Pitch Auditory Feedback Research Articles

Task-Dependent Modulation of Auditory Feedback Control of Vocal Intensity

Auditory feedback control of fundamental frequency (fo) is modulated in a task-dependent manner. When voice pitch auditory feedback perturbations are applied in sentence versus sustained-vowel production, larger and faster vocal fo responses are measured in sentence production. This task-dependency reflects the scaling of auditory targets for pitch for the precision required in each speech task. When the range for the pitch auditory target is scaled down for precision (as in the sentence-production task), a greater degree of mismatch is detected from the feedback perturbation and a larger vocal response is measured. The purpose of this study was to determine whether auditory feedback control of vocal intensity is also modulated in a task-dependent manner similar to the control of vocal pitch. Twenty-five English speakers produced repetitions of a sentence and a sustained vowel while hearing their voice auditory feedback briefly perturbed in loudness (+/- 3 or 6 dB SPL, 200 ms duration). The resulting vocal intensity responses were measured, and response magnitudes were robustly larger in the sentence (mean: 1.96 dB) than vowel production (mean: 0.89 dB). Additionally, response magnitudes increased as a function of perturbation magnitude only in sentence production for downward perturbations but decreased in magnitude by perturbation magnitude for upward perturbations. Peak response latencies were robustly shorter in sentence (mean: 184.94 ms) than in vowel production (mean: 214.92 ms). Overall, these results support the hypothesis that auditory feedback control of pitch and loudness are modulated by task and that both pitch and loudness auditory targets are scaled for the precision required for the speaking task.

Semantic focus mediates pitch auditory feedback control in phrasal prosody

ABSTRACT This study investigated the effect of semantic focus on pitch auditory feedback control in the production of phrasal prosody through an experiment using pitch-shifted auditory feedback. We hypothesized that pitch-shift responses would be mediated by semantic focus because highly informative focus types, such as corrective focus, impose more specific constraints on the prosodic form of a phrase and require greater consistency in the production of pitch excursions compared to sentences with no such focus elements. Twenty-eight participants produced sentences with and without corrective focus while their auditory feedback was briefly and unexpectedly perturbed in pitch by +/−200 cents at the start of the sentence. The magnitude and latency of the reflexive pitch-shift responses were measured as a reflection of auditory feedback control. Our results matched our prediction that corrective focus would elicit larger pitch-shift responses, supporting our hypothesis that auditory feedback control is mediated by semantic focus.

The Effect of Pitch Auditory Feedback Perturbations on the Production of Anticipatory Phrasal Prominence and Boundary.

Purpose In this study, we investigated how the direction and timing of a perturbation in voice pitch auditory feedback during phrasal production modulated the magnitude and latency of the pitch-shift reflex as well as the scaling of acoustic production of anticipatory intonation targets for phrasal prominence and boundary. Method Brief pitch auditory feedback perturbations (±200 cents for 200-ms duration) were applied during the production of a target phrase on the first or the second word of the phrase. To replicate previous work, we first measured the magnitude and latency of the pitch-shift reflex as a function of the direction and timing of the perturbation within the phrase. As a novel approach, we also measured the adjustment in the production of the phrase-final prominent word as a function of perturbation direction and timing by extracting the acoustic correlates of pitch, loudness, and duration. Results The pitch-shift reflex was greater in magnitude after perturbations on the first word of the phrase, replicating the results from Mandarin speakers in an American English-speaking population. Additionally, the production of the phrase-final prominent word was acoustically enhanced (lengthened vowel duration and increased intensity and fundamental frequency) after perturbations earlier in the phrase, but more so after perturbations on the first word in the phrase. Conclusion The timing of the pitch perturbation within the phrase modulated both the magnitude of the pitch-shift reflex and the production of the prominent word, supporting our hypothesis that speakers use auditory feedback to correct for immediate production errors and to scale anticipatory intonation targets during phrasal production.

Dopamine Depletion Affects Vocal Acoustics and Disrupts Sensorimotor Adaptation in Songbirds.

