Experiments and models of cortical oscillations as a target for noninvasive brain stimulation

Noninvasive brain stimulation is attracting substantial attention due to its potential for safe and effective modulation of brain network dynamics. Promising applications include cognitive enhancement and treatment of disorders of the central nervous system. Recently, targeting of cortical oscillations by brain stimulation with periodic electromagnetic waveforms has emerged as a particularly appealing approach for understanding the causal role of cortical oscillations in human cognition and behavior. Two main approaches exist: repetitive transcranial magnetic stimulation (rTMS) and transcranial alternating current stimulation (tACS); rTMS is more widely used as a research and clinical tool but only recently has it been suggested to selectively engage frequency-matched cortical oscillations. In contrast, tACS is an offspring of transcranial direct current stimulation and has been introduced with the specific aim of engaging cortical oscillations. One of the main lessons that the field of noninvasive brain stimulation has learned over the last few years is that without a mechanistic understanding of how stimulation engages neuronal circuits, little progress can be made toward the rational design of individualized, adaptive stimulation treatments. Computer simulations of cellular and network models from the field of computational neuroscience are a key tool to gain such a mechanistic understanding. However, the insights gained from such modeling strategies can only be fully leveraged when used in tight conjunction with experimental approaches in both human and animal model studies. Here, I provide an in-depth review of the pioneering experimental and computational studies that together provide the basis for understanding how periodic noninvasive brain stimulation targets cortical oscillations to enable the rational design of brain stimulation treatments for disorders associated with specific deficits in cortical oscillations.