Abstract
Single pulses of transcranial magnetic stimulation (TMS) result in distal and long-lasting oscillations, a finding directly challenging the virtual lesion hypothesis. Previous research supporting this finding has primarily come from stimulation of the motor cortex. We have used single-pulse TMS with simultaneous EEG to target seven brain regions, six of which belong to the visual system [left and right primary visual area V1, motion-sensitive human middle temporal cortex, and a ventral temporal region], as determined with functional MRI-guided neuronavigation, and a vertex "control" site to measure the network effects of the TMS pulse. We found the TMS-evoked potential (TMS-EP) over visual cortex consists mostly of site-dependent theta- and alphaband oscillations. These site-dependent oscillations extended beyond the stimulation site to functionally connected cortical regions and correspond to time windows where the EEG responses maximally diverge (40, 200, and 385 ms). Correlations revealed two site-independent oscillations ∼350 ms after the TMS pulse: a theta-band oscillation carried by the frontal cortex, and an alpha-band oscillation over parietal and frontal cortical regions. A manipulation of stimulation intensity at one stimulation site (right hemisphere V1-V3) revealed sensitivity to the stimulation intensity at different regions of cortex, evidence of intensity tuning in regions distal to the site of stimulation. Together these results suggest that a TMS pulse applied to the visual cortex has a complex effect on brain function, engaging multiple brain networks functionally connected to the visual system with both invariant and site-specific spatiotemporal dynamics. With this characterization of TMS, we propose an alternative to the virtual lesion hypothesis. Rather than a technique that simulates lesions, we propose TMS generates natural brain signals and engages functional networks.