Dopamine is hypothesized to convey error information in reinforcement learning tasks with explicit appetitive or aversive cues. However, during motor skill learning feedback signals arise from an animal’s evaluation of sensory feedback resulting from its own behavior, rather than any external reward or punishment. It has previously been shown that intact dopaminergic signaling from the ventral tegmental area/substantia nigra pars compacta (VTA/SNc) complex is necessary for vocal learning when songbirds modify their vocalizations to avoid hearing distorted auditory feedback (playbacks of white noise). However, it remains unclear whether dopaminergic signaling underlies vocal learning in response to more naturalistic errors (pitch-shifted feedback delivered via headphones). We used male Bengalese finches (Lonchura striata var. domestica) to test the hypothesis that the necessity of dopamine signaling is shared between the two types of learning. We combined 6-hydroxydopamine (6-OHDA) lesions of dopaminergic terminals within Area X, a basal ganglia nucleus critical for song learning, with a headphones learning paradigm that shifted the pitch of auditory feedback and compared their learning to that of unlesioned controls. We found that 6-OHDA lesions affected song behavior in two ways. First, over a period of days lesioned birds systematically lowered their pitch regardless of the presence or absence of auditory errors. Second, 6-OHDA lesioned birds also displayed severe deficits in sensorimotor learning in response to pitch-shifted feedback. Our results suggest roles for dopamine in both motor production and auditory error processing, and a shared mechanism underlying vocal learning in response to both distorted and pitch-shifted auditory feedback.

The effect of pitch of auditory feedback on synchronized tapping

The effect of pitch of auditory feedback on synchronized tapping

Bioelectrical brain effects of one's own voice identification in pitch of voice auditory feedback.

Control of voice fundamental frequency (F0) relies in part on comparison of the intended F0 level and auditory feedback. This comparison impacts "sense of agency", or SoA, commonly defined as being the agent of one's own actions and plays a key role for self-awareness and social interactions. SoA is aberrant in several psychiatric disorders. Knowledge about brain activity reflecting SoA can be used in clinical practice for these disorders. It was shown that perception of voice feedback as one's own voice, reflecting the recognition of SoA, alters auditory sensory processing. Using a voice perturbation paradigm we contrasted vocal and bioelectrical brain responses to auditory stimuli that differed in magnitude: 100 and 400 cents. Results suggest the different magnitudes were perceived as a pitch error in self-vocalization (100 cents) or as a pitch shift generated externally (400 cents). Vocalizations and neural responses to changes in pitch of self-vocalization were defined as those made to small magnitude pitch-shifts (100 cents) and which did not show differential neural responses to upward versus downward changes in voice pitch auditory feedback. Vocal responses to large magnitude pitch shifts (400 cents) were smaller than those made to small pitch shifts, and neural responses differed according to upwards versus downward changes in pitch. Our results suggest that the presence of SoA for self-produced sounds may modify bioelectrical brain responses reflecting differences in auditory processing of the direction of a pitch shift. We suggest that this modification of bioelectrical response can be used as a biological index of SoA. Possible neuronal mechanisms of this modification of bioelectrical brain response are discussed.

A Bayesian Account of Vocal Adaptation to Pitch-Shifted Auditory Feedback.

Motor systems are highly adaptive. Both birds and humans compensate for synthetically induced shifts in the pitch (fundamental frequency) of auditory feedback stemming from their vocalizations. Pitch-shift compensation is partial in the sense that large shifts lead to smaller relative compensatory adjustments of vocal pitch than small shifts. Also, compensation is larger in subjects with high motor variability. To formulate a mechanistic description of these findings, we adapt a Bayesian model of error relevance. We assume that vocal-auditory feedback loops in the brain cope optimally with known sensory and motor variability. Based on measurements of motor variability, optimal compensatory responses in our model provide accurate fits to published experimental data. Optimal compensation correctly predicts sensory acuity, which has been estimated in psychophysical experiments as just-noticeable pitch differences. Our model extends the utility of Bayesian approaches to adaptive vocal behaviors.

The Effects of Pitch Shifts on Delay-Induced Changes in Vocal Sequencing in a Songbird.

