Compensatory mechanisms affect sensorimotor integration during ongoing vocal motor acts in marmoset monkeys

Any transmission of vocal signals faces the challenge of acoustic interferences such as heavy rain, wind, animal or urban sounds. Consequently, several mechanisms and strategies have evolved to optimize signal‐to‐noise ratio. Examples to increase detectability are the Lombard effect, an involuntary rise in call amplitude in response to masking ambient noise, which is often associated with other vocal changes such as call frequency and duration, as well as the animals’ capability of limiting calling to periods where noise perturbation is absent. Previous studies revealed vocal flexibility and various audio‐vocal integration mechanisms in marmoset monkeys. Using acoustic perturbation triggered by vocal behaviour, we investigated whether marmosets are capable of exhibiting changes in call structure when perturbing noise starts after call onset or whether such effects only occur if noise perturbation starts prior to call onset. We show that marmosets are capable of rapidly modulating call amplitude and frequency in response to such noise perturbation. Vocalizations swiftly increased call frequency after noise onset indicating a rapid effect of perturbing noise on vocal motor production. Call amplitudes were also affected. Interestingly, however, the marmosets did not exhibit the Lombard effect as previously reported but decreased call intensity in response to noise. Our findings indicate that marmosets possess a general avoidance strategy to call in the presence of ambient noise and suggest that these animals are capable of counteracting a previously thought involuntary audio‐vocal mechanism, the Lombard effect. These findings will pave the way to investigate the underlying audio‐vocal integration mechanisms explaining these behaviours.     

Effects of voice harmonic complexity on ERP responses to pitch-shifted auditory feedback

Objective

The present study investigated the neural mechanisms of voice pitch control for different levels of harmonic complexity in the auditory feedback.

Methods

Event-related potentials (ERPs) were recorded in response to +200 cents pitch perturbations in the auditory feedback of self-produced natural human vocalizations, complex and pure tone stimuli during active vocalization and passive listening conditions.

Results

During active vocal production, ERP amplitudes were largest in response to pitch shifts in the natural voice, moderately large for non-voice complex stimuli and smallest for the pure tones. However, during passive listening, neural responses were equally large for pitch shifts in voice and non-voice complex stimuli but still larger than that for pure tones.

Conclusions

These findings suggest that pitch change detection is facilitated for spectrally rich sounds such as natural human voice and non-voice complex stimuli compared with pure tones. Vocalization-induced increase in neural responses for voice feedback suggests that sensory processing of naturally-produced complex sounds such as human voice is enhanced by means of motor-driven mechanisms (e.g. efference copies) during vocal production.

Significance

This enhancement may enable the audio-vocal system to more effectively detect and correct for vocal errors in the feedback of natural human vocalizations to maintain an intended vocal output for speaking.     

Audio-vocal interaction in the pontine brainstem during self-initiated vocalization in the squirrel monkey

The adjustment of the voice by auditory input happens at several brain levels. The caudal pontine brainstem, though rarely investigated, is one candidate area for such audio-vocal integration. We recorded neuronal activity in this area in awake, behaving squirrel monkeys (Saimiri sciureus) during vocal communication, using telemetric single-unit recording techniques. We found audio-vocal neurons at locations not described before, namely in the periolivary region of the superior olivary complex and the adjacent pontine reticular formation. They showed various responses to external sounds (noise bursts) and activity increases (excitation) or decreases (inhibition) to self-produced vocalizations, starting prior to vocal onset and continuing through vocalizations. In most of them, the responses to noise bursts and self-produced vocalizations were similar, with the only difference that neuronal activity started prior to vocal onset. About one-third responded phasically to noise bursts, independent of whether they increased or decreased their activity to vocalization. The activity of most audio-vocal neurons correlated with basic acoustic features of the vocalization, such as call duration and/or syllable structure. Auditory neurons near audio-vocal neurons showed significantly more frequent phasic response patterns than those in areas without audio-vocal activity. Based on these findings, we propose that audio-vocal neurons showing similar activity to external acoustical stimuli and vocalization play a role in olivocochlear regulation. Specifically, audio-vocal neurons with a phasic response to external auditory stimuli are candidates for the mediation of basal audio-vocal reflexes such as the Lombard reflex. Thus, our findings suggest that complex audio-vocal integration mechanisms exist in the ventrolateral pontine brainstem.     

