Human brain activation during phonation and exhalation: Common volitional Control for two upper airway functions

Laryngeal and Speech Section, Medical Neurology Branch, National Institute of Neurological Disorders and Stroke, National Institutes of Health, Bethesda, MD, USA.
NeuroImage (Impact Factor: 6.36). 06/2007; 36(1):131-43. DOI: 10.1016/j.neuroimage.2007.01.049
Source: PubMed


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.

    • "This represents a striking species difference in the cortical control of the larynx between monkeys and humans, one supported by neuroanatomy. In monkeys, the cortical larynx area is restricted to the premotor cortex (Hast et al., 1974), while in the human brain it extends into primary motor cortex as well (Loucks et al., 2007; Brown et al., 2008; Simonyan et al., 2009; Belyk and Brown, 2014b). The human larynx area is activated by volitional movement of the laryngeal muscles, phonation and forced expiration, leading us to refer to it as the 'larynx phonation area' (LPA; Brown et al., 2008). "
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    ABSTRACT: Affective prosody is that aspect of speech that conveys a speaker's emotional state through modulations in various vocal parameters, most prominently pitch. While a large body of research implicates the cingulate vocalization area in controlling affective vocalizations in monkeys, no systematic test of this functional homology has yet been reported in humans. In the present study, we used functional MRI to compare brain activations when subjects produced affective vocalizations in the form of exclamations versus non-affective vocalizations with similar pitch contours. We also examined the perception of affective vocalizations by having participants make judgments about either the emotions being conveyed by recorded affective vocalizations or the pitch contours of acoustically similar but non-affective vocalizations. Production of affective vocalizations and matched pitch contours activated a highly overlapping set of brain areas, including the larynx-phonation area of the primary motor cortex and a region of the anterior cingulate cortex that is consistent with the macro-anatomical position of the cingulate vocalization area. This overlap contradicts the dominant view that these areas form two distinct vocal pathways with dissociable functions. Instead, we propose that these brain areas are nodes in a single vocal network, with an emphasis on pitch modulation as a vehicle for affective expression. © The Author (2015). Published by Oxford University Press. For Permissions, please email:
    No preview · Article · Jun 2015 · Social Cognitive and Affective Neuroscience
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    • "When we delivered TMS over the dominant hemisphere, differently from HS, patients had increased MEPs also during the 'syllabic phonation' task that we used to investigate non-symbolic linguistic processes. This apparently contrasts with our proposed abnormal SRN activation pattern in ASD, as phonation and emotional speech are thought to activate preferentially the non-dominant hemisphere (Schulz et al., 2005; Hickok & Poeppel, 2007; Loucks et al., 2007; Simonyan et al., 2009). However, the 'syllabic phonation' task we used consisted of three repeated syllabic phonations implying a simple 'linguistic' task rather than a 'phonation' task, and thus probably requiring dominant hemisphere SRN activation. "

    Full-text · Article · Jan 2015
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    • "f these muscles , Brown et al . ( 2008 ) had subjects perform both non - vocal ( glottal stops ) and vocal ( phonation ) laryn - geal tasks , and showed that the same part of the motor cortex was activated by both types of tasks , leading to a characterization of a multi - functional larynx motor cortex ( for a related set of observa - tions , see Loucks et al . , 2007 and Belyk & Brown , 2014 ) . Two dis - tinct regions of activation were found , namely a ventromedial peak in the primary motor cortex ( BA 4 ) and a dorsolateral peak in the premotor cortex ( BA 6 ) . The two meta - analyses performed in the current study specifically implicated the ventromedial primary motor peak in stuttering , consi"
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    ABSTRACT: Stuttering is a speech disorder characterised by repetitions, prolongations and blocks that disrupt the forward movement of speech. An earlier meta-analysis of brain imaging studies of stuttering (Brown et al., 2005) revealed a general trend towards rightward lateralization of brain activations and hyperactivity in the larynx motor cortex bilaterally. The present study sought not only to update that meta-analysis with recent work but to introduce an important distinction not present in the first study, namely the difference between ‘trait’ and ‘state’ stuttering. The analysis of trait stuttering compares people who stutter (PWS) with people who do not stutter when behaviour is controlled for, i.e., when speech is fluent in both groups. In contrast, the analysis of state stuttering examines PWS during episodes of stuttered speech compared with episodes of fluent speech. Seventeen studies were analysed using activation likelihood estimation. Trait stuttering was characterised by the well-known rightward shift in lateralization for language and speech areas. State stuttering revealed a more diverse pattern. Abnormal activation of larynx and lip motor cortex was common to the two analyses. State stuttering was associated with overactivation in the right hemisphere larynx and lip motor cortex. Trait stuttering was associated with overactivation of lip motor cortex in the right hemisphere but underactivation of larynx motor cortex in the left hemisphere. These results support a large literature highlighting laryngeal and lip involvement in the symptomatology of stuttering, and disambiguate two possible sources of activation in neuroimaging studies of persistent developmental stuttering.
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