies the speech production deficit in conduction aphasia. We therefore expect to observe a commonality in phonological errors regardless of task - speaking, reading, or writing. A case report is presented of an oral reading task performed by a Thai conduction aphasic with evidence of localized damage in the left temporoparietal zone. He was instructed to read aloud selections from elementary school materials printed in Thai script at his own pace. A phonological analysis of substitution errors revealed that syllable-initial consonants were more vulnerable to disruption than vowels or tones. Tonal errors were seen to be a secondary consequence of a substitution error involving the syllable-initial consonant. His impaired performance is interpreted as evidence in support of a sensorimotor interface system that converts phonological representations derived from visual orthographic input into articulatory motor representations for speech output.

Relationship between brainstem, cortical and behavioral measures relevant to pitch salience in humans.

Neural representation of pitch-relevant information at both the brainstem and cortical levels of processing is influenced by language or music experience. However, the functional roles of brainstem and cortical neural mechanisms in the hierarchical network for language processing, and how they drive and maintain experience-dependent reorganization are not known. In an effort to evaluate the possible interplay between these two levels of pitch processing, we introduce a novel electrophysiological approach to evaluate pitch-relevant neural activity at the brainstem and auditory cortex concurrently. Brainstem frequency-following responses and cortical pitch responses were recorded from participants in response to iterated rippled noise stimuli that varied in stimulus periodicity (pitch salience). A control condition using iterated rippled noise devoid of pitch was employed to ensure pitch specificity of the cortical pitch response. Neural data were compared with behavioral pitch discrimination thresholds. Results showed that magnitudes of neural responses increase systematically and that behavioral pitch discrimination improves with increasing stimulus periodicity, indicating more robust encoding for salient pitch. Absence of cortical pitch response in the control condition confirms that the cortical pitch response is specific to pitch. Behavioral pitch discrimination was better predicted by brainstem and cortical responses together as compared to each separately. The close correspondence between neural and behavioral data suggest that neural correlates of pitch salience that emerge in early, preattentive stages of processing in the brainstem may drive and maintain with high fidelity the early cortical representations of pitch. These neural representations together contain adequate information for the development of perceptual pitch salience.

Distortion products and their influence on representation of pitch-relevant information in the human brainstem for unresolved harmonic complex tones.

Pitch experiments aimed at evaluating temporal pitch mechanism(s) often utilize complex sounds with only unresolved harmonic components, and a low-pass noise masker to eliminate the potential contribution of audible distortion products to the pitch percept. Herein we examine how: (i) masker induced reduction of neural distortion products (difference tone: DT; and cubic difference tone: CDT) alters the representation of pitch relevant information in the brainstem; and (ii) the pitch salience is altered when distortion products are reduced and/or eliminated. Scalp recorded brainstem frequency following responses (FFR) were recorded in normal hearing individuals using a complex tone with only unresolved harmonics presented in quiet, and in the presence of a low-pass masker at SNRs of +15, +5, and -5 dB. Difference limen for F0 discrimination (F0 DL) was obtained in quiet and in the presence of low-pass noise. Magnitude of DT components (with the exception of components at F0 and 2F0), and the CDT components decreased with increasing masker level. Neural pitch strength decreased with increasing masker level for both the envelope-related (FFR(ENV)) and spectral-related (FFR(SPEC)) phase-locked activity. Finally, F0 DLs increased with decreasing SNRs suggesting poorer F0 discrimination with reduction of the distortion products. Collectively, these findings support the notion that both DT and CDT, as reflected in the FFR(ENV) and FFR(SPEC), respectively, influence both the brainstem representation of pitch relevant information and the pitch salience of the complex sounds.

Linguistic status of timbre influences pitch encoding in the brainstem.

The aim of this experiment is to assess the effects of the linguistic status of timbre on pitch processing in the brainstem. Brainstem frequency following responses were evoked by the Mandarin high-rising lexical tone superimposed on a native vowel quality ([i]), nonnative vowel quality ([œ]), and iterated rippled noise (nonspeech). Results revealed that voice fundamental frequency magnitudes were larger when concomitant with a native vowel quality compared with either nonnative vowel quality or nonspeech timbre. Such experience-dependent effects suggest that subcortical sensory encoding of pitch interacts with timbre in the human brainstem. As a consequence, responses of the perceptual system can be differentially shaped to pitch patterns in relation to the linguistic status of their concomitant timbre.