Vocalization-induced enhancement of the auditory cortex responsiveness during voice F0 feedback perturbation

ObjectiveThe present study investigated whether self-vocalization enhances auditory neural responsiveness to voice pitch feedback perturbation and how this vocalization-induced neural modulation can be affected by the extent of the feedback deviation.MethodsEvent-related potentials (ERPs) were recorded in 15 subjects in response to +100, +200 and +500 cents pitch-shifted voice auditory feedback during active vocalization and passive listening to the playback of the self-produced vocalizations.ResultsThe amplitude of the evoked P₁ (latency: 73.51 ms) and P₂ (latency: 199.55 ms) ERP components in response to feedback perturbation were significantly larger during vocalization than listening. The difference between P₂ peak amplitudes during vocalization vs. listening was shown to be significantly larger for +100 than +500 cents stimulus.ConclusionsResults indicate that the human auditory cortex is more responsive to voice F₀ feedback perturbations during vocalization than passive listening. Greater vocalization-induced enhancement of the auditory responsiveness to smaller feedback perturbations may imply that the audio–vocal system detects and corrects for errors in vocal production that closely match the expected vocal output.SignificanceFindings of this study support previous suggestions regarding the enhanced auditory sensitivity to feedback alterations during self-vocalization, which may serve the purpose of feedback-based monitoring of one’s voice.     

Bihemispheric network dynamics coordinating vocal feedback control

Modulation of vocal pitch is a key speech feature that conveys important linguistic and affective information. Auditory feedback is used to monitor and maintain pitch. We examined induced neural high gamma power (HGP) (65–150 Hz) using magnetoencephalography during pitch feedback control. Participants phonated into a microphone while hearing their auditory feedback through headphones. During each phonation, a single real‐time 400 ms pitch shift was applied to the auditory feedback. Participants compensated by rapidly changing their pitch to oppose the pitch shifts. This behavioral change required coordination of the neural speech motor control network, including integration of auditory and somatosensory feedback to initiate change in motor plans. We found increases in HGP across both hemispheres within 200 ms of pitch shifts, covering left sensory and right premotor, parietal, temporal, and frontal regions, involved in sensory detection and processing of the pitch shift. Later responses to pitch shifts (200–300 ms) were right dominant, in parietal, frontal, and temporal regions. Timing of activity in these regions indicates their role in coordinating motor change and detecting and processing of the sensory consequences of this change. Subtracting out cortical responses during passive listening to recordings of the phonations isolated HGP increases specific to speech production, highlighting right parietal and premotor cortex, and left posterior temporal cortex involvement in the motor response. Correlation of HGP with behavioral compensation demonstrated right frontal region involvement in modulating participant's compensatory response. This study highlights the bihemispheric sensorimotor cortical network involvement in auditory feedback‐based control of vocal pitch. Hum Brain Mapp 37:1474‐1485, 2016. © 2016 Wiley Periodicals, Inc.     

