Spontaneous α Brain Dynamics Track the Episodic "When".

Across species, neurons track time over the course of seconds to minutes, which may feed the sense of time passing. Here, we asked whether neural signatures of time-tracking could be found in humans. Participants stayed quietly awake for a few minutes while being recorded with magnetoencephalography (MEG). They were unaware they would be asked how long the recording lasted (retrospective time) or instructed beforehand to estimate how long it will last (prospective timing). At rest, rhythmic brain activity is nonstationary and displays bursts of activity in the alpha range (α: 7-14 Hz). When participants were not instructed to attend to time, the relative duration of α bursts linearly predicted individuals' retrospective estimates of how long their quiet wakefulness lasted. The relative duration of α bursts was a better predictor than α power or burst amplitude. No other rhythmic or arrhythmic activity predicted retrospective duration. However, when participants timed prospectively, the relative duration of α bursts failed to predict their duration estimates. Consistent with this, the amount of α bursts was discriminant between prospective and retrospective timing. Last, with a control experiment, we demonstrate that the relation between α bursts and retrospective time is preserved even when participants are engaged in a visual counting task. Thus, at the time scale of minutes, we report that the relative time of spontaneous α burstiness predicts conscious retrospective time. We conclude that in the absence of overt attention to time, α bursts embody discrete states of awareness constitutive of episodic timing.SIGNIFICANCE STATEMENT The feeling that time passes is a core component of consciousness and episodic memory. A century ago, brain rhythms called "α" were hypothesized to embody an internal clock. However, rhythmic brain activity is nonstationary and displays on-and-off oscillatory bursts, which would serve irregular ticks to the hypothetical clock. Here, we discovered that in a given lapse of time, the relative bursting time of α rhythms is a good indicator of how much time an individual will report to have elapsed. Remarkably, this relation only holds true when the individual does not attend to time and vanishes when attending to it. Our observations suggest that at the scale of minutes, α brain activity tracks episodic time.

Rapid encoding of task regularities in the human hippocampus guides sensorimotor timing.

The brain encodes the statistical regularities of the environment in a task-specific yet flexible and generalizable format. Here, we seek to understand this process by bridging two parallel lines of research, one centered on sensorimotor timing, and the other on cognitive mapping in the hippocampal system. By combining functional magnetic resonance imaging (fMRI) with a fast-paced time-to-contact (TTC) estimation task, we found that the hippocampus signaled behavioral feedback received in each trial as well as performance improvements across trials along with reward-processing regions. Critically, it signaled performance improvements independent from the tested intervals, and its activity accounted for the trial-wise regression-to-the-mean biases in TTC estimation. This is in line with the idea that the hippocampus supports the rapid encoding of temporal context even on short time scales in a behavior-dependent manner. Our results emphasize the central role of the hippocampus in statistical learning and position it at the core of a brain-wide network updating sensorimotor representations in real time for flexible behavior.

Sequence Memory in the Hippocampal-Entorhinal Region.

Episodic memories are constructed from sequences of events. When recalling such a memory, we not only recall individual events, but we also retrieve information about how the sequence of events unfolded. Here, we focus on the role of the hippocampal-entorhinal region in processing and remembering sequences of events, which are thought to be stored in relational networks. We summarize evidence that temporal relations are a central organizational principle for memories in the hippocampus. Importantly, we incorporate novel insights from recent studies about the role of the adjacent entorhinal cortex in sequence memory. In rodents, the lateral entorhinal subregion carries temporal information during ongoing behavior. The human homologue is recruited during memory recall where its representations reflect the temporal relationships between events encountered in a sequence. We further introduce the idea that the hippocampal-entorhinal region might enable temporal scaling of sequence representations. Flexible changes of sequence progression speed could underlie the traversal of episodic memories and mental simulations at different paces. In conclusion, we describe how the entorhinal cortex and hippocampus contribute to remembering event sequences-a core component of episodic memory.

Publisher Correction: The effect of attention and working memory on the estimation of elapsed time.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

The effect of attention and working memory on the estimation of elapsed time.

Psychological models of time perception involve attention and memory: while attention typically regulates the flow of events, memory maintains timed events or intervals. The precise, and possibly distinct, roles of attention and memory in time perception remain debated. In this behavioral study, we tested 48 participants in a prospective duration estimation task while they fully attended to time or performed a working memory (WM) task. We report that paying attention to time lengthened perceived duration in the range of seconds to minutes, whereas diverting attention away from time shortened perceived duration. The overestimation due to attending to time did not scale with durations. To the contrary, increasing WM load systematically decreased subjective duration and this effect scaled with durations. Herein, we discuss the dissociation between attention and WM in timing and scalar variability from the perspective of Bayesian models of time estimations.

Event-related potential correlates of stimulus equivalence classes: A study of task order of the equivalence based priming probes with respect to the stimulus equivalence tests, and among the distinct trial types with each other.

This study investigates the influences of: 1) the task order of two stimulus equivalence classes (SEC) probes, and 2) the possible differences within the equivalence trial types. These factors were analyzed together on both behavioral and event-related potentials (ERP) data. Two groups of normal subjects participated in two successive sessions. In the first session, all participants were trained in the baseline relations among visual stimuli (pseudo-words). In the second session, one group performed the matching-to-sample (MTS) equivalence tests before the equivalence-relatedness-priming (EBRP) task, while the other group performed both tasks in reverse order. In the EBRP task related trial types included trained, symmetrical and equivalence relationships while the unrelated trial types included the same stimuli but without relationships. Event related potentials were recorded separately for related and unrelated conditions during the EBRP task. Results showed that response times to related trials were shorter than those to unrelated ones. At the electrophysiological level, two late waveforms were sensitive to the differences among the stimulus pairs of the EBRP task: Both waveforms were larger for the unrelated than the related conditions. Conversely, there were no main influences of the task order or of the trial types with each other. These results provide evidence that 1) the EBRP task exhibits priming effects among the SEC stimuli, 2) the behavioral and electrophysiological effects were similar regardless of whether the EBRP task was done before or after the MTS tests, and 3) there were no differences within the baseline and derived trial types in the EBRP task.