The Spatiotemporal Link of Temporal Expectations: Contextual Temporal Expectation Is Independent of Spatial Attention.

Temporal expectation is the ability to construct predictions regarding the timing of events, based on previously experienced temporal regularities of different types. For example, cue-based expectations are constructed when a cue validly indicates when a target is expected to occur. However, in the absence of such cues, expectations can be constructed based on contextual temporal information, including the onset distribution of the event and recent prior experiences, both providing implicit probabilistic information regarding the timing of the event. It was previously suggested that cue-based temporal expectation is exerted via synchronization of spatially specific neural activity at a predictable time of a target, within receptive fields corresponding to the expected location of the target. Here, we tested whether the same theoretical model holds for contextual temporal effects. Participants (n = 40, 25 females) performed a speeded spatial-cuing detection task with two-thirds valid spatial cues. The hazard-rate function of the target was modulated by varying the foreperiod-the interval between the spatial cue and the target-among trials and was manipulated between groups by changing the interval distribution. Reaction times were analyzed using both frequentist and Bayesian generalized linear mixed models, accounting for hazard and sequential effects. Results showed that the effects of contextual temporal structures on reaction times were independent of spatial attention. This suggests that the spatiotemporal mechanisms, thought to account for cue-based expectation, cannot explain other sources of temporal expectations. We conclude that expectations based on contextual structures have different characteristics than cue-based temporal expectation, suggesting reliance on distinct neural mechanisms.SIGNIFICANCE STATEMENT Temporal expectation is the ability to predict an event onset based on temporal regularities. A neurophysiological model suggested that temporal expectation relies on the synchronization of spatially specific neurons whose receptive fields represent the attended location. This model predicts that temporal expectation would be evident solely within the locus of spatial attention. Existing evidence supported this model for expectation based on associations between a temporal cue and a target, but here we show that it cannot account for temporal expectation that is based on contextual information, that is, the distribution of intervals and recent priors. These findings reveal the existence of different predictive mechanisms for cued and contextual temporal predictions, with the former depending on spatial attention and the latter nonspatially specific.

Sequential effect and temporal orienting in prestimulus oculomotor inhibition.

When faced with unfamiliar circumstances, we often turn to our past experiences with similar situations to shape our expectations. This results in the well-established sequential effect, in which previous trials influence the expectations of the current trial. Studies have revealed that, in addition to the classical behavioral metrics, the inhibition of eye movement could be used as a biomarker to study temporal expectations. This prestimulus oculomotor inhibition is found a few hundred milliseconds prior to predictable events, with a stronger inhibition for predictable than unpredictable events. The phenomenon has been found to occur in various temporal structures, such as rhythms, cue-association, and conditional probability, yet it is still unknown whether it reflects local sequential information of the previous trial. To explore this, we examined the relationship between the sequential effect and the prestimulus oculomotor inhibition. Our results (N = 40) revealed that inhibition was weaker when the previous trial was longer than the current trial, in line with findings of behavioral metrics. These findings indicate that the prestimulus oculomotor inhibition covaries with expectation based on local sequential information, demonstrating the tight connection between this phenomenon and expectation and providing a novel measurement for studying sequential effects in temporal expectation.

Neurofeedback Therapy for Sensory Over-Responsiveness-A Feasibility Study.

Background: Difficulty in modulating multisensory input, specifically the sensory over-responsive (SOR) type, is linked to pain hypersensitivity and anxiety, impacting daily function and quality of life in children and adults. Reduced cortical activity recorded under resting state has been reported, suggestive of neuromodulation as a potential therapeutic modality. This feasibility study aimed to explore neurofeedback intervention in SOR. Methods: Healthy women with SOR (n = 10) underwent an experimental feasibility study comprising four measurement time points (T1­baseline; T2­preintervention; T3­postintervention; T4­follow-up). Outcome measures included resting-state EEG recording, in addition to behavioral assessments of life satisfaction, attaining functional goals, pain sensitivity, and anxiety. Intervention targeted the upregulation of alpha oscillatory power over ten sessions. Results: No changes were detected in all measures between T1 and T2. Exploring the changes in brain activity between T2 and T4 revealed power enhancement in delta, theta, beta, and gamma oscillatory bands, detected in the frontal region (p = 0.03−<0.001; Cohen's d = 0.637−1.126) but not in alpha oscillations. Furthermore, a large effect was found in enhancing life satisfaction and goal attainment (Cohen's d = 1.18; 1.04, respectively), and reduced pain sensitivity and anxiety trait (Cohen's d = 0.70). Conclusion: This is the first study demonstrating the feasibility of neurofeedback intervention in SOR.

Directing Voluntary Temporal Attention Increases Fixational Stability.

Our visual input is constantly changing, but not all moments are equally relevant. Visual temporal attention, the prioritization of visual information at specific points in time, increases perceptual sensitivity at behaviorally relevant times. The dynamic processes underlying this increase are unclear. During fixation, humans make small eye movements called microsaccades, and inhibiting microsaccades improves perception of brief stimuli. Here, we investigated whether temporal attention changes the pattern of microsaccades in anticipation of brief stimuli. Human observers (female and male) judged stimuli presented within a short sequence. Observers were given either an informative precue to attend to one of the stimuli, which was likely to be probed, or an uninformative (neutral) precue. We found strong microsaccadic inhibition before the stimulus sequence, likely due to its predictable onset. Critically, this anticipatory inhibition was stronger when the first target in the sequence (T1) was precued (task-relevant) than when the precue was uninformative. Moreover, the timing of the last microsaccade before T1 and the first microsaccade after T1 shifted such that both occurred earlier when T1 was precued than when the precue was uninformative. Finally, the timing of the nearest pre- and post-T1 microsaccades affected task performance. Directing voluntary temporal attention therefore affects microsaccades, helping to stabilize fixation at the most relevant moments over and above the effect of predictability. Just as saccading to a relevant stimulus can be an overt correlate of the allocation of spatial attention, precisely timed gaze stabilization can be an overt correlate of the allocation of temporal attention.SIGNIFICANCE STATEMENT We pay attention at moments in time when a relevant event is likely to occur. Such temporal attention improves our visual perception, but how it does so is not well understood. Here, we discovered a new behavioral correlate of voluntary, or goal-directed, temporal attention. We found that the pattern of small fixational eye movements called microsaccades changes around behaviorally relevant moments in a way that stabilizes the position of the eyes. Microsaccades during a brief visual stimulus can impair perception of that stimulus. Therefore, such fixation stabilization may contribute to the improvement of visual perception at attended times. This link suggests that, in addition to cortical areas, subcortical areas mediating eye movements may be recruited with temporal attention.

The coordinate system of endogenous spatial attention during smooth pursuit.

A central question in vision is whether spatial attention is represented in an eye-centered (retinotopic) or world-centered (spatiotopic) reference-frame. Most previous studies on this question focused on how coordinates are modulated across saccades. In the present study, we investigated the reference-frame of attention across smooth pursuit eye-movements using a goal-directed saccade task. In two experiments, participants were asked to pursue a moving target while attending to one or two grating stimuli. On each trial, one stimulus was constant in its retinal position and the other was constant in its spatial position. Upon detection of a slight change in stimulus orientation, participants were asked to stop pursuing and perform a fast saccade toward the modified stimulus. In the focused attention condition, they attended one, predefined, stimulus, and in the divided attention condition they attended both. In Experiment 1 the angle of the orientation change marking the target event was constant across participants and conditions. In Experiment 2, the angle was individually adapted to equate performance across participants and conditions. Findings of the two experiments were consistent and showed that the enhancement of mean visual sensitivity in the focused relative to the divided attention condition was similar in magnitude for both retinotopic and spatiotopic targets. This indicates that during smooth pursuit, endogenous attention was proportionally divided between targets in retinotopic and spatiotopic frames of reference.

Oculomotor inhibition reflects temporal expectations.

The accurate extraction of signals out of noisy environments is a major challenge of the perceptual system. Forming temporal expectations and continuously matching them with perceptual input can facilitate this process. In humans, temporal expectations are typically assessed using behavioral measures, which provide only retrospective but no real-time estimates during target anticipation, or by using electrophysiological measures, which require extensive preprocessing and are difficult to interpret. Here we show a new correlate of temporal expectations based on oculomotor behavior. Observers performed an orientation-discrimination task on a central grating target, while their gaze position and EEG were monitored. In each trial, a cue preceded the target by a varying interval ("foreperiod"). In separate blocks, the cue was either predictive or non-predictive regarding the timing of the target. Results showed that saccades and blinks were inhibited more prior to an anticipated regular target than a less-anticipated irregular one. This consistent oculomotor inhibition effect enabled a trial-by-trial classification according to interval-regularity. Additionally, in the regular condition the slope of saccade-rate and drift were shallower for longer than shorter foreperiods, indicating their adjustment according to temporal expectations. Comparing the sensitivity of this oculomotor marker with those of other common predictability markers (e.g. alpha-suppression) showed that it is a sensitive marker for cue-related anticipation. In contrast, temporal changes in conditional probabilities (hazard-rate) modulated alpha-suppression more than cue-related anticipation. We conclude that pre-target oculomotor inhibition is a correlate of temporal predictions induced by cue-target associations, whereas alpha-suppression is more sensitive to conditional probabilities across time.

Transient and sustained effects of stimulus properties on the generation of microsaccades.

Saccades shift the gaze rapidly every few hundred milliseconds from one fixated location to the next, producing a flow of visual input into the visual system even in the absence of changes in the environment. During fixation, small saccades called microsaccades are produced 1-3 times per second, generating a flow of visual input. The characteristics of this visual flow are determined by the timings of the saccades and by the characteristics of the visual stimuli on which they are performed. Previous models of microsaccade generation have accounted for the effects of external stimulation on the production of microsaccades, but they have not considered the effects of the prolonged background stimulus on which microsaccades are performed. The effects of this stimulus on the process of microsaccade generation could be sustained, following its prolonged presentation, or transient, through the visual transients produced by the microsaccades themselves. In four experiments, we varied the properties of the constant displays and examined the resulting modulation of microsaccade properties: their sizes, their timings, and the correlations between properties of consecutive microsaccades. Findings show that displays of higher spatial frequency and contrast produce smaller microsaccades and longer minimal intervals between consecutive microsaccades; and smaller microsaccades are followed by smaller and delayed microsaccades. We explain these findings in light of previous models and suggest a conceptual model by which both sustained and transient effects of the stimulus have central roles in determining the generation of microsaccades.

Gaze Control During Simulator Driving in Adolescents With and Without Attention Deficit Hyperactivity Disorder.

IMPORTANCE: Attention deficit hyperactivity disorder (ADHD) is associated with driving deficits. Visual standards for driving define minimum qualifications for safe driving, including acuity and field of vision, but they do not consider the ability to explore the environment efficiently by shifting the gaze, which is a critical element of safe driving. OBJECTIVE: To examine visual exploration during simulated driving in adolescents with and without ADHD. DESIGN: Adolescents with and without ADHD drove a driving simulator for approximately 10 min while their gaze was monitored. They then completed a battery of questionnaires. SETTING: University lab. PARTICIPANTS: Participants with (n = 16) and without (n = 15) ADHD were included. Participants had a history of neurological disorders other than ADHD and normal or corrected-to-normal vision. Control participants reported not having a diagnosis of ADHD. Participants with ADHD had been previously diagnosed by a qualified professional. