Decoding Remapped Spatial Information in the Peri-Saccadic Period.

It has been suggested that, prior to a saccade, visual neurons predictively respond to stimuli that will fall in their receptive fields after completion of the saccade. This saccadic remapping process is thought to compensate for the shift of the visual world across the retina caused by eye movements. To map the timing of this predictive process in the brain, we recorded neural activity using electroencephalography (EEG) during a saccade task. Human participants (male and female) made saccades between two fixation points while covertly attending to oriented gratings briefly presented at various locations on the screen. Data recorded during trials in which participants maintained fixation were used to train classifiers on stimuli in different positions. Subsequently, data collected during saccade trials were used to test for the presence of remapped stimulus information at the post-saccadic retinotopic location in the peri-saccadic period, providing unique insight into when remapped information becomes available. We found that the stimulus could be decoded at the remapped location ∼180 ms post-stimulus onset, but only when the stimulus was presented 100-200 ms before saccade onset. Within this range, we found that the timing of remapping was dictated by stimulus onset rather than saccade onset. We conclude that presenting the stimulus immediately before the saccade allows for optimal integration of the corollary discharge signal with the incoming peripheral visual information, resulting in a remapping of activation to the relevant post-saccadic retinotopic neurons.Significance Statement Each eye movement leads to a shift of the visual world across the retina, such that the visual input before and after the eye movement do not match. Despite this, we perceive the visual world as stable. A predictive mechanism known as saccadic remapping is thought to contribute to this stability. We use a saccade task with time-resolved EEG decoding to obtain a fine-grained analysis of the temporal dynamics of the saccadic remapping process. Probing different stimulus-saccade latencies and an array of stimulus locations, we identify when remapped information becomes available in the visual cortex. We describe a critical window in which feedforward visual information and the preparatory motor signals interact to allow for predictive remapping of a stimulus.

Sensory Drive Modifies Brain Dynamics and the Temporal Integration Window.

Perception is suggested to occur in discrete temporal windows, clocked by cycles of neural oscillations. An important testable prediction of this theory is that individuals' peak frequencies of oscillations should correlate with their ability to segregate the appearance of two successive stimuli. An influential study tested this prediction and showed that individual peak frequency of spontaneously occurring alpha (8-12 Hz) correlated with the temporal segregation threshold between two successive flashes of light [Samaha, J., & Postle, B. R. The speed of alpha-band oscillations predicts the temporal resolution of visual perception. Current Biology, 25, 2985-2990, 2015]. However, these findings were recently challenged [Buergers, S., & Noppeney, U. The role of alpha oscillations in temporal binding within and across the senses. Nature Human Behaviour, 6, 732-742, 2022]. To advance our understanding of the link between oscillations and temporal segregation, we devised a novel experimental approach. Rather than relying entirely on spontaneous brain dynamics, we presented a visual grating before the flash stimuli that is known to induce continuous oscillations in the gamma band (45-65 Hz). By manipulating the contrast of the grating, we found that high contrast induces a stronger gamma response and a shorter temporal segregation threshold, compared to low-contrast trials. In addition, we used a novel tool to characterize sustained oscillations and found that, for half of the participants, both the low- and high-contrast gratings were accompanied by a sustained and phase-locked alpha oscillation. These participants tended to have longer temporal segregation thresholds. Our results suggest that visual stimulus drive, reflected by oscillations in specific bands, is related to the temporal resolution of visual perception.

Hello from the other side: Robust contralateral interference in tactile detection.