Correlation between neural discharges in cat primary auditory cortex and tone-detection behaviors

Understanding the physiological role of the auditory cortex (AC) in acoustic perception is an essential issue in auditory neuroscience. By comparing sound discrimination behaviors in animals before and after AC lesion, many studies have demonstrated that AC is necessary for the perceptual process of human vowels and animal vocalizations, but is not necessary to discriminate simple acoustic parameters such as sound onset, intensity and duration. Because a lesion study cannot fully reveal the function of AC under normal conditions, in this study, we combined electrophysiological recording and psychophysical experiments on the same animal to investigate whether AC is involved in a simple auditory task. We recorded the neural activities of the primary auditory cortex (A1) using implanted electrodes, while freely-moving cats performed a tone-detection task in which they were required to lick a metal tube to obtain a food reward after hearing a tone pip. The performance of the cats' behavioral response increased with the increase of tone intensity, and the neural activities of A1 covaried with the behavioral performance. Also, whether the tone-detection behavior was interfered by a wideband noise was dependent on whether the tone-evoked neural response was masked by the noise-evoked response. Our results did not support that A1 neurons directly associate with the cat's behavioral decision; instead, they may mainly generate a neural representation of stimulus amplitude for further processing to determine whether a tone occurred or not.     

Cerebellar contribution to auditory feedback control of speech production: Evidence from patients with spinocerebellar ataxia

The cerebellum has been implicated in the feedforward control of speech production. However, the role of the cerebellum in the feedback control of speech production remains unclear. To address this question, the present event‐related potential study examined the behavioral and neural correlates of auditory feedback control of vocal production in patients with spinocerebellar ataxia (SCA) and healthy controls. All participants were instructed to produce sustained vowels while hearing their voice unexpectedly pitch‐shifted ?200 or ?500 cents. The behavioral results revealed significantly larger vocal compensations for pitch perturbations in patients with SCA relative to healthy controls. At the cortical level, patients with SCA exhibited significantly smaller cortical P2 responses that were source localized in the right superior temporal gyrus, primary auditory cortex, and supramarginal gyrus than healthy controls. These findings indicate that reduced brain activity in the right temporal and parietal regions are significant neural contributors to abnormal auditory‐motor processing of vocal pitch regulation as a consequence of cerebellar degeneration, which may be related to disrupted reciprocal interactions between the cerebellum and cortical regions that support the top‐down modulation of auditory‐vocal integration. These differences in behavior and cortical activity between healthy controls and patients with SCA demonstrate that the cerebellum is not only essential for feedforward control but also plays a crucial role in the feedback‐based control of speech production.     

Intermittent theta burst stimulation over right somatosensory larynx cortex enhances vocal pitch‐regulation in nonsingers

While the significance of auditory cortical regions for the development and maintenance of speech motor coordination is well established, the contribution of somatosensory brain areas to learned vocalizations such as singing is less well understood. To address these mechanisms, we applied intermittent theta burst stimulation (iTBS), a facilitatory repetitive transcranial magnetic stimulation (rTMS) protocol, over right somatosensory larynx cortex (S1) and a nonvocal dorsal S1 control area in participants without singing experience. A pitch‐matching singing task was performed before and after iTBS to assess corresponding effects on vocal pitch regulation. When participants could monitor auditory feedback from their own voice during singing (Experiment I), no difference in pitch‐matching performance was found between iTBS sessions. However, when auditory feedback was masked with noise (Experiment II), only larynx‐S1 iTBS enhanced pitch accuracy (50–250 ms after sound onset) and pitch stability (>250 ms after sound onset until the end). Results indicate that somatosensory feedback plays a dominant role in vocal pitch regulation when acoustic feedback is masked. The acoustic changes moreover suggest that right larynx‐S1 stimulation affected the preparation and involuntary regulation of vocal pitch accuracy, and that kinesthetic‐proprioceptive processes play a role in the voluntary control of pitch stability in nonsingers. Together, these data provide evidence for a causal involvement of right larynx‐S1 in vocal pitch regulation during singing.     

Neurophysiological evidence of differential mechanisms involved in producing opposing and following responses to altered auditory feedback

Objective

When hearing perturbations in voice auditory feedback, people produce responses that mostly oppose the perturbation direction, whereas a few responses follow the direction of feedback perturbation. The causes of opposing and following responses, however, remain poorly understood. The present event-related potential (ERP) study sought to examine the neurophysiological processing of opposing and following responses to pitch feedback perturbations during self-monitoring of vocal production.

