Attention modulates cortical processing of pitch feedback errors in voice control.
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ABSTRACT:
SUBMITTER: Hu H
PROVIDER: S-EPMC4295089 | biostudies-other | 2015
REPOSITORIES: biostudies-other
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Project description:The control of vocalization is critically dependent on auditory feedback. Here, we determined the human peri-Sylvian speech network that mediates feedback control of pitch using direct cortical recordings. Subjects phonated while a real-time signal processor briefly perturbed their output pitch (speak condition). Subjects later heard the same recordings of their auditory feedback (listen condition). In posterior superior temporal gyrus, a proportion of sites had suppressed responses to normal feedback, whereas other spatially independent sites had enhanced responses to altered feedback. Behaviorally, speakers compensated for perturbations by changing their pitch. Single-trial analyses revealed that compensatory vocal changes were predicted by the magnitude of both auditory and subsequent ventral premotor responses to perturbations. Furthermore, sites whose responses to perturbation were enhanced in the speaking condition exhibited stronger correlations with behavior. This sensorimotor cortical network appears to underlie auditory feedback-based control of vocal pitch in humans.
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Project description:Monitoring voice pitch is a fine-tuned process in daily conversations as conveying accurately the linguistic and affective cues in a given utterance depends on the precise control of phonation and intonation. This monitoring is thought to depend on whether the error is treated as self-generated or externally-generated, resulting in either a correction or inflation of errors. The present study reports on two separate paradigms of adaptation to altered feedback to explore whether participants could behave in a more cohesive manner once the error is of comparable size perceptually. The vocal behavior of normal-hearing and fluent speakers was recorded in response to a personalized size of pitch shift versus a non-specific size, one semitone. The personalized size of shift was determined based on the just-noticeable difference in fundamental frequency (F0) of each participant's voice. Here we show that both tasks successfully demonstrated opposing responses to a constant and predictable F0 perturbation (on from the production onset) but these effects barely carried over once the feedback was back to normal, depicting a pattern that bears some resemblance to compensatory responses. Experiencing a F0 shift that is perceived as self-generated (because it was precisely just-noticeable) is not enough to force speakers to behave more consistently and more homogeneously in an opposing manner. On the contrary, our results suggest that the type of the response as well as the magnitude of the response do not depend in any trivial way on the sensitivity of participants to their own voice pitch. Based on this finding, we speculate that error correction could possibly occur even with a bionic ear, typically even when F0 cues are too subtle for cochlear implant users to detect accurately.
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Project description:The present study describes a technique for analysis of vocal responses to auditory feedback pitch perturbations in which individual trials are first sorted according to response direction and then separately averaged in groups of upward or downward responses. In experiment 1, the stimulus direction was predictable (all upward) but magnitude was randomized between +100, +200, or +500 cents (unpredictable). Results showed that pitch-shift stimuli (PSS) of +100 and +200 cents elicited significantly larger opposing (compensatory) responses than +500 cent stimuli, but no such effect was observed for "following" responses. In experiment 2, subjects were tested in three blocks of trials where for the first two, PSS magnitude and direction were predictable (block 1+100 and block 2-100 cents), and in block 3, the magnitude was predictable (±100 cents) but direction was randomized (upward or downward). Results showed there were slightly more opposing than following responses for predictable PSS direction, but randomized directions led to significantly more opposing than following responses. Results suggest that predictability of stimulus direction and magnitude can modulate vocal responses to feedback pitch perturbations. The function and causes of the opposing and following responses are unknown, but there may be two different neural mechanisms involved in their production.
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Project description:The present study was conducted to test the hypothesis that intrinsic laryngeal muscles are involved in producing voice fundamental frequency (F(0)) responses to perturbations in voice pitch auditory feedback. Electromyography (EMG) recordings of the cricothyroid and thyroarytenoid muscles were made with hooked-wire electrodes, while subjects sustained vowel phonations at three different voice F(0) levels (conversational, high pitch in head register, and falsetto register) and received randomized pitch shifts (±100 or ±300 cents) in their voice auditory feedback. The median latencies from stimulus onset to the peak in the EMG and voice F(0) responses were 167 and 224 ms, respectively. Among the three different F(0) levels, the falsetto register produced compensatory EMG responses that occurred prior to vocal responses and increased along with rising voice F(0) responses and decreased for falling F(0) responses. For the conversational and high voice levels, the EMG response timing was more variable than in the falsetto voice, and changes in EMG activity with relevance to the vocal responses did not follow the consistent trend observed in the falsetto condition. The data from the falsetto condition suggest that both the cricothyroid and thyroarytenoid muscles are involved in generating the compensatory vocal responses to pitch-shifted voice feedback.