Touch is unique among the sensory modalities in that our tactile receptors are spread across the body surface and continuously receive different inputs at the same time. These inputs vary in type, properties, relevance according to current goals, and, of course, location on the body. Sometimes, they must be integrated, and other times set apart and distinguished. Here, we investigate how simultaneous stimulation to different body sites affects tactile cognition. Specifically, we characterized the impact of irrelevant tactile sensations on tactile change detection. To this end, we embedded detection targets amidst ongoing performance, akin to the conditions encountered in everyday life, where we are constantly confronted with new events within ongoing stimuli. In the set of experiments presented here, participants detected a brief intensity change (.04 s) within an ongoing vibrotactile stimulus (1.6 s) that was always presented in a constantly attended location. The intensity change (i.e., the detection target) varied parametrically, from hardly detectable to easily detectable. In half of the trials, irrelevant ongoing stimulation was simultaneously presented to a site across the body midline, but participants were instructed to ignore it. In line with previous bimanual studies employing brief onset targets, we document robust interference on performance due to the irrelevant stimulation at each of the measured body sites (homologous and nonhomologous fingers, and the contralateral ankle). After describing this basic phenomenon, we further examine the conditions under which such interference occurs in three additional tasks. In each task, we honed in on a different aspect of the stimulation protocol (e.g., hand distance, the strength of the irrelevant stimulation, the detection target itself) in order to better understand the principles governing the observed interference effects. Our findings suggest a minimal role for exogenous attentional capture in producing the observed interference effects (Exp. 2), and a principled distribution of attentional resources or sensory integration between body sides (Exps. 3, 4). In our last study (Exp. 4), we presented bilateral tactile targets of varying intensities to both the relevant and irrelevant stimulation sites. We then characterized the degree to which the irrelevant stimulation is also processed. Our results-that participants' perception of target intensity is always proportional to the combined bilateral signal-suggest that both body sites are equally weighed and processed despite clear instructions to attend only the target site. In light of this observation and participants' inability to use selection processes to guide their perception, we propose that bilateral tactile inputs are automatically combined, quite possibly early in the hierarchy of somatosensory processing.

A Role for Bottom-Up Alpha Oscillations in Temporal Integration.

Neural oscillations in the 8-12 Hz alpha band are thought to represent top-down inhibitory control and to influence temporal resolution: Individuals with faster peak frequencies segregate stimuli appearing closer in time. Recently, this theory has been challenged. Here, we investigate a special case in which alpha does not correlate with temporal resolution: when stimuli are presented amidst strong visual drive. Based on findings regarding alpha rhythmogenesis and wave spatial propagation, we suggest that stimulus-induced, bottom-up alpha oscillations play a role in temporal integration. We propose a theoretical model, informed by visual persistence, lateral inhibition, and network refractory periods, and simulate physiologically plausible scenarios of the interaction between bottom-up alpha and the temporal segregation. Our simulations reveal that different features of oscillations, including frequency, phase, and power, can influence temporal perception and provide a theoretically informed starting point for future empirical studies.

Intonation Units in Spontaneous Speech Evoke a Neural Response.

Spontaneous speech is produced in chunks called intonation units (IUs). IUs are defined by a set of prosodic cues and presumably occur in all human languages. Recent work has shown that across different grammatical and sociocultural conditions IUs form rhythms of ∼1 unit per second. Linguistic theory suggests that IUs pace the flow of information in the discourse. As a result, IUs provide a promising and hitherto unexplored theoretical framework for studying the neural mechanisms of communication. In this article, we identify a neural response unique to the boundary defined by the IU. We measured the EEG of human participants (of either sex), who listened to different speakers recounting an emotional life event. We analyzed the speech stimuli linguistically and modeled the EEG response at word offset using a GLM approach. We find that the EEG response to IU-final words differs from the response to IU-nonfinal words even when equating acoustic boundary strength. Finally, we relate our findings to the body of research on rhythmic brain mechanisms in speech processing. We study the unique contribution of IUs and acoustic boundary strength in predicting delta-band EEG. This analysis suggests that IU-related neural activity, which is tightly linked to the classic Closure Positive Shift (CPS), could be a time-locked component that captures the previously characterized delta-band neural speech tracking.SIGNIFICANCE STATEMENT Linguistic communication is central to human experience, and its neural underpinnings are a topic of much research in recent years. Neuroscientific research has benefited from studying human behavior in naturalistic settings, an endeavor that requires explicit models of complex behavior. Usage-based linguistic theory suggests that spoken language is prosodically structured in intonation units. We reveal that the neural system is attuned to intonation units by explicitly modeling their impact on the EEG response beyond mere acoustics. To our understanding, this is the first time this is demonstrated in spontaneous speech under naturalistic conditions and under a theoretical framework that connects the prosodic chunking of speech, on the one hand, with the flow of information during communication, on the other.

Attentional Sampling between Eye Channels.