Method

Twelve Mandarin-native speakers participated in the experiment. Vocal and neurophysiological responses to pitch perturbations (±50 and ±200 cents) in voice auditory feedback were measured. Individual-trial responses were categorized according to the response direction and then separately averaged in groups of opposing and following responses. ERPs indexed by the P1-N1-P2 complex corresponding to two types of vocal responses were also obtained.

Results

Opposing and following vocal responses did not differ in the magnitude, but there were greater proportions of opposing to following responses to 50 cents stimuli. The amplitude and latency of the P1 and N1 components showed none of significance across conditions, whereas there was a direction × magnitude effect on the P2 response. Following responses elicited greater P2 amplitudes than opposing responses only when pitch feedback was perturbed for downward 200 cents, and upward pitch perturbation elicited greater P2 amplitudes than those with downward direction only in the production of opposing responses.

Conclusion

These findings demonstrate that cortical processing of opposing responses is different from that of following responses, which can be modulated by the physical properties of feedback perturbation.

Significance

Different neural mechanisms are involved in the production of opposing and following responses to feedback perturbations during self-monitoring of vocal production.     

Neural correlates of verbal feedback processing: an fMRI study employing overt speech

Speakers use external auditory feedback to monitor their own speech. Feedback distortion has been found to increase activity in the superior temporal areas. Using fMRI, the present study investigates the neural correlates of processing verbal feedback without distortion. In a blocked design, the following conditions were presented: (1) overt picture-naming, (2) overt picture-naming while pink noise was presented to mask external feedback, (3) covert picture-naming, (4) listening to the picture names (previously recorded from participants' own voices), and (5) listening to pink noise. The results show that auditory feedback processing involves a network of different areas related to general performance monitoring and speech-motor control. These include the cingulate cortex and the bilateral insula, supplementary motor area, bilateral motor areas, cerebellum, thalamus and basal ganglia. Our findings suggest that the anterior cingulate cortex, which is often implicated in error-processing and conflict-monitoring, is also engaged in ongoing speech monitoring. Furthermore, in the superior temporal gyrus, we found a reduced response to speaking under normal feedback conditions. This finding is interpreted in the framework of a forward model according to which, during speech production, the sensory consequence of the speech-motor act is predicted to attenuate the sensitivity of the auditory cortex.     

Human temporal-lobe response to vocal sounds

Voice is not only the vehicle of speech, it is also an 'auditory face' that conveys a wealth of information on a person's identity and affective state. In contrast to speech perception, little is known about the neural bases of our ability to perceive these various types of paralinguistic vocal information. Using functional magnetic resonance imaging (fMRI), we identified regions along the superior temporal sulcus (STS) that were not only sensitive, but also highly selective to vocal sounds. In the present study, we asked how neural activity in the voice areas was influenced by (i) the presence or not of linguistic information in the vocal input (speech vs. nonspeech) and (ii) frequency scrambling. Speech sounds were found to elicit greater responses than nonspeech vocalizations in most parts of auditory cortex, including primary auditory cortex (A1), on both sides of the brain. In contrast, response attenuation due to frequency scrambling was much more pronounced in anterior STS areas than at the level of A1. Importantly, only right anterior STS regions responded more strongly to nonspeech vocal sounds than to their scrambled version, suggesting that these regions could be specifically involved in paralinguistic aspects of voice perception.     

