André M. Cravo

Temporal expectation enhances contrast sensitivity by phase entrainment of low-frequency oscillations in visual cortex.

Although it is increasingly accepted that temporal expectation can modulate early perceptual processing, the underlying neural computations remain unknown. In the present study, we combined a psychophysical paradigm with electrophysiological recordings to investigate the putative contribution of low-frequency oscillatory activity in mediating the modulation of visual perception by temporal expectation. Human participants judged the orientation of brief targets (visual Gabor patterns tilted clockwise or counterclockwise) embedded within temporally regular or irregular streams of noise-patches used as temporal cues. Psychophysical results indicated that temporal expectation enhanced the contrast sensitivity of visual targets. A diffusion model indicated that rhythmic temporal expectation modulated the signal-to-noise gain of visual processing. The concurrent electrophysiological data revealed that the phase of delta oscillations overlying human visual cortex (1–4 Hz) was predictive of the quality of target processing only in regular streams of events. Moreover, in the regular condition, the optimum phase of these perception-predictive oscillations occurred in anticipation of the expected events. Together, these results show a strong correspondence between psychophysical and neurophysiological data, suggesting that the phase entrainment of low-frequency oscillations to external sensory cues can serve as an important and flexible mechanism for enhancing sensory processing.

Temporal Expectation Improves the Quality of Sensory Information

It is increasingly clear that we extract patterns of temporal regularity between events to optimize information processing. Whereas some of the mechanisms for facilitating action preparation and execution have been well documented, much less is understood about whether and how temporal expectations influence visual perception. We used a psychophysical paradigm and computational modeling to investigate the mechanisms by which temporal expectation can modulate visual perception. Visual targets appeared in a stream of noise-patches separated by a fixed (400 ms regular condition) or jittered (200/300/400/500/600 ms irregular condition) intervals. Targets were visual gratings tilted 45° clockwise or counter-clockwise, presented at one of seven contrast levels. Human observers were required to perform an orientation discrimination (i.e., left or right). Psychometric functions for contrast sensitivity fitted for the regular and irregular conditions indicated that temporal expectation modulates perceptual processing by enhancing the contrast sensitivity of visual targets. This increase in the signal strength was accompanied by a reduction in reaction times. A diffusion model indicated that rhythmic temporal expectation enhanced the signal-to-noise gain of the sensory evidence upon which decisions were made. These effects support the idea that temporal structure of external events can entrain the attentional focus and psychophysical data, optimizing the processing of relevant sensory information.

Endogenous modulation of low frequency oscillations by temporal expectations.

Recent studies have associated increasing temporal expectations with synchronization of higher frequency oscillations and suppression of lower frequencies. In this experiment, we explore a proposal that low-frequency oscillations provide a mechanism for regulating temporal expectations. We used a speeded Go/No-go task and manipulated temporal expectations by changing the probability of target presentation after certain intervals. Across two conditions, the temporal conditional probability of target events differed substantially at the first of three possible intervals. We found that reactions times differed significantly at this first interval across conditions, decreasing with higher temporal expectations. Interestingly, the power of theta activity (4–8 Hz), distributed over central midline sites, also differed significantly across conditions at this first interval. Furthermore, we found a transient coupling between theta phase and beta power after the first interval in the condition with high temporal expectation for targets at this time point. Our results suggest that the adjustments in theta power and the phase-power coupling between theta and beta contribute to a central mechanism for controlling neural excitability according to temporal expectations.

Attention Restores Discrete Items to Visual Short-Term Memory

When a memory is forgotten, is it lost forever? Our study shows that selective attention can restore forgotten items to visual short-term memory (VSTM). In our two experiments, all stimuli presented in a memory array were designed to be equally task relevant during encoding. During the retention interval, however, participants were sometimes given a cue predicting which of the memory items would be probed at the end of the delay. This shift in task relevance improved recall for that item. We found that this type of cuing improved recall for items that otherwise would have been irretrievable, providing critical evidence that attention can restore forgotten information to VSTM. Psychophysical modeling of memory performance has confirmed that restoration of information in VSTM increases the probability that the cued item is available for recall but does not improve the representational quality of the memory. We further suggest that attention can restore discrete items to VSTM.

Voluntary action and causality in temporal binding

Previous studies have documented temporal attraction in perceived times of actions and their effects. While some authors argue that voluntary action is a necessary condition for this phenomenon, others claim that the causal relationship between action and effect is the crucial ingredient. In the present study, we investigate voluntary action and causality as the necessary and sufficient conditions for temporal binding. We used a variation of the launching effect proposed by Michotte, in which participants controlled the launch stimulus in some blocks. Volunteers reported causality ratings and estimated the interval between the two events. Our results show dissociations between causality ratings and temporal estimation. While causality ratings are not affected by voluntary action, temporal bindings were only found in the presence of both voluntary action and high causality. Our results indicate that voluntary action and causality are both necessary for the emergence of temporal binding.

The relation between action, predictability and temporal contiguity in temporal binding

Previous studies have documented a subjective temporal attraction between actions and their effects. This finding, named intentional binding, is thought to be the result of a cognitive function that links actions to their consequences. Although several studies have tried to outline the necessary and sufficient conditions for intentional binding, a quantitative comparison between the roles of temporal contiguity, predictability and voluntary action and the evaluation of their interactions is difficult due to the high variability of the temporal binding measurements. In the present study, we used a novel methodology to investigate the properties of intentional binding. Subjects judged whether an auditory stimulus, which could either be triggered by a voluntary finger lift or be presented after a visual temporal marker unrelated to any action, was presented synchronously with a reference stimulus. In three experiments, the predictability, the interval between action and consequence and the presence of action itself were manipulated. The results indicate that (1) action is a necessary condition for temporal binding; (2) a fixed interval between the two events is not sufficient to cause the effect and (3) only in the presence of voluntary action do temporal predictability and contiguity play a significant role in modulating the effect.These findings are discussed in the context of the relationship between intentional binding and temporal expectation.

Visual causality judgments correlate with the phase of alpha oscillations

The detection of causality is essential for our understanding of whether distinct events relate. A central requirement for the sensation of causality is temporal contiguity: As the interval between events increases, causality ratings decrease; for intervals longer than approximately 100 msec, the events start to appear independent. It has been suggested that this effect might be due to perception relying on discrete processing. According to this view, two events may be judged as sequential or simultaneous depending on their temporal relationship within a discrete neuronal process. To assess if alpha oscillations underlie this discrete neuronal process, we investigated how these oscillations modulate the judgment of causality. We used the classic launching effect with concurrent recording of EEG signal. In each trial, a disk moved horizontally toward a second disk at the center of the screen and stopped when they touched each other. After a delay that varied between 0 and 400 msec after contact, the right disk began to move. Participants were instructed to judge whether or not they had a feeling that the first disk caused the movement of the second disk. We found that frontocentral alpha phase significantly biased causality estimates. Moreover, we found that alpha phase was concentrated around different angles for trials in which participants judged events as causally related versus not causally related. We conclude that alpha phase plays a key role in biasing causality judgments.

A psychophysical and computational analysis of the spatio-temporal mechanisms underlying the flash-lag effect.

Several accounts put forth to explain the flash-lag effect (FLE) rely mainly on either spatial or temporal mechanisms. Here we investigated the relationship between these mechanisms by psychophysical and theoretical approaches. In a first experiment we assessed the magnitudes of the FLE and temporal-order judgments performed under identical visual stimulation. The results were interpreted by means of simulations of an artificial neural network, that was also employed to make predictions concerning the FLE. The model predicted that a spatio-temporal mislocalisation would emerge from two, continuous and abrupt-onset, moving stimuli. Additionally, a straightforward prediction of the model revealed that the magnitude of this mislocalisation should be task-dependent, increasing when the use of the abrupt-onset moving stimulus switches from a temporal marker only to both temporal and spatial markers. Our findings confirmed the models predictions and point to an indissoluble interplay between spatial facilitation and processing delays in the FLE.

The Time of Perception and the Other Way Around

The world we perceive is delayed in relation to its flowing content, as well as the outcome of our actions on the world in relation to the moment we decide to act. This mosaic of different latencies permeating both perception and action has to be taken into account critically in order for us to cope with the temporal challenges constantly imposed by the environment. Fundamental notions, such as the sense of agency and causality, depend on the temporal relationship of events occurring in well-defined windows of time. Here, we offer a broad, yet abridged, historical view of some thought-provoking issues concerning the time of perception and action. From the pioneering work of Wundt, Titchener, and Libet to recent findings and ideas related to the employment of visual illusions as psychophysical probes (such as the flash-lag effect), we have tried to expose some problems inherent to the act of measuring the time of both perception and action, and devise possible solutions as well.

Task and exposure time modulate laterality of spatial frequency for faces.

The present study psychophysically investigated the laterality of low spatial frequencies (LSFs) and high spatial frequencies (HSFs) during face recognition at different exposure times. Spatial frequency–filtered faces were presented in a divided visual field at high and low temporal constraints in 2 tasks: face recognition (Experiment 1) and face gender recognition (Experiment 2). Both experiments revealed general primacy in the recognition of LSF over HSF faces. In Experiment 1, LSF and HSF facial information was more efficiently processed in the right and left hemispheres, respectively, and exposure time had no effect. Experiment 2 showed right hemisphere asymmetry for LSF faces at a low temporal constraint. These results suggest that the spatial frequency processing for face recognition is lateralized in the brain hemispheres. However, the contributions of LSFs and HSFs depend on the task and exposure time.