Like human speech, vocal behavior in songbirds depends critically on auditory feedback. In both humans and songbirds, vocal skills are acquired by a process of imitation whereby current vocal production is compared to an acoustic target. Similarly, performance in adulthood relies strongly on auditory feedback, and online manipulations of auditory signals can dramatically alter acoustic production even after vocalizations have been well learned. Artificially delaying auditory feedback can disrupt both speech and birdsong, and internal delays in auditory feedback have been hypothesized as a cause of vocal dysfluency in persons who stutter. Furthermore, in both song and speech, online shifts of the pitch (fundamental frequency) of auditory feedback lead to compensatory changes in vocal pitch for small perturbations, but larger pitch shifts produce smaller changes in vocal output. Intriguingly, large pitch shifts can partially restore normal speech in some dysfluent speakers, suggesting that the effects of auditory feedback delays might be ameliorated by online pitch manipulations. Although birdsong provides a promising model system for understanding speech production, the interactions between sensory feedback delays and pitch shifts have not yet been assessed in songbirds. To investigate this, we asked whether the addition of a pitch shift modulates delay-induced changes in Bengalese finch song, hypothesizing that pitch shifts would reduce the effects of feedback delays. Compared with the effects of delays alone, combined delays and pitch shifts resulted in a significant reduction in behavioral changes in one type of sequencing (branch points) but not another (distribution of repeated syllables).

Sensory Processing: Advances in Understanding Structure and Function of Pitch-Shifted Auditory Feedback in Voice Control

The pitch-shift paradigm has become a widely used method for studying the role of voice pitch auditory feedback in voice control. This paradigm introduces small, brief pitch shifts in voice auditory feedback to vocalizing subjects. The perturbations trigger a reflexive mechanism that counteracts the change in pitch. The underlying mechanisms of the vocal responses are thought to reflect a negative feedback control system that is similar to constructs developed to explain other forms of motor control. Another use of this technique requires subjects to voluntarily change the pitch of their voice when they hear a pitch shift stimulus. Under these conditions, short latency responses are produced that change voice pitch to match that of the stimulus. The pitch-shift technique has been used with magnetoencephalography (MEG) and electroencephalography (EEG) recordings, and has shown that at vocal onset there is normally a suppression of neural activity related to vocalization. However, if a pitch-shift is also presented at voice onset, there is a cancellation of this suppression, which has been interpreted to mean that one way in which a person distinguishes self-vocalization from vocalization of others is by a comparison of the intended voice and the actual voice. Studies of the pitch shift reflex in the fMRI environment show that the superior temporal gyrus (STG) plays an important role in the process of controlling voice F0 based on auditory feedback. Additional studies using fMRI for effective connectivity modeling show that the left and right STG play critical roles in correcting for an error in voice production. While both the left and right STG are involved in this process, a feedback loop develops between left and right STG during perturbations, in which the left to right connection becomes stronger, and a new negative right to left connection emerges along with the emergence of other feedback loops within the cortical network tested.

Vocal and Neural Responses to Unexpected Changes in Voice Pitch Auditory Feedback During Register Transitions

It is known that singers are able to control their voice to maintain a relatively constant vocal quality while transitioning between vocal registers; however, the neural mechanisms underlying this effect are not understood. It was hypothesized that greater attention to the acoustical feedback of the voice and increased control of the vocal musculature during register transitions compared with singing within a register would be represented as neurological differences in event-related potentials. Nine singers sang musical notes at the high end of the modal register (the boundary between the modal and the head/falsetto registers) and at the low end (the boundary between the modal and the fry/pulse registers). While singing, the pitch of the voice auditory feedback was unexpectedly shifted either into the adjacent register ("toward" the register boundary) or within the modal register ("away from" the boundary). Singers were instructed to maintain a constant pitch and ignore any changes to their voice feedback. Vocal response latencies and magnitude of the accompanying N1 and P2 event-related potentials were greatest at the lower (modal-to-fry) boundary when the pitch shift carried the subjects' voices into the fry register as opposed to remaining within the modal register. These findings suggest that when a singer lowers the pitch of his or her voice such that it enters the fry register from the modal register, there is increased sensory-motor control of the voice, reflected as increased magnitude of the neural potentials to help minimize qualitative changes in the voice.

