Entrainment and Spike-Timing Dependent Plasticity - A Review of Proposed Mechanisms of Transcranial Alternating Current Stimulation.

Specific frequency bands of neural oscillations have been correlated with a range of cognitive and behavioral effects (e.g., memory and attention). The causal role of specific frequencies may be investigated using transcranial alternating current stimulation (tACS), a non-invasive brain stimulation method. TACS involves applying a sinusoidal current between two or more electrodes attached on the scalp, above neural regions that are implicated in cognitive processes of interest. The theorized mechanisms by which tACS affects neural oscillations have implications for the exact stimulation frequency used, as well as its anticipated effects. This review outlines two main mechanisms that are thought to underlie tACS effects - entrainment, and spike-timing dependent plasticity (STDP). Entrainment suggests that the stimulated frequency synchronizes the ongoing neural oscillations, and is thought to be most effective when the stimulated frequency is at or close to the endogenous frequency of the targeted neural network. STDP suggests that stimulation leads to synaptic changes based on the timing of neuronal firing in the target neural network. According to the principles of STDP, synaptic strength is thought to increase when pre-synaptic events occur prior to post-synaptic events (referred to as long-term potentiation, LTP). Conversely, when post-synaptic events occur prior to pre-synaptic events, synapses are thought to be weakened (referred to as long-term depression, LTD). In this review, we summarize the theoretical frameworks and critically review the tACS evidence for each hypothesis. We also discuss whether each mechanism alone can account for tACS effects or whether a combined account is necessary.