Our ability to detect targets in the environment fluctuates in time. When individuals focus attention on a single location, the ongoing temporal structure of performance fluctuates at 8 Hz. When task demands require the distribution of attention over two objects defined by their location, color or motion direction, ongoing performance fluctuates at 4 Hz per object. This suggests that distributing attention entails the division of the sampling process found for focused attention. It is unknown, however, at what stage of the processing hierarchy this sampling occurs, and whether attentional sampling depends on awareness. Here, we show that unaware selection between the two eyes leads to rhythmic sampling. We presented a display with a single central object to both eyes and manipulated the presentation of a reset event (i.e., cue) and a detection target to either both eyes (binocular) or separately to the different eyes (monocular). We assume that presenting a cue to one eye biases the selection process to content presented in that eye. Although participants were unaware of this manipulation, target detection fluctuated at 8 Hz under the binocular condition, and at 4 Hz when the right (and dominant) eye was cued. These results are consistent with recent findings reporting that competition between receptive fields leads to attentional sampling and demonstrate that this competition does not rely on aware processes. Furthermore, attentional sampling occurs at an early site of competition among monocular channels, before they are fused in the primary visual cortex.

Rhythmic modulation of visual discrimination is linked to individuals' spontaneous motor tempo.

The impact of external rhythmic structure on perception has been demonstrated across different modalities and experimental paradigms. However, recent findings emphasize substantial individual differences in rhythm-based perceptual modulation. Here, we examine the link between spontaneous rhythmic preferences, as measured through the motor system, and individual differences in rhythmic modulation of visual discrimination. As a first step, we measure individual rhythmic preferences using the spontaneous tapping task. Then we assess perceptual rhythmic modulation using a visual discrimination task in which targets can appear either in-phase or out-of-phase with a preceding rhythmic stream of visual stimuli. The tempo of the preceding stream was manipulated over different experimental blocks (0.77 Hz, 1.4 Hz, 2 Hz). We find that visual rhythmic stimulation modulates discrimination performance. The modulation is dependent on the tempo of stimulation, with maximal perceptual benefits for the slowest tempo of stimulation (0.77 Hz). Most importantly, the strength of modulation is also linked to individuals' spontaneous motor tempo. Individuals with slower spontaneous tempi show greater rhythmic modulation compared to individuals with faster spontaneous tempi. This finding suggests that different tempi affect the cognitive system with varying levels of efficiency and that self-generated rhythms impact our ability to utilize rhythmic structure in the environment for guiding perception and performance.

Neural signatures of evidence accumulation in temporal decisions.

Cognitive models of interval timing can be formulated as an accumulation-to-bound process.1-5 However, the physiological manifestation of such processes has not yet been identified. We used electroencephalography (EEG) to measure the neural responses of participants while they performed a temporal bisection task in which they were requested to categorize the duration of visual stimuli as short or long.6 We found that the stimulus-offset and response-locked activity depends on both stimulus duration and the participants' decision. To relate this activity to the underlying cognitive processes, we used a drift-diffusion model.7 The model includes a noisy accumulator starting with the stimulus onset and a decision threshold. According to the model, a stimulus duration will be categorized as "long" if the accumulator reaches the threshold during stimulus presentation. Otherwise, it will be categorized as "short." We found that at the offset of stimulus presentation, an EEG response marks the distance of the accumulator from the threshold. Therefore, this model offers an accurate description of our behavioral data as well as the EEG response using the same two model parameters. We then replicated this finding in an identical experiment conducted in the tactile domain. We also extended this finding to two different temporal ranges (sub- and supra-second). Taken together, the work provides a new way to study the cognitive processes underlying temporal decisions, using a combination of behavior, EEG, and modeling.

Sequences of Intonation Units form a ~ 1 Hz rhythm.

