Estimating Time and Rhythm by Predicting External Stimuli.

In this chapter, we present recent findings from our group showing that elapsed time, interval timing, and rhythm maintenance might be achieved by the well-known ability of the brain to predict the future states of the world. The difference between predictions and actual sensory evidence is used to generate perceptual and behavioral adjustments that help subjects achieve desired behavioral goals. Concretely, we show that (1) accumulating prediction errors is a plausible strategy humans could use to determine whether a train of consecutive stimuli arrives at regular or irregular intervals. By analyzing the behavior of human and non-human primate subjects performing rhythm perception tasks, we demonstrate that (2) the ability to estimate elapsed time and internally maintain rhythms is shared across primates and humans. Neurophysiological recordings show that (3) the medial premotor cortex engages in rhythm entrainment and maintains oscillatory activity that reveals an internal metronome's spatial and temporal characteristics. Finally, we demonstrate that (4) the amplitude of gamma oscillations within this cortex increases proportionally to the total elapsed time. In conjunction with our most recent experiments, our results suggest that timing might be achieved by an internal simulation of the sensory stimuli and the motor commands that define the timing task that needs to be performed.

Keeping time and rhythm by internal simulation of sensory stimuli and behavioral actions.

Effective behavior often requires synchronizing our actions with changes in the environment. Rhythmic changes in the environment are easy to predict, and we can readily time our actions to them. Yet, how the brain encodes and maintains rhythms is not known. Here, we trained primates to internally maintain rhythms of different tempos and performed large-scale recordings of neuronal activity across the sensory-motor hierarchy. Results show that maintaining rhythms engages multiple brain areas, including visual, parietal, premotor, prefrontal, and hippocampal regions. Each recorded area displayed oscillations in firing rates and oscillations in broadband local field potential power that reflected the temporal and spatial characteristics of an internal metronome, which flexibly encoded fast, medium, and slow tempos. The presence of widespread metronome-related activity, in the absence of stimuli and motor activity, suggests that internal simulation of stimuli and actions underlies timekeeping and rhythm maintenance.