Sensory–motor networks involved in speech production and motor control: An fMRI study

Speaking is one of the most complex motor behaviors developed to facilitate human communication. The underlying neural mechanisms of speech involve sensory–motor interactions that incorporate feedback information for online monitoring and control of produced speech sounds. In the present study, we adopted an auditory feedback pitch perturbation paradigm and combined it with functional magnetic resonance imaging (fMRI) recordings in order to identify brain areas involved in speech production and motor control. Subjects underwent fMRI scanning while they produced a steady vowel sound /a/ (speaking) or listened to the playback of their own vowel production (playback). During each condition, the auditory feedback from vowel production was either normal (no perturbation) or perturbed by an upward (+600 cents) pitch-shift stimulus randomly. Analysis of BOLD responses during speaking (with and without shift) vs. rest revealed activation of a complex network including bilateral superior temporal gyrus (STG), Heschl's gyrus, precentral gyrus, supplementary motor area (SMA), Rolandic operculum, postcentral gyrus and right inferior frontal gyrus (IFG). Performance correlation analysis showed that the subjects produced compensatory vocal responses that significantly correlated with BOLD response increases in bilateral STG and left precentral gyrus. However, during playback, the activation network was limited to cortical auditory areas including bilateral STG and Heschl's gyrus. Moreover, the contrast between speaking vs. playback highlighted a distinct functional network that included bilateral precentral gyrus, SMA, IFG, postcentral gyrus and insula. These findings suggest that speech motor control involves feedback error detection in sensory (e.g. auditory) cortices that subsequently activate motor-related areas for the adjustment of speech parameters during speaking.

Vocal generalization depends on gesture identity and sequence.

Generalization, the brain's ability to transfer motor learning from one context to another, occurs in a wide range of complex behaviors. However, the rules of generalization in vocal behavior are poorly understood, and it is unknown how vocal learning generalizes across an animal's entire repertoire of natural vocalizations and sequences. Here, we asked whether generalization occurs in a nonhuman vocal learner and quantified its properties. We hypothesized that adaptive error correction of a vocal gesture produced in one sequence would generalize to the same gesture produced in other sequences. To test our hypothesis, we manipulated the fundamental frequency (pitch) of auditory feedback in Bengalese finches (Lonchura striata var. domestica) to create sensory errors during vocal gestures (song syllables) produced in particular sequences. As hypothesized, error-corrective learning on pitch-shifted vocal gestures generalized to the same gestures produced in other sequential contexts. Surprisingly, generalization magnitude depended strongly on sequential distance from the pitch-shifted syllables, with greater adaptation for gestures produced near to the pitch-shifted syllable. A further unexpected result was that nonshifted syllables changed their pitch in the direction opposite from the shifted syllables. This apparently antiadaptive pattern of generalization could not be explained by correlations between generalization and the acoustic similarity to the pitch-shifted syllable. These findings therefore suggest that generalization depends on the type of vocal gesture and its sequential context relative to other gestures and may reflect an advantageous strategy for vocal learning and maintenance.

Left-hemisphere activation is associated with enhanced vocal pitch error detection in musicians with absolute pitch

The ability to process auditory feedback for vocal pitch control is crucial during speaking and singing. Previous studies have suggested that musicians with absolute pitch (AP) develop specialized left-hemisphere mechanisms for pitch processing. The present study adopted an auditory feedback pitch perturbation paradigm combined with ERP recordings to test the hypothesis whether the neural mechanisms of the left-hemisphere enhance vocal pitch error detection and control in AP musicians compared with relative pitch (RP) musicians and non-musicians (NM). Results showed a stronger N1 response to pitch-shifted voice feedback in the right-hemisphere for both AP and RP musicians compared with the NM group. However, the left-hemisphere P2 component activation was greater in AP and RP musicians compared with NMs and also for the AP compared with RP musicians. The NM group was slower in generating compensatory vocal reactions to feedback pitch perturbation compared with musicians, and they failed to re-adjust their vocal pitch after the feedback perturbation was removed. These findings suggest that in the earlier stages of cortical neural processing, the right hemisphere is more active in musicians for detecting pitch changes in voice feedback. In the later stages, the left-hemisphere is more active during the processing of auditory feedback for vocal motor control and seems to involve specialized mechanisms that facilitate pitch processing in the AP compared with RP musicians. These findings indicate that the left hemisphere mechanisms of AP ability are associated with improved auditory feedback pitch processing during vocal pitch control in tasks such as speaking or singing.

