Transcranial magnetic stimulation of visual cortex in memory: cortical state, interference and reactivation of visual content in memory

Memory for perceptual events includes the neural representation of the sensory information at short or longer time scales. Recent transcranial magnetic stimulation (TMS) studies of human visual cortex provided evidence that sensory cortex contributes to memory functions. In this review, we provide an exhaustive overview of these studies and ascertain how well the available evidence supports the idea of a causal role of sensory cortex in memory retention and retrieval. We discuss the validity and implications of the studies using a number of methodological and theoretical criteria that are relevant for brain stimulation of visual cortex. While most studies applied TMS to visual cortex to interfere with memory functions, a handful of pioneering studies used TMS to 'reactivate' memories in visual cortex. Interestingly, similar effects of TMS on memory were found in different memory tasks, which suggests that different memory systems share a neural mechanism of memory in visual cortex. At the same time, this neural mechanism likely interacts with higher order brain areas. Based on this overview and evaluation, we provide a first attempt to an integrative framework that describes how sensory processes contribute to memory in visual cortex, and how higher order areas contribute to this mechanism.