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Cerebral Cortex Advance Access first published online on March 2, 2005
This version published online on May 2, 2005
Cerebral Cortex, doi:10.1093/cercor/bhi061
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Published by Oxford University Press 2005.

Article

Functional Neuroanatomy of Human Vocalization: An H215O PET Study

G.M. Schulz 1*, M. Varga 2, K. Jeffires 2, C.L. Ludlow 3, and A.R. Braun 2

1 Department of Speech and Hearing Science, The George Washington University, USA
2 Language Section, Voice, Speech and Language Branch, NIDCD, NIH, Bethesda, MD 2089, USA
3 Laryngeal and Speech Section, Medical Neurology Branch, NINDS, NIH, Bethesda, MD 2089, USA

* To whom correspondence should be addressed.
G.M. Schulz, E-mail: schulz{at}egwu.edu


  Abstract

Vocalization in lower animals is associated with a well-described visceromotor call system centered on the mesencephalic periacqueductal grey matter (PAG), which is itself regulated by paramedian cortical structures. To determine the role this phylogenetically older system plays in human phonation, we contrasted voiced and unvoiced speech using positron emission tomography and then evaluated functional connectivity of regions that significantly differentiated these conditions. Vocalization was associated with increased and highly correlated activity within the midline structures -- PAG and paramedian cortices -- described in lower mammalian species. Concurrent activation and connectivity of neocortical and subcortical motor regions -- medial and lateral premotor structures and elements of basal ganglia thalamocortical circuitry -- suggest a mechanism by which this system may have come under an increasing degree of voluntary control in humans. Additionally, areas in the temporal lobe and cerebellum were selectively activated during voiced but not unvoiced speech. These regions are functionally coupled to both visceromotor and neocortical motor areas during production of voiced speech, suggesting they may play a central role in self-monitoring and feedback regulation of human phonation.

Keywords: PAG; self-monitoring; species-specific calls; speech production; voice.
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