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Project description:In single-sided deafness patients fitted with a cochlear implant (CI) in the affected ear and preserved normal hearing in the other ear, acoustic and electric hearing can be directly compared without the need for an external control group. Although poor pitch perception is a crucial limitation when listening through CIs, it remains unclear how exactly the cortical processing of pitch information differs between acoustic and electric hearing. Hence, we separately presented both ears of 20 of these patients with vowel sequences in which the pitch contours were either repetitive or variable, while simultaneously recording functional near-infrared spectroscopy (fNIRS) and EEG data. Overall, the results showed smaller and delayed auditory cortex activity in electric hearing, particularly for the P2 event-related potential component, which appears to reflect the processing of voice pitch information. Both the fNIRS data and EEG source reconstructions furthermore showed that vowel sequences with variable pitch contours evoked additional activity in posterior right auditory cortex in electric but not acoustic hearing. This surprising discrepancy demonstrates, firstly, that the acoustic detail transmitted by CIs is sufficient to distinguish between speech sounds that only vary regarding their pitch information. Secondly, the absence of a condition difference when stimulating the normal-hearing ears suggests a saturation of cortical activity levels following unilateral deafness. Taken together, these results provide strong evidence in favour of using CIs in this patient group.
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Project description:The pitch perturbation technique is a validated technique that has been used for over 30 years to understand how people control their voice. This technique involves altering a person's voice pitch in real-time while they produce a vowel (commonly, a prolonged /a/ sound). Although post-task changes in the voice have been observed in several studies (e.g., a change in mean fo across the duration of the experiment), the potential for using the pitch perturbation technique as a training tool for voice pitch regulation and/or modification has not been explored. The present study examined changes in event related potentials (ERPs) and voice pitch in three groups of subjects due to altered voice auditory feedback following a brief, four-day training period. Participants in the opposing group were trained to change their voice fo in the opposite direction of a pitch perturbation stimulus. Participants in the following group were trained to change their voice fo in the same direction as the pitch perturbation stimulus. Participants in the non-varying group did not voluntarily change their pitch, but instead were asked to hold their voice constant when they heard pitch perturbations. Results showed that all three types of training affected the ERPs and the voice pitch-shift response from pre-training to post-training (i.e., "hold your voice pitch steady" task; an indicator of voice pitch regulation). Across all training tasks, the N1 and P2 components of the ERPs occurred earlier, and the P2 component of the ERPs occurred with larger amplitude post-training. The voice responses also occurred earlier but with a smaller amplitude following training. These results demonstrate that participation in pitch-shifted auditory feedback tasks even for brief periods of time can modulate the automatic tendency to compensate for alterations in voice pitch feedback and has therapeutic potential.
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Project description:Auditory-guided vocal learning is a mechanism that operates both in humans and other animal species making us capable to imitate arbitrary sounds. Both auditory memories and auditory feedback interact to guide vocal learning. This may explain why it is easier for humans to imitate the pitch of a human voice than the pitch of a synthesized sound. In this study, we compared the effects of two different feedback modalities in learning pitch-matching abilities using a synthesized pure tone in 47 participants with no prior music experience. Participants were divided into three groups: a feedback group (N = 15) receiving real-time visual feedback of their pitch as well as knowledge of results; an equal-timbre group (N = 17) receiving additional auditory feedback of the target note with a similar timbre to the instrument being used (i.e., violin or human voice); and a control group (N = 15) practicing without any feedback or knowledge of results. An additional fourth group of violin experts performed the same task for comparative purposes (N = 15). All groups were posteriorly evaluated in a transfer phase. Both experimental groups (i.e., the feedback and equal-timbre groups) improved their intonation abilities with the synthesized sound after receiving feedback. Participants from the equal-timber group seemed as capable as the feedback group of producing the required pitch with the voice after listening to the human voice, but not with the violin (although they also showed improvement). In addition, only participants receiving real-time visual feedback learned and retained in the transfer phase the mapping between the synthesized pitch and its correspondence with the produced vocal or violin pitch. It is suggested that both the effect of an objective external reward, together with the experience of exploring the pitch space with their instrument in an explicit manner, helped participants to understand how to control their pitch production, strengthening their schemas, and favoring retention.