Musicians demonstrate experience-dependent brainstem enhancement of musical scale features within continuously gliding pitch.

In contrast to language, where pitch patterns consist of continuous and curvilinear contours, musical pitch consists of relatively discrete, stair-stepped sequences of notes. Behavioral and neurophysiological studies suggest that both tone-language and music experience enhance the representation of pitch cues associated with a listener's domain of expertise, e.g., curvilinear pitch in language, discrete scale steps in music. We compared brainstem frequency-following responses (FFRs) of English-speaking musicians (musical pitch experience) and native speakers of Mandarin Chinese (linguistic pitch experience) elicited by rising and falling tonal sweeps that are exemplary of Mandarin tonal contours but uncharacteristic of the pitch patterns typically found in music. In spite of musicians' unfamiliarity with such glides, we find that their brainstem FFRs show enhancement of the stimulus where the curvilinear sweep traverses discrete notes along the diatonic musical scale. This enhancement was note specific in that it was not observed immediately preceding or following the scale tone of interest (passing note). No such enhancements were observed in Chinese listeners. These findings suggest that the musician's brainstem may be differentially tuned by long-term exposure to the pitch patterns inherent to music, extracting pitch in relation to a fixed, hierarchical scale.

Musicians and tone-language speakers share enhanced brainstem encoding but not perceptual benefits for musical pitch.

Behavioral and neurophysiological transfer effects from music experience to language processing are well-established but it is currently unclear whether or not linguistic expertise (e.g., speaking a tone language) benefits music-related processing and its perception. Here, we compare brainstem responses of English-speaking musicians/non-musicians and native speakers of Mandarin Chinese elicited by tuned and detuned musical chords, to determine if enhancements in subcortical processing translate to improvements in the perceptual discrimination of musical pitch. Relative to non-musicians, both musicians and Chinese had stronger brainstem representation of the defining pitches of musical sequences. In contrast, two behavioral pitch discrimination tasks revealed that neither Chinese nor non-musicians were able to discriminate subtle changes in musical pitch with the same accuracy as musicians. Pooled across all listeners, brainstem magnitudes predicted behavioral pitch discrimination performance but considering each group individually, only musicians showed connections between neural and behavioral measures. No brain-behavior correlations were found for tone language speakers or non-musicians. These findings point to a dissociation between subcortical neurophysiological processing and behavioral measures of pitch perception in Chinese listeners. We infer that sensory-level enhancement of musical pitch information yields cognitive-level perceptual benefits only when that information is behaviorally relevant to the listener.

Functional ear (a)symmetry in brainstem neural activity relevant to encoding of voice pitch: a precursor for hemispheric specialization?

Pitch processing is lateralized to the right hemisphere; linguistic pitch is further mediated by left cortical areas. This experiment investigates whether ear asymmetries vary in brainstem representation of pitch depending on linguistic status. Brainstem frequency-following responses (FFRs) were elicited by monaural stimulation of the left and right ear of 15 native speakers of Mandarin Chinese using two synthetic speech stimuli that differ in linguistic status of tone. One represented a native lexical tone (Tone 2: T2); the other, T2', a nonnative variant in which the pitch contour was a mirror image of T2 with the same starting and ending frequencies. Two 40-ms portions of f(0) contours were selected in order to compare two regions (R1, early; R2 late) differing in pitch acceleration rate and perceptual saliency. In R2, linguistic status effects revealed that T2 exhibited a larger degree of FFR rightward ear asymmetry as reflected in f(0) amplitude relative to T2'. Relative to midline (ear asymmetry=0), the only ear asymmetry reaching significance was that favoring left ear stimulation elicited by T2'. By left- and right-ear stimulation separately, FFRs elicited by T2 were larger than T2' in the right ear only. Within T2', FFRs elicited by the earlier region were larger than the later in both ears. Within T2, no significant differences in FFRS were observed between regions in either ear. Collectively, these findings support the idea that origins of cortical processing preferences for perceptually-salient portions of pitch are rooted in early, preattentive stages of processing in the brainstem.