Morphology of axonal projections from the high vocal center to vocal motor cortex in songbirds

Only birds that learn complex vocalizations have telencephalic brain regions that control vocal learning and production, including HVC (high vocal center), a cortical nucleus that encodes vocal motor output in adult songbirds. HVC projects to RA (robust nucleus of the arcopallium), a nucleus in motor cortex that in turn projects topographically onto hindbrain neurons innervating vocal muscles. Individual neurons projecting from HVC to RA (HVC(RA) ) fire sparsely to drive RA activity during song production. To advance understanding of how individual HVC neurons encode production of learned vocalizations, we reconstructed single HVC axons innervating RA in adult male zebra finches. Individual HVC(RA) axons were not topographically organized within RA: 1) axon arbors of HVC cell bodies located near each other sent branches to different subregions of RA, and 2) branches of single HVC axons terminated in different locations within RA. HVC(RA) axons also had a simple, sparse morphology, suggesting that a single HVC neuron activates a limited population of postsynaptic RA neurons. These morphological data are consistent with previous work showing that single HVC(RA) neurons burst sparsely for a brief period of time during the production of a song, indicating that ensembles of HVC(RA) neurons fire simultaneously to drive small temporal segments of song behavior. We also examined the morphology of axons projecting from HVC to RA cup, a region surrounding RA that receives input from auditory cortex. Axons projecting to RA cup also sent some branches into RA, suggesting direct integration between the sensory and motor circuits for song control.     

A paradox in the evolution of primate vocal learning

The importance of auditory feedback in the development of spoken language in humans is striking. Paradoxically, although auditory-feedback-dependent vocal plasticity has been shown in a variety of taxonomic groups, there is little evidence that our nearest relatives--non-human primates--require auditory feedback for the development of species-typical vocal signals. Because of the apparent lack of developmental plasticity in the vocal production system, neuroscientists have largely ignored the neural mechanisms of non-human primate vocal production and perception. Recently, the absence of evidence for vocal plasticity from developmental studies has been contrasted with evidence for vocal plasticity in adults. We argue that this new evidence makes non-human primate vocal behavior an attractive model system for neurobiological analysis.     

Vocalization-related stapedius muscle activity in different age chickens (Gallus gallus), and its role in vocal development

The stapedius muscle activity associated with vocalization was analyzed in young and adult roosters. Our results show that remarkable differences in the behavior of vocalization-related stapedius muscle activity exist between these two ages. Unlike young roosters, electrical stimulation in the midbrain of adult cocks yields vocalizations associated with stapedius muscle EMG responses that always show a higher threshold and a longer latency than those of the vocalization induced. Moreover, the maximal amplitude of the stapedius muscle EMG response is consistently lower than that detected in young roosters, despite the fact that the maximal vocalization amplitude of the adult birds is much higher. On the whole our results demonstrate that vocalization-related stapedius muscle activity is strongly reduced in adulthood. The possibility that stapedius muscle may play a role during the vocal development was verified by comparing the crow of normal roosters with that of cocks from which the stapedius muscle had been removed shortly after hatching. Strong differences exist in the amplitude/frequency distribution of the crowing of normal and stapedectomized roosters, suggesting that the stapedius muscle exerts an important role in auditory feedback modulation, and that this feedback is used for normal vocal development.     

Role of neocortex in binsural hearing in the cat. I. Contralateral masking.

Cats with earphones were trained with a shock avoidance procedure to detect the occurrence of 1 kHz tone pulses at one ear while continuous noise pulses were simultaneously presented to the opposite ear. For normal cats the presence of the noise produced a mean increase of 5.4 dB in the thresholds for detection of tones at the opposite ears. After large unilateral auditory cortex ablations the same cats exhibited an asymmetry between the ears in the size of the contralateral masking effect. There was a mean increase of 10.9 dB in the detection thresholds for tones at the ear contralateral to the damaged hemisphere when noise was presented to the ear opposite the intact hemisphere. Noise of the same physical intensity when presented to the ear contralateral to the damaged cortex produced no significant changes from the preoperative masking levels. Subsequent ablation of the auditory cortex of the opposite hemisphere resulted in a cancellation of the unilateral lesion effect; the cats exhibited interaurally symmetrical masking levels of the same magnitude as those observed prior to the first operation. Additional control tests indicate that the unilateral lesion effect is a central nervous system phenomenon and is specific to lesions of auditory cortex.     