Studies of speech processing investigate the relationship between temporal structure in speech stimuli and neural activity. Despite clear evidence that the brain tracks speech at low frequencies (~ 1 Hz), it is not well understood what linguistic information gives rise to this rhythm. In this study, we harness linguistic theory to draw attention to Intonation Units (IUs), a fundamental prosodic unit of human language, and characterize their temporal structure as captured in the speech envelope, an acoustic representation relevant to the neural processing of speech. IUs are defined by a specific pattern of syllable delivery, together with resets in pitch and articulatory force. Linguistic studies of spontaneous speech indicate that this prosodic segmentation paces new information in language use across diverse languages. Therefore, IUs provide a universal structural cue for the cognitive dynamics of speech production and comprehension. We study the relation between IUs and periodicities in the speech envelope, applying methods from investigations of neural synchronization. Our sample includes recordings from every-day speech contexts of over 100 speakers and six languages. We find that sequences of IUs form a consistent low-frequency rhythm and constitute a significant periodic cue within the speech envelope. Our findings allow to predict that IUs are utilized by the neural system when tracking speech. The methods we introduce here facilitate testing this prediction in the future (i.e., with physiological data).

Beta-Band Activity Is a Signature of Statistical Learning.

Through statistical learning (SL), cognitive systems may discover the underlying regularities in the environment. Testing human adults (n = 35, 21 females), we document, in the context of a classical visual SL task, divergent rhythmic EEG activity in the interstimulus delay periods within patterns versus between patterns (i.e., pattern transitions). Our findings reveal increased oscillatory activity in the beta band (∼20 Hz) at triplet transitions that indexes learning: it emerges with increased pattern repetitions; and importantly, it is highly correlated with behavioral learning outcomes. These findings hold the promise of converging on an online measure of learning regularities and provide important theoretical insights regarding the mechanisms of SL and prediction.SIGNIFICANCE STATEMENT Statistical learning has become a major theoretical construct in cognitive science, providing the primary means by which organisms learn about regularities in the environment. As such, it is a critical building block for basic and higher-order cognitive functions. Here we identify, for the first time, a spectral neural index in the time window before stimulus presentation, which evolves with increased pattern exposure, and is predictive of learning performance. The manifestation of learning that is revealed, not in stimulus processing but in the blank interval between stimuli, makes a direct link between the fields of statistical learning on the one hand and either prediction or consolidation on the other hand, suggesting a possible mechanistic account of visual statistical learning.

When Temporal Certainty Doesn't Help.

In a dynamically changing environment, the ability to capture regularities in our sensory input helps us generate predictions about future events. In most sensory systems, the basic finding is clear: Knowing when something will happen improves performance on it [Nobre, A. C., & van Ede, F. (2017). Anticipated moments: Temporal structure in attention. Nature Reviews Neuroscience, 19, 34-48, 2017]. We here examined the impact of temporal predictions on a less-explored modality: touch. Participants were instructed to detect a brief target embedded in an ongoing vibrotactile stimulus. Unbeknownst to them, the experiment had two timing conditions: In one part, the time of target onset was fixed and thus temporally predictable, whereas in the other, it could appear at a random time within the ongoing stimulation. We found a clear modulation of detection thresholds due to temporal predictability: Contrary to other sensory systems, detecting a predictable tactile target was worse relative to unpredictable targets. We discuss our findings within the framework of tactile suppression.

Where Does Time Go When You Blink?

Retinal input is frequently lost because of eye blinks, yet humans rarely notice these gaps in visual input. Although previous studies focused on the perceptual and neural correlates of diminished awareness to blinks, the impact of these correlates on the perceived time of concurrent events is unknown. Here, we investigated whether the subjective sense of time is altered by spontaneous blinks. We found that participants (N = 22) significantly underestimated the duration of a visual stimulus when a spontaneous blink occurred during stimulus presentation and that this underestimation was correlated with the blink duration of individual participants. Importantly, the effect was not present when durations of an auditory stimulus were judged (N = 23). The results point to a link between spontaneous blinks, previously demonstrated to induce activity suppression in the visual cortex, and a compression of subjective time. They suggest that ongoing encoding within modality-specific sensory cortices, independent of conscious awareness, informs the subjective sense of time.

Feature-Based Attention Samples Stimuli Rhythmically.

Attention supports the allocation of resources to relevant locations and objects in a scene. Under most conditions, several stimuli compete for neural representation. Attention biases neural representation toward the response associated with the attended object [1, 2]. Therefore, an attended stimulus enjoys a neural response that resembles the response to that stimulus in isolation. Factors that determine and generate attentional bias have been researched, ranging from endogenously controlled processes to exogenous capture of attention [1-4]. Recent studies investigate the temporal structure governing attention. When participants monitor a single location, visual-target detection depends on the phase of an ∼8-Hz brain rhythm [5, 6]. When two locations are monitored, performance fluctuates at 4 Hz for each location [7, 8]. The hypothesis is that 4-Hz sampling for two locations may reflect a common sampler that operates at 8 Hz globally, which is divided between relevant locations [5-7, 9]. The present study targets two properties of this phenomenon, called rhythmic-attentional sampling: first, sampling is typically described for selection over different locations. We examined whether rhythmic sampling is limited to selection over space or whether it extends to feature-based attention. Second, we examined whether sampling at 4 Hz results from the division of an 8-Hz rhythm over two objects. We found that two overlapping objects defined by features are sampled at ∼4 Hz per object. In addition, performance on a single object fluctuated at 8 Hz. Rhythmic sampling of features did not result from temporal structure in eye movements.

Neuroscience: A Mechanism for Rhythmic Sampling in Vision.

Ongoing perception ebbs and flows rhythmically. Understanding the source and scope of this phenomenon is an important step in unraveling the foundations of sensory processing. A new study demonstrates that local neuronal interactions generate rhythmic brain activity and correspond to rhythmic performance patterns on a visual-detection task.

Spatial attention and environmental information.

Navigating through our perceptual environment requires constant selection of behaviorally relevant information and irrelevant information. Spatial cues guide attention to information in the environment that is relevant to the current task. How does the amount of information provided by a location cue and irrelevant information influence the deployment of attention and what are the processes underlying this effect? To address these questions, we used a spatial cueing paradigm to measure the relationship between cue predictability (measured in bits of information) and the voluntary attention effect, the benefit in reaction time (RT) because of cueing a target. We found a linear relationship between cue predictability and the attention effect. To analyze the cognitive processes producing this effect, we used a simple RT model, the Linear Ballistic Accumulator model. We found that informative cues reduced the amount of evidence necessary to make a response (the threshold), regardless of the presence of irrelevant information (i.e., distractors). However, a change in the rate of evidence accumulation occurred when distractors were present in the display. Thus, the mechanisms underlying the deployment of attention are exquisitely tuned to the amount and behavioral relevancy of statistical information in the environment. (PsycINFO Database Record

Improved control of exogenous attention in action video game players.

Action video game players (VGPs) have demonstrated a number of attentional advantages over non-players. Here, we propose that many of those benefits might be underpinned by improved control over exogenous (i.e., stimulus-driven) attention. To test this we used an anti-cueing task, in which a sudden-onset cue indicated that the target would likely appear in a separate location on the opposite side of the fixation point. When the time between the cue onset and the target onset was short (40 ms), non-players (nVGPs) showed a typical exogenous attention effect. Their response times were faster to targets presented at the cued (but less probable) location compared with the opposite (more probable) location. VGPs, however, were less likely to have their attention drawn to the location of the cue. When the onset asynchrony was long (600 ms), VGPs and nVGPs were equally able to endogenously shift their attention to the likely (opposite) target location. In order to rule out processing-speed differences as an explanation for this result, we also tested VGPs and nVGPs on an attentional blink (AB) task. In a version of the AB task that minimized demands on task switching and iconic memory, VGPs and nVGPs did not differ in second target identification performance (i.e., VGPs had the same magnitude of AB as nVGPs), suggesting that the anti-cueing results were due to flexible control over exogenous attention rather than to more general speed-of-processing differences.

Extracting the mean size across the visual field in patients with mild, chronic unilateral neglect.

Previous studies suggest that normal vision pools information from groups of objects in a display to extract statistical summaries (e.g., mean size). Here we explored whether patients with mild, chronic left neglect were able to extract statistical summaries on the right and left sides of space in a typical manner. We tested four patients using a visual search task and varied the mean size of a group of circles within the display. On each trial, a single circle first appeared in the center of the screen (the target). This circle varied in size from trial to trial. Then a multi-item display appeared with circles of various sizes grouped together either on the left or right side of the display. The instructions were to search the circles and determine whether the target was present or not. The circles were always accompanied by a group of task-irrelevant triangles that appeared on the opposite side of the display. On half the trials, the mean size of the circles was the size of the target. On the other half the mean size was different from the target. The patients were not told that this was the case, and no explicit report of the statistics was required. The results showed that when the targets were absent patients produced more false alarms to the mean than non-mean size when the circles were on the left (neglected) side of the display. This finding demonstrates that statistical information was implicitly extracted from the left group of circles. However, summary statistics on the right side were not limited to the circles. Rather it appears that participants pooled the distractors with the target circles, yielding a skewed statistical summary on the right side. These findings are discussed as they relate to statistical summary processing, visual search and segregation of right and left items in patients with mild, chronic unilateral neglect.

