Coordinating voicing onset with articulation: a potential role for sensory cues in shaping phonological distinctions.

In the typical speech of any language, voicing onset and offset are effortlessly coordinated with articulation as part of the intrinsic coordination of sound production. In this paper, we argue that voicing-articulatory coordination patterns could be shaped by sensory feedback during early speech learning and these patterns persist in mature syllable productions. Our experimental results show that voicing onset is closely associated with the peak velocity and peak amplitude of jaw and upper lip movements for VC syllables in adults. This robust coordination in the onset position may function to increase the salience of VC syllables and provide a phonetically natural explanation for vowels to undergo phonological lengthening and to avoid phonological reduction in word-initial onset position.

Common neural substrates support speech and non-speech vocal tract gestures.

The issue of whether speech is supported by the same neural substrates as non-speech vocal tract gestures has been contentious. In this fMRI study we tested whether producing non-speech vocal tract gestures in humans shares the same functional neuroanatomy as non-sense speech syllables. Production of non-speech vocal tract gestures, devoid of phonological content but similar to speech in that they had familiar acoustic and somatosensory targets, was compared to the production of speech syllables without meaning. Brain activation related to overt production was captured with BOLD fMRI using a sparse sampling design for both conditions. Speech and non-speech were compared using voxel-wise whole brain analyses, and ROI analyses focused on frontal and temporoparietal structures previously reported to support speech production. Results showed substantial activation overlap between speech and non-speech function in regions. Although non-speech gesture production showed greater extent and amplitude of activation in the regions examined, both speech and non-speech showed comparable left laterality in activation for both target perception and production. These findings posit a more general role of the previously proposed "auditory dorsal stream" in the left hemisphere--to support the production of vocal tract gestures that are not limited to speech processing.

Sensory stimulation activates both motor and sensory components of the swallowing system.

Volitional swallowing in humans involves the coordination of both brainstem and cerebral swallowing control regions. Peripheral sensory inputs are necessary for safe and efficient swallowing, and their importance to the patterned components of swallowing has been demonstrated. However, the role of sensory inputs to the cerebral system during volitional swallowing is less clear. We used four conditions applied during functional magnetic resonance imaging to differentiate between sensory, motor planning, and motor execution components for cerebral control of swallowing. Oral air pulse stimulation was used to examine the effect of sensory input, covert swallowing was used to engage motor planning for swallowing, and overt swallowing was used to activate the volitional swallowing system. Breath-holding was also included to determine whether its effects could account for the activation seen during overt swallowing. Oral air pulse stimulation, covert swallowing and overt swallowing all produced activation in the primary motor cortex, cingulate cortex, putamen and insula. Additional regions of the swallowing cerebral system that were activated by the oral air pulse stimulation condition included the primary and secondary somatosensory cortex and thalamus. Although air pulse stimulation was on the right side only, bilateral cerebral activation occurred. On the other hand, covert swallowing minimally activated sensory regions, but did activate the supplementary motor area and other motor regions. Breath-holding did not account for the activation during overt swallowing. The effectiveness of oral-sensory stimulation for engaging both sensory and motor components of the cerebral swallowing system demonstrates the importance of sensory input in cerebral swallowing control.

The effect of surface electrical stimulation on vocal fold position.

OBJECTIVES/HYPOTHESIS: Closure of the true and false vocal folds is a normal part of airway protection during swallowing. Individuals with reduced or delayed true vocal fold closure can be at risk for aspiration and may benefit from intervention to ameliorate the problem. Surface electrical stimulation is currently used during therapy for dysphagia, despite limited knowledge of its physiological effects. DESIGN: Prospective single effects study. METHODS: The immediate physiological effect of surface stimulation on true vocal fold angle was examined at rest in 27 healthy adults using 10 different electrode placements on the submental and neck regions. Fiberoptic nasolaryngoscopic recordings during passive inspiration were used to measure change in true vocal fold angle with stimulation. RESULTS: Vocal fold angles changed only to a small extent during two electrode placements (P < or = .05). When two sets of electrodes were placed vertically on the neck, the mean true vocal fold abduction was 2.4 degrees; while horizontal placements of electrodes in the submental region produced a mean adduction of 2.8 degrees (P = .03). CONCLUSIONS: Surface electrical stimulation to the submental and neck regions does not produce immediate true vocal fold adduction adequate for airway protection during swallowing, and one position may produce a slight increase in true vocal fold opening.

The effect of surface electrical stimulation on hyolaryngeal movement in normal individuals at rest and during swallowing.

Surface electrical stimulation is currently used in therapy for swallowing problems, although little is known about its physiological effects on neck muscles or swallowing. Previously, when one surface electrode placement was used in dysphagic patients at rest, it lowered the hyolaryngeal complex. Here we examined the effects of nine other placements in normal volunteers to determine 1) whether movements induced by surface stimulation using other placements differ, and 2) whether lowering the hyolaryngeal complex by surface electrical stimulation interfered with swallowing in healthy adults. Ten bipolar surface electrode placements overlying the submental and laryngeal regions were tested. Maximum tolerated stimulation levels were applied at rest while participants held their mouths closed. Videofluoroscopic recordings were used to measure hyoid bone and subglottic air column (laryngeal) movements from resting position and while swallowing 5 ml of liquid barium, with and without stimulation. Videofluoroscopic recordings of swallows were rated blind to condition using the National Institutes of Health-Swallowing Safety Scale. Significant (P < 0.0001) laryngeal and hyoid descent occurred with stimulation at rest. During swallowing, significant (P

Laryngeal muscle responses to mechanical displacement of the thyroid cartilage in humans.

Speakers may use laryngeal sensory feedback to adjust vocal fold tension and length before initiating voice. The mechanism for accurately initiating voice at an intended pitch is unknown, given the absence of laryngeal muscle spindles in animals and conflicting findings regarding their existence in humans. Previous reports of rapid changes in voice fundamental frequency following thyroid cartilage displacement suggest that changes in vocal fold length modulate laryngeal muscle contraction in humans. We tested the hypothesis that voice changes resulting from mechanical perturbation are due to rapid responses in the intrinsic laryngeal muscles. Hooked wire electrodes were used to record from the thyroarytenoid, cricothyroid, and sternothyroid muscles along with surface electrodes on the skin overlying the thyroid cartilage in 10 normal adults. Servomotor displacements produced consistent changes in the subjects' vocal fundamental frequency at 70-80 ms, demonstrating changes in vocal fold length and tension. No simultaneous electromyographic responses occurred in the thyroarytenoid or cricothyroid muscles in any subjects. Instead, short-latency responses at 25-40 ms following stimulus onset occurred in the sternothyroid muscles, simultaneous with responses in the surface recordings. The sternothyroid responses may modulate long-latency changes in voice fundamental frequency (approximately 150 ms). The absence of intrinsic laryngeal muscle responses is consistent with a lack of spindles in these muscles. Our results suggest that other sensory receptors, such as mucosal mechanoreceptors, provide feedback for voice control.

Suppression of thyroarytenoid muscle responses during repeated air pressure stimulation of the laryngeal mucosa in awake humans.

Repeated stimulation of the laryngeal mucosa occurs during speech. Single stimuli, however, can elicit the laryngeal adductor response (LAR). Our hypothesis was that the LAR to repeated rapid air pressure stimuli is centrally suppressed in humans. Hooked-wire electrodes were inserted into the thyroarytenoid and cricothyroid muscles on both sides and into the posterior cricoarytenoid muscle on one side. Pairs of air puff stimuli were presented to the mucosa over the arytenoids at pressure levels three times threshold with interstimulus intervals from 250 to 5,000 ms. Bilateral thyroarytenoid responses occurred at around 150 ms to more than 70% of the initial stimuli. With repeated presentation at intervals of 2 seconds or less, the percent occurrence decreased to less than 40% and response amplitudes were reduced by 50%. Central suppression of adductor responses to repeated air puff stimuli may allow speakers to produce voice without eliciting reflexive spasms that could disrupt speech.

Correspondence between laryngeal vocal fold movement and muscle activity during speech and nonspeech gestures.

To better understand the role of each of the laryngeal muscles in producing vocal fold movement, activation of these muscles was correlated with laryngeal movement during different tasks such as sniff, cough or throat clear, and speech syllable production. Four muscles [the posterior cricoarytenoid, lateral cricoarytenoid, cricothyroid (CT), and thyroarytenoid (TA)] were recorded with bipolar hooked wire electrodes placed bilaterally in four normal subjects. A nasoendoscope was used to record vocal fold movement while simultaneously recording muscle activity. Muscle activation level was correlated with ipsilateral vocal fold angle for vocal fold opening and closing. Pearson correlation coefficients and their statistical significance were computed for each trial. Significant effects of muscle (P < or = 0.0005) and task (P = 0.034) were found on the r (transformed to Fisher's Z') values. All of the posterior cricoarytenoid recordings related significantly with vocal opening, whereas CT activity was significantly correlated with opening only during sniff. The TA and lateral cricoarytenoid activities were significantly correlated with vocal fold closing during cough. During speech, the CT and TA activity correlated with both opening and closing. Laryngeal muscle patterning to produce vocal fold movement differed across tasks; reciprocal muscle activity only occurred on cough, whereas speech and sniff often involved simultaneous contraction of muscle antagonists. In conclusion, different combinations of muscle activation are used for biomechanical control of vocal fold opening and closing movements during respiratory, airway protection, and speech tasks.

Mucosal afferents mediate laryngeal adductor responses in the cat.

Laryngeal adductor responses (LAR) close the airway in response to stimulation of peripheral afferents in the superior laryngeal nerve. Although both mucosal afferents and proprioceptive receptors are present in the larynx, their relative contribution for reflex elicitation is unknown. Our purpose was to determine which receptor types are of importance in eliciting the LAR. A servomotor with displacement feedback was used to deliver punctate displacements to the body of the arytenoid cartilage and overlying mucosa on each side of the larynx in eight anesthetized cats. The same displacements were delivered both before and after surgical excision of the overlying mucosa. With the mucosa intact, early short-latency component R1 LAR responses recorded from the thyroarytenoid muscles were frequent (ipsilateral > 92%, contralateral > 95%). After the mucosa was removed, the LAR became infrequent (<3%) and was reduced in amplitude in both the ipsilateral and contralateral thyroarytenoid muscle recording sites (P < 0.0005). These findings demonstrate that mucosal mechanoreceptors and not proprioceptive afferents contribute to the elicitation of LAR responses in the cat.

Elevating pain thresholds in humans using depolarizing prepulses.

Electrocutaneous stimulation is a potentially useful communication tool for applications in virtual reality, sensory substitution, and sensory augmentation. Many of these applications require the use of arrays of small electrodes. Stimulation through small electrodes is often painful, however, limiting the practicality of such arrays. The purpose of this study was to test a method for elevating the pain threshold to electrocutaneous stimulation through small (1-mm diameter) electrodes on the fingertip. We hypothesized that long, subthreshold, depolarizing prepulses (PP) would elevate the pain threshold so that a subsequent stimulus pulse (SP) would be less likely to be painful. We used psychophysical methods to measure the probability that an SP would be perceived as painful both by itself and when preceded by a PP that was 2, 4, 6, 8, or 10 dB lower in amplitude than the SP. We found that the PPs significantly increased the pain threshold, reducing the likelihood that the SP was painful (p < .0001). The dose effect of PP amplitude was also highly significant (p < .0001), with larger PPs elevating pain thresholds more. To our knowledge, this is the first report of PPs being used to elevate electrical stimulation thresholds in humans. PPs may be useful for selective inactivation of neural subpopulations in many human neuroprosthetic applications.

Functional neuroanatomy of human voluntary cough and sniff production.

Cough and sniff are both spontaneous respiratory behaviors that can be initiated voluntarily in humans. Disturbances of cough may be life threatening, while inability to sniff impairs the sense of smell in neurological patients. Cortical mechanisms of voluntary cough and sniff production have been predicted to exist; however, the localization and function of supramedullary areas responsible for these behaviors are poorly understood. We used functional magnetic resonance imaging to identify the central control of voluntary cough and sniff compared with breathing. We determined that both voluntary cough and sniff require a widespread pattern of sensorimotor activation along the Sylvian fissure convergent with voluntary breathing. Task-specific activation occurred in a pontomesencephalic region during voluntary coughing and in the hippocampus and piriform cortex during voluntary sniffing. Identification of the localization of cortical activation for cough control in humans may help potential drug development to target these regions in patients with chronic cough. Understanding the sensorimotor sniff control mechanisms may provide a new view on the cerebral functional reorganization of olfactory control in patients with neurological disorders.

Human brain activation during phonation and exhalation: common volitional control for two upper airway functions.

Phonation is defined as a laryngeal motor behavior used for speech production, which involves a highly specialized coordination of laryngeal and respiratory neuromuscular control. During speech, brief periods of vocal fold vibration for vowels are interspersed by voiced and unvoiced consonants, glottal stops and glottal fricatives (/h/). It remains unknown whether laryngeal/respiratory coordination of phonation for speech relies on separate neural systems from respiratory control or whether a common system controls both behaviors. To identify the central control system for human phonation, we used event-related fMRI to contrast brain activity during phonation with activity during prolonged exhalation in healthy adults. Both whole-brain analyses and region of interest comparisons were conducted. Production of syllables containing glottal stops and vowels was accompanied by activity in left sensorimotor, bilateral temporoparietal and medial motor areas. Prolonged exhalation similarly involved activity in left sensorimotor and temporoparietal areas but not medial motor areas. Significant differences between phonation and exhalation were found primarily in the bilateral auditory cortices with whole-brain analysis. The ROI analysis similarly indicated task differences in the auditory cortex with differences also detected in the inferolateral motor cortex and dentate nucleus of the cerebellum. A second experiment confirmed that activity in the auditory cortex only occurred during phonation for speech and did not depend upon sound production. Overall, a similar central neural system was identified for both speech phonation and voluntary exhalation that primarily differed in auditory monitoring.