The impact of parkinson's disease on the cortical mechanisms that support auditory–motor integration for voice control

Several studies have shown sensorimotor deficits in speech processing in individuals with idiopathic Parkinson's disease (PD). The underlying neural mechanisms, however, remain poorly understood. In the present event‐related potential (ERP) study, 18 individuals with PD and 18 healthy controls were exposed to frequency‐altered feedback (FAF) while producing a sustained vowel and listening to the playback of their own voice. Behavioral results revealed that individuals with PD produced significantly larger vocal compensation for pitch feedback errors than healthy controls, and exhibited a significant positive correlation between the magnitude of their vocal responses and the variability of their unaltered vocal pitch. At the cortical level, larger P2 responses were observed for individuals with PD compared with healthy controls during active vocalization due to left‐lateralized enhanced activity in the superior and inferior frontal gyrus, premotor cortex, inferior parietal lobule, and superior temporal gyrus. These two groups did not differ, however, when they passively listened to the playback of their own voice. Individuals with PD also exhibited larger P2 responses during active vocalization when compared with passive listening due to enhanced activity in the inferior frontal gyrus, precental gyrus, postcentral gyrus, and middle temporal gyrus. This enhancement effect, however, was not observed for healthy controls. These findings provide neural evidence for the abnormal auditory–vocal integration for voice control in individuals with PD, which may be caused by their deficits in the detection and correction of errors in voice auditory feedback. Hum Brain Mapp 37:4248–4261, 2016. © 2016 Wiley Periodicals, Inc.     

Auditory response following vocalization: a magnetoencephalographic study.

OBJECTIVE: We recorded vocalization-related cortical fields (VRCF) under complete masking of a subject's own voice to identify the auditory component evoked by a subject's own voice in the VRCF complex. METHODS: We recorded VRCF during simple vowel (/u/) vocalization in 10 right-handed healthy volunteers under two conditions: (1) no masking (control) and (2) masking of the subject's own voice by weighted-white noise during vocalization. In the second experiment, we recorded auditory evoked magnetic fields (AEF) following stimulation of a speech sound applied by voice-recorder. RESULTS: The onset of VRCF appeared gradually before the vocalization onset, and a clear phase-reversed deflection was identified after the onset of vocalization. The difference waveform obtained by subtracting the VRCF of the masking condition from that of the control showed a deflection (1M) at 81.3+/-20.5 (mean+/-SD) ms after the onset of vocalization, but there was no consistent deflection before the vocalization onset. The AEF following voice sound in the second experiment showed the M100 component at 94.3+/-18.4 ms. The equivalent current dipole of the 1M component for different waveforms was located close in the auditory cortex to that of the M100 for AEF waveforms in each hemisphere. CONCLUSION: We successfully separated the auditory feedback response from the VRCF complex, using an adequate masking condition during vocalization of a subject's own voice. The masking effect was crucial to the auditory feedback process after the onset of vocalization. The present results suggested that the 1M component was mainly generated from the auditory feedback process by the subject's own voice. The activated auditory area for simple own voice might be similar to that for simple external sound.     

Intrinsic and extrinsic contributions to auditory selectivity in a song nucleus critical for vocal plasticity.

The development, maintenance, and perception of learned vocalizations in songbirds are likely to require auditory neurons that respond selectively to song. Neurons with song-selective responses have been described in several brain nuclei critical to singing, but the mechanisms by which such response properties arise, are modified, and propagate are poorly understood. The lateral magnocellular nucleus of the anterior neostriatum (LMAN) is the output of an anterior forebrain pathway (AFP) essential for learning and maintenance of song, processes dependent on auditory feedback. Although neurons throughout this pathway respond selectively to auditory presentation of the bird's own song, LMAN is the last stage at which responses to this auditory information could be transformed before being transmitted to vocal motor areas, where such responses may influence vocal production. Indeed, previous extracellular studies have indicated that LMAN's auditory selectivity is greater than that at earlier stages of the AFP. To determine whether LMAN local circuitry transforms or simply relays song-related auditory information to vocal control neurons, it is essential to distinguish local from extrinsic contributions to LMAN's auditory selectivity. In vivo intracellular recordings from LMAN projection neurons, coupled with local circuit inactivation, reveal that much of LMAN's song selectivity is supplied by its extrinsic inputs, but selective blockade of GABA receptors indicates that local inhibition is required for the expression of song selectivity. Therefore, LMAN neurons receive highly song-selective information, but LMAN's local circuitry can mask these selective inputs, providing a mechanism for context-dependent auditory feedback.     

Wireless multi-channel single unit recording in freely moving and vocalizing primates

The ability to record well-isolated action potentials from individual neurons in naturally behaving animals is crucial for understanding neural mechanisms underlying natural behaviors. Traditional neurophysiology techniques, however, require the animal to be restrained which often restricts natural behavior. An example is the common marmoset (Callithrix jacchus), a highly vocal New World primate species, used in our laboratory to study the neural correlates of vocal production and sensory feedback. When restrained by traditional neurophysiological techniques marmoset vocal behavior is severely inhibited. Tethered recording systems, while proven effective in rodents pose limitations in arboreal animals such as the marmoset that typically roam in a three-dimensional environment. To overcome these obstacles, we have developed a wireless neural recording technique that is capable of collecting single-unit data from chronically implanted multi-electrodes in freely moving marmosets. A lightweight, low power and low noise wireless transmitter (headstage) is attached to a multi-electrode array placed in the premotor cortex of the marmoset. The wireless headstage is capable of transmitting 15 channels of neural data with signal-to-noise ratio (SNR) comparable to a tethered system. To minimize radio-frequency (RF) and electro-magnetic interference (EMI), the experiments were conducted within a custom designed RF/EMI and acoustically shielded chamber. The individual electrodes of the multi-electrode array were periodically advanced to densely sample the cortical layers. We recorded single-unit data over a period of several months from the frontal cortex of two marmosets. These recordings demonstrate the feasibility of using our wireless recording method to study single neuron activity in freely roaming primates.     

Repetitive vocalizations evoked by electrical stimulation of avian brains. IV. Evoked and spontaneous activity in expiratory and inspiratory nerves and muscles of the chicken (Gallus gallus).

The activity in respiratory nerves and muscles in response to electrical stimulation of vocal substrates in the brain and to CO2 stimulation of the respiratory centers was studied in 28 adult chickens. It was found that the same nerves and muscles were active during both vocalization and respiration. Stimulation of vocal substrates resulted in short latency bursting in the expiratory nerves and muscles. As stimulation intensity increased, progressively longer duration bursts composed of numerous subbursts were produced. By relating muscle activity with sound production , such bursting was shown to underlie evoked vocalizations. Background activity in inspiratory nerves and muscles continued uninterruptedly past stimulus onset only stopping when expiratory activity began. Thereafter inspiratory bursting reciprocated with expiratory bursting and was shown to underlie the intervals between vocalizations. The pattern of activity which was evoked by stimulating vocal substrates was found to strongly interact with the pattern of activity evoked by CO2 stimulation of the respiratory system. Simultaneous records of respiratory and tracheal muscles demonstrated that the same information was sent to both groups of muscles during evoked vocalization. Activity in the respiratory muscles was recorded during spontaneous vocalization of a free-moving bird and was found to resemble that recorded from anesthetized birds. Finally the activity of single units in the obex region of the medulla was recorded during electrical stimulation of vocal substrates and during CO2 stimulation of the respiratory system. Rhythmically active units were found only in the medulla. Unit activity paralleled that found in the nerves and muscles. On the basis on the data accumulated, two models of the chicken vocal system are presented. The first is a model of the sound-producing structures of the chicken. The second is a model of the neural machinery which controls the sound-producing structures. The two models are used as a basis for an explanation of the production of voclizations by the chick of the same species.