Neurons as oscillators.

Regularly spiking neurons can be described as oscillators. In this article we review some of the insights gained from this conceptualization and their relevance for systems neuroscience. First, we explain how a regularly spiking neuron can be viewed as an oscillator and how the phase-response curve (PRC) describes the response of the neuron's spike times to small perturbations. We then discuss the meaning of the PRC for a single neuron's spiking behavior and review the PRCs measured from a variety of neurons in a range of spiking regimes. Next, we show how the PRC can be related to a number of common measures used to quantify neuronal firing, such as the spike-triggered average and the peristimulus histogram. We further show that the response of a neuron to correlated inputs depends on the shape of the PRC. We then explain how the PRC of single neurons can be used to predict neural network behavior. Given the PRC, conduction delays, and the waveform and time course of the synaptic potentials, it is possible to predict neural population behavior such as synchronization. The PRC also allows us to quantify the robustness of the synchronization to heterogeneity and noise. We finally ask how to combine the measured PRCs and the predictions based on PRC to further the understanding of systems neuroscience. As an example, we discuss how the change of the PRC by the neuromodulator acetylcholine could lead to a destabilization of cortical network dynamics. Although all of these studies are grounded in mathematical abstractions that do not strictly hold in biology, they provide good estimates for the emergence of the brain's network activity from the properties of individual neurons. The study of neurons as oscillators can provide testable hypotheses and mechanistic explanations for systems neuroscience.