Modality-Specific Cognitive Function of Medial and Lateral Human Brodmann Area 6

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Abstract
Despite the fact that human Brodmann area 6 (BA6), a traditional “motor” area, is active during higher motor control involving various cognitive operations, the functional specialization within BA6 in the cognitive domain is essentially unknown. Furthermore, its functional relevance in cognition has been questioned because brain activity in BA6 during cognitive tasks has often been explained away as a concomitant, latent motor process. Therefore, we examined the structural-functional relationship of human BA6 in nonmotor cognitive functions and its functional relevance using both functional magnetic resonance imaging (fMRI) and repetitive transcranial magnetic stimulation (rTMS). Subjects performed mental-operation (MO) tasks in which they serially updated verbal and spatial mental representations (MO-v and MO-s). In the fMRI experiments, activity in the medial BA6 was more increased in MO-v, whereas the activity in the lateral BA6 in both hemispheres was more in MO-s. Low-frequency rTMS to the medial BA6 disrupted only the performance of MO-v, whereas rTMS to the lateral BA6 in both hemispheres disrupted only MO-s. Hence the converging results demonstrate a functional double dissociation in which medial BA6 has a critical role in updating verbal information and lateral BA6 has a role in updating spatial information. The present study provides direct physiological evidence of modality-specific cognitive function within human BA6.