Cholinergic enhancement reduces orientation-specific surround suppression but not visual crowding.

Acetylcholine (ACh) reduces the spatial spread of excitatory fMRI responses in early visual cortex and receptive field size of V1 neurons. We investigated the perceptual consequences of these physiological effects of ACh with surround suppression and crowding, two phenomena that involve spatial interactions between visual field locations. Surround suppression refers to the reduction in perceived stimulus contrast by a high-contrast surround stimulus. For grating stimuli, surround suppression is selective for the relative orientations of the center and surround, suggesting that it results from inhibitory interactions in early visual cortex. Crowding refers to impaired identification of a peripheral stimulus in the presence of flankers and is thought to result from excessive integration of visual features. We increased synaptic ACh levels by administering the cholinesterase inhibitor donepezil to healthy human subjects in a placebo-controlled, double-blind design. In Experiment 1, we measured surround suppression of a central grating using a contrast discrimination task with three conditions: (1) surround grating with the same orientation as the center (parallel), (2) surround orthogonal to the center, or (3) no surround. Contrast discrimination thresholds were higher in the parallel than in the orthogonal condition, demonstrating orientation-specific surround suppression (OSSS). Cholinergic enhancement decreased thresholds only in the parallel condition, thereby reducing OSSS. In Experiment 2, subjects performed a crowding task in which they reported the identity of a peripheral letter flanked by letters on either side. We measured the critical spacing between the targets and flanking letters that allowed reliable identification. Cholinergic enhancement with donepezil had no effect on critical spacing. Our findings suggest that ACh reduces spatial interactions in tasks involving segmentation of visual field locations but that these effects may be limited to early visual cortical processing.

Action video game experience reduces the cost of switching tasks.

Video game expertise has been shown to have beneficial effects for visual attention processes, but the effects of action video game playing on executive functions, such as task switching and filtering out distracting information, are less well understood. In the main experiment presented here, video game players (VGPs) and nonplayers (nVGPs) switched between two tasks of unequal familiarity: a familiar task of responding in the direction indicated by an arrow, and a novel task of responding in the opposite direction. nVGPs had large response time costs for switching from the novel task to the familiar task, and small costs for switching from the familiar task to the novel task, replicating prior findings. However, as compared to the nVGPs, VGPs were more facile in switching between tasks, producing overall smaller and more symmetric switching costs, suggesting that experience with action video games produces improvements in executive functioning. In contrast, VGPs and nVGPs did not differ in filtering out the irrelevant flanking stimuli or in remembering details of aurally presented stories. The lack of global differences between the groups suggests that the improved task-switching performance seen in VGPs was not due to differences in global factors, such as VGPs being more motivated than nVGPs.

Voluntary and involuntary attention vary as a function of impulsivity.

In the present study we examined, first, whether voluntary and involuntary attention manifest differently in people who differ in impulsivity (measured with the Barratt Impulsivity Scale). For Experiment 1, we used the spatial cueing task with informative and noninformative spatial cues to probe voluntary and involuntary attention, respectively. We found that participants with high impulsivity scores exhibited larger involuntary attention effects, whereas participants with low impulsivity scores exhibited larger voluntary attention effects. For Experiment 2, we used the correlated-flanker task to determine whether the differences between groups in Experiment 1 were due to high-impulsive participants being less sensitive to the display contingencies or to high-impulsive participants having a greater spread of spatial attention. Surprisingly, high-impulsive participants showed a greater sensitivity to contingencies in the environment (correlated-flanker effect). Our results illustrate one situation in which involuntary attention associated with high impulsivity can play a useful role.