Critical time course of right frontoparietal involvement in mental number space

Neuropsychological, neurophysiological, and neuroimaging studies suggest that right frontoparietal circuits may be necessary for the processing of mental number space, also known as the mental number line (MNL). Here we sought to specify the critical time course of three nodes that have previously been related to MNL processing: right posterior parietal cortex (rPPC), right FEF (rFEF), and right inferior frontal gyrus (rIFG). The effects of single-pulse TMS delivered at 120% distance-adjusted individual motor threshold were investigated in 21 participants, within a window of 0-400 msec (sampling interval = 33 msec) from the onset of a central digit (1-9, 5 excluded). Pulses were delivered in a random order and with equal probability at each time point, intermixed with noTMS trials. To analyze whether and when TMS interfered with MNL processing, we fitted bimodal Gaussian functions to the observed data and measured effects on changes in the Spatial-Numerical Association of Response Codes (SNARC) effect (i.e., an advantage for left- over right-key responses to small numbers and right- over left-key responses to large numbers) and in overall performance efficiency. We found that, during magnitude judgment with unimanual key-press responses, TMS reduced the SNARC effect in the earlier period of the fitted functions (∼25-60 msec) when delivered over rFEF (small and large numbers) and rIFG (small numbers); TMS further reduced the SNARC effect for small numbers in a later period when delivered to rFEF (∼200 msec). In contrast, TMS of rPPC did not interfere with the SNARC effect but generally reduced performance for small numbers and enhanced it for large numbers, thus producing a pattern reminiscent of "neglect" in mental number space. Our results confirm the causal role of an intact right frontoparietal network in the processing of mental number space. They also indicate that rPPC is specifically tied to explicit number magnitude processing and that rFEF and rIFG contribute to interfacing mental visuospatial codes with lateralized response codes. Overall, our findings suggest that both ventral and dorsal frontoparietal circuits are causally involved and functionally connected in the mapping of numbers to space.