Transcranial magnetic stimulation reveals the content of visual short-term memory in the visual cortex

Cortical areas involved in sensory analysis are also believed to be involved in short-term storage of that sensory information. Here we investigated whether transcranial magnetic stimulation (TMS) can reveal the content of visual short-term memory (VSTM) by bringing this information to visual awareness. Subjects were presented with two random-dot displays (moving either to the left or to the right) and they were required to maintain one of these in VSTM. In Experiment 1, TMS was applied over the motion-selective area V5/MT+ above phosphene threshold during the maintenance phase. The reported phosphene contained motion features of the memory item, when the phosphene spatially overlapped with memory item. Specifically, phosphene motion was enhanced when the memory item moved in the same direction as the subjects' V5/MT+ baseline phosphene, whereas it was reduced when the motion direction of the memory item was incongruent with that of the baseline V5/MT+ phosphene. There was no effect on phosphene reports when there was no spatial overlap between the phosphene and the memory item. In Experiment 2, VSTM maintenance did not influence the appearance of phosphenes induced from the lateral occipital region. These interactions between VSTM maintenance and phosphene appearance demonstrate that activity in V5/MT+ reflects the motion qualities of items maintained in VSTM. Furthermore, these results also demonstrate that information in VSTM can modulate the pattern of visual activation reaching awareness, providing evidence for the view that overlapping neuronal populations are involved in conscious visual perception and VSTM.