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Project description:In the olfactory system, the olfactory cortex sends glutamatergic projections back to the first stage of olfactory processing, the olfactory bulb (OB). Such corticofugal excitatory circuits - a canonical circuit motif described in all sensory systems- dynamically adjust early sensory processing. Here, we uncover a corticofugal inhibitory feedback to OB, originating from a subpopulation of GABAergic neurons in the anterior olfactory cortex and innervating both local and output OB neurons. In vivo imaging and network modeling showed that optogenetic activation of cortical GABAergic projections drives a net subtractive inhibition of both spontaneous and odor-evoked activity in local as well as output neurons. In output neurons, stimulation of cortical GABAergic feedback enhances separation of population odor responses in tufted cells, but not mitral cells. Targeted pharmacogenetic silencing of cortical GABAergic axon terminals impaired discrimination of similar odor mixtures. Thus, corticofugal GABAergic projections represent an additional circuit motif in cortical feedback control of sensory processing.
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Project description:Pitch patterns, such as melodies, consist of two levels of structure: a global level, comprising the pattern of ups and downs, or contour; and a local level, comprising the precise intervals that make up this contour. An influential neuropsychological model suggests that these two levels of processing are hierarchically linked, with processing of the global structure occurring within the right hemisphere in advance of local processing within the left. However, the predictions of this model and its anatomical basis have not been tested in neurologically normal individuals. The present study used fMRI and required participants to listen to consecutive pitch sequences while performing a same/different one-back task. Sequences, when different, either preserved (local) or violated (global) the contour of the sequence preceding them. When the activations for the local and global conditions were contrasted directly, additional activation was seen for local processing in right planum temporale and posterior superior temporal sulcus (pSTS). The presence of additional activation for local over global processing supports the hierarchical view that the global structure of a pitch sequence acts as a "framework" on which the local detail is subsequently hung. However, the lateralisation of activation seen in the present study, with global processing occurring in left pSTS and local processing occurring bilaterally, differed from that predicted by the neuroanatomical model. A re-examination of the individual lesion data on which the neuroanatomical model is based revealed that the lesion data equally well support the laterality scheme suggested by our data. While the present study supports the hierarchical view of local and global processing, there is an evident need for further research, both in patients and neurologically normal individuals, before an understanding of the functional lateralisation of local and global processing can be considered established.
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Project description:BackgroundSensory suppression occurs when hearing one's self-generated voice, as opposed to passively listening to one's own voice. Quality changes in sensory feedback to the self-generated voice can increase attentional control. These changes affect the self-other voice distinction and might lead to hearing voices in the absence of an external source (ie, auditory verbal hallucinations). However, it is unclear how changes in sensory feedback processing and attention allocation interact and how this interaction might relate to hallucination proneness (HP).Study designParticipants varying in HP self-generated (via a button-press) and passively listened to their voice that varied in emotional quality and certainty of recognition-100% neutral, 60%-40% neutral-angry, 50%-50% neutral-angry, 40%-60% neutral-angry, 100% angry, during electroencephalography (EEG) recordings.Study resultsThe N1 auditory evoked potential was more suppressed for self-generated than externally generated voices. Increased HP was associated with (1) an increased N1 response to the self- compared with externally generated voices, (2) a reduced N1 response for angry compared with neutral voices, and (3) a reduced N2 response to unexpected voice quality in sensory feedback (60%-40% neutral-angry) compared with neutral voices.ConclusionsThe current study highlights an association between increased HP and systematic changes in the emotional quality and certainty in sensory feedback processing (N1) and attentional control (N2) in self-voice production in a nonclinical population. Considering that voice hearers also display these changes, these findings support the continuum hypothesis.
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