Modulation of effective connectivity during vocalization with perturbed auditory feedback

The integration of auditory feedback with vocal motor output is important for the control of voice fundamental frequency (F0). We used a pitch-shift paradigm where subjects respond to an alteration, or shift, of voice pitch auditory feedback with a reflexive change in F0. We presented varying magnitudes of pitch shifted auditory feedback to subjects during vocalization and passive listening and measured event related potentials (ERPs) to the feedback shifts. Shifts were delivered at +100 and +400 cents (200ms duration). The ERP data were modeled with dynamic causal modeling (DCM) techniques where the effective connectivity between the superior temporal gyrus (STG), inferior frontal gyrus and premotor areas were tested. We compared three main factors: the effect of intrinsic STG connectivity, STG modulation across hemispheres and the specific effect of hemisphere. A Bayesian model selection procedure was used to make inference about model families. Results suggest that both intrinsic STG and left to right STG connections are important in the identification of self-voice error and sensory motor integration. We identified differences in left-to-right STG connections between 100 cent and 400 cent shift conditions suggesting that self- and non-self-voice error are processed differently in the left and right hemisphere. These results also highlight the potential of DCM modeling of ERP responses to characterize specific network properties of forward models of voice control.

Effects of vocal training on voluntary responses to pitch-shifted voice auditory feedback

Control of voice fundamental frequency (F0) relies on the interaction between various forms of sensory feedback and neural motor control mechanisms. Several studies have shown that unexpected changes in pitch in the auditory feedback lead to reflexive compensatory vocal responses. We examined voluntary vocal responses to unpredictable perturbations in pitch auditory feedback. Ten subjects were trained over a five-day period to change their voice F0 in the opposite direction to the pitch-shifted feedback (±100 cents, 1000 ms) and 10 in the same direction as the feedback. Results showed that responses that followed the stimulus direction had significantly shorter latencies (200 ms) than opposing responses (324 ms). The reduced latency of the following responses suggests a switch from a feedback to a feedforward control strategy. The feedback strategy requires monitoring feedback and correcting for errors between the feedback signal and the intended vocal goal. The feedforward strategy relies less on auditory feedback and more on an internal model of the desired vocal pitch goal. Furthermore, feedback systems generally are slower than feedforward strategies, which would explain the shorter latencies of the responses that followed the stimulus direction. Results of this study will be discussed in terms of the differing strategies that may be used in various vocal contexts.

Cortical plasticity in the sensorimotor control of voice induced by auditory cognitive training

Multiple lines of evidence have shown that motor-related brain regions can be activated during passively musical listening or beat perception, indicating a connection between auditory and sensorimotor systems. In the present study, we sought to examine whether the neural processing of auditory-vocal integration can be shaped by short-term cognitive training related to auditory attention and working memory. Auditory cognitive training consisted of a ten-day backward digit span task, in which digits embedded in various noise at different SNR levels were presented and subjects were required to repeat the digits in the reverse order. Before and after the cognitive training, subjects also participated in a vocal motor task, in which they heard their pitch auditory feedback unexpectedly altered upwards (50 and 200 cents) during sustained vocalization and their neurophysiological responses were recorded. The results revealed a significantly improved performance on the backward digit span task after the training. Moreover, cortical responses indexed by P2 amplitude to pitch perturbations in voice auditory feedback were significantly increased after the training compared with those before the training. These findings provide evidence that plastic cortical changes in the sensorimotor control of voice can be caused by auditory cognitive training.

Vocal learning is constrained by the statistics of sensorimotor experience

The brain uses sensory feedback to correct behavioral errors. Larger errors by definition require greater corrections, and many models of learning assume that larger sensory feedback errors drive larger motor changes. However, an alternative perspective is that larger errors drive learning less effectively because such errors fall outside the range of errors normally experienced and are therefore unlikely to reflect accurate feedback. This is especially crucial in vocal control because auditory feedback can be contaminated by environmental noise or sensory processing errors. A successful control strategy must therefore rely on feedback to correct errors while disregarding aberrant auditory signals that would lead to maladaptive vocal corrections. We hypothesized that these constraints result in compensation that is greatest for smaller imposed errors and least for larger errors. To test this hypothesis, we manipulated the pitch of auditory feedback in singing Bengalese finches. We found that learning driven by larger sensory errors was both slower than that resulting from smaller errors and showed less complete compensation for the imposed error. Additionally, we found that a simple principle could account for these data: the amount of compensation was proportional to the overlap between the baseline distribution of pitch production and the distribution experienced during the shift. Correspondingly, the fraction of compensation approached zero when pitch was shifted outside of the song's baseline pitch distribution. Our data demonstrate that sensory errors drive learning best when they fall within the range of production variability, suggesting that learning is constrained by the statistics of sensorimotor experience.

A Lightweight, Headphones-based System for Manipulating Auditory Feedback in Songbirds

Experimental manipulations of sensory feedback during complex behavior have provided valuable insights into the computations underlying motor control and sensorimotor plasticity1. Consistent sensory perturbations result in compensatory changes in motor output, reflecting changes in feedforward motor control that reduce the experienced feedback error. By quantifying how different sensory feedback errors affect human behavior, prior studies have explored how visual signals are used to recalibrate arm movements2,3 and auditory feedback is used to modify speech production4-7. The strength of this approach rests on the ability to mimic naturalistic errors in behavior, allowing the experimenter to observe how experienced errors in production are used to recalibrate motor output.Songbirds provide an excellent animal model for investigating the neural basis of sensorimotor control and plasticity8,9. The songbird brain provides a well-defined circuit in which the areas necessary for song learning are spatially separated from those required for song production, and neural recording and lesion studies have made significant advances in understanding how different brain areas contribute to vocal behavior9-12. However, the lack of a naturalistic error-correction paradigm - in which a known acoustic parameter is perturbed by the experimenter and then corrected by the songbird - has made it difficult to understand the computations underlying vocal learning or how different elements of the neural circuit contribute to the correction of vocal errors13.The technique described here gives the experimenter precise control over auditory feedback errors in singing birds, allowing the introduction of arbitrary sensory errors that can be used to drive vocal learning. Online sound-processing equipment is used to introduce a known perturbation to the acoustics of song, and a miniaturized headphones apparatus is used to replace a songbird's natural auditory feedback with the perturbed signal in real time. We have used this paradigm to perturb the fundamental frequency (pitch) of auditory feedback in adult songbirds, providing the first demonstration that adult birds maintain vocal performance using error correction14. The present protocol can be used to implement a wide range of sensory feedback perturbations (including but not limited to pitch shifts) to investigate the computational and neurophysiological basis of vocal learning.

A Lightweight, Headphones-based System for Manipulating Auditory Feedback in Songbirds

Experimental manipulations of sensory feedback during complex behavior have provided valuable insights into the computations underlying motor control and sensorimotor plasticity1. Consistent sensory perturbations result in compensatory changes in motor output, reflecting changes in feedforward motor control that reduce the experienced feedback error. By quantifying how different sensory feedback errors affect human behavior, prior studies have explored how visual signals are used to recalibrate arm movements2,3 and auditory feedback is used to modify speech production4-7. The strength of this approach rests on the ability to mimic naturalistic errors in behavior, allowing the experimenter to observe how experienced errors in production are used to recalibrate motor output. Songbirds provide an excellent animal model for investigating the neural basis of sensorimotor control and plasticity8,9. The songbird brain provides a well-defined circuit in which the areas necessary for song learning are spatially separated from those required for song production, and neural recording and lesion studies have made significant advances in understanding how different brain areas contribute to vocal behavior9-12. However, the lack of a naturalistic error-correction paradigm - in which a known acoustic parameter is perturbed by the experimenter and then corrected by the songbird - has made it difficult to understand the computations underlying vocal learning or how different elements of the neural circuit contribute to the correction of vocal errors13. The technique described here gives the experimenter precise control over auditory feedback errors in singing birds, allowing the introduction of arbitrary sensory errors that can be used to drive vocal learning. Online sound-processing equipment is used to introduce a known perturbation to the acoustics of song, and a miniaturized headphones apparatus is used to replace a songbird's natural auditory feedback with the perturbed signal in real time. We have used this paradigm to perturb the fundamental frequency (pitch) of auditory feedback in adult songbirds, providing the first demonstration that adult birds maintain vocal performance using error correction14. The present protocol can be used to implement a wide range of sensory feedback perturbations (including but not limited to pitch shifts) to investigate the computational and neurophysiological basis of vocal learning.

Opposing and following vocal responses to pitch-shifted auditory feedback: Evidence for different mechanisms of voice pitch control

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.