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1Division of Human Brain Research, Department of Neuroscience, Karolinska Institute, Stockholm, Sweden; 2Graduate School of Human and Environmental Studies, Kyoto University, Kyoto, Japan; 3C. and O. Vogt Institute for Brain Research, University of Dusseldorf, Germany; and 4Wellcome Department of Imaging Neuroscience, Institute of Neurology, London, United Kingdom
Submitted 23 June 2004; accepted in final form 11 September 2004
We have previously shown that motor areas are engaged when subjects experience illusory limb movements elicited by tendon vibration. However, traditionally cytoarchitectonic area 2 is held responsible for kinesthesia. Here we use functional magnetic resonance imaging and cytoarchitectural mapping to examine whether area 2 is engaged in kinesthesia, whether it is engaged bilaterally because area 2 in non-human primates has strong callosal connections, which other areas are active members of the network for kinesthesia, and if there is a dominance for the right hemisphere in kinesthesia as has been suggested. Ten right-handed blindfolded healthy subjects participated. The tendon of the extensor carpi ulnaris muscles of the right or left hand was vibrated at 80 Hz, which elicited illusory palmar flexion in an immobile hand (illusion). As control we applied identical stimuli to the skin over the processus styloideus ulnae, which did not elicit any illusions (vibration). We found robust activations in cortical motor areas [areas 4a, 4p, 6; dorsal premotor cortex (PMD) and bilateral supplementary motor area (SMA)] and ipsilateral cerebellum during kinesthetic illusions (illusion-vibration). The illusions also activated contralateral area 2 and right area 2 was active in common irrespective of illusions of right or left hand. Right areas 44, 45, anterior part of intraparietal region (IP1) and caudo-lateral part of parietal opercular region (OP1), cortex rostral to PMD, anterior insula and superior temporal gyrus were also activated in common during illusions of right or left hand. These right-sided areas were significantly more activated than the corresponding areas in the left hemisphere. The present data, together with our previous results, suggest that human kinesthesia is associated with a network of active brain areas that consists of motor areas, cerebellum, and the right fronto-parietal areas including high-order somatosensory areas. Furthermore, our results provide evidence for a right hemisphere dominance for perception of limb movement.
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