Contribution of the frontal lobe to externally and internally specified verbal responses: fMRI evidence

It has been suggested that within the frontal cortex there is a lateral to medial shift in the control of action, with the lateral premotor area (PMA) involved in externally specified actions and the medial supplementary motor areas (SMA) involved in internally specified actions. Recent brain imaging studies demonstrate, however, that the control of externally and internally specified actions may involve more complex and overlapping networks involving not only the PMA and the SMA, but also the pre-SMA and the lateral prefrontal cortex (PFC). The aim of the present study was to determine whether these frontal regions are differentially involved in the production of verbal responses, when they are externally specified and when they are internally specified. Participants engaged in three overt speaking tasks in which the degree of response specification differed. The tasks involved reading aloud words (externally specified), or generating words aloud from narrow or broad semantic categories (internally specified). Using fMRI, the location and magnitude of the BOLD activity for these tasks was measured in a group of ten participants. Compared with rest, all tasks activated the primary motor area and the SMA-proper, reflecting their common role in speech production. The magnitude of the activity in the PFC (Brodmann area 45), the left PMAv and the pre-SMA increased for word generation, suggesting that each of these three regions plays a role in internally specified action selection. This confirms previous reports concerning the participation of the pre-SMA in verbal response selection. The pattern of activity in PMAv suggests participation in both externally and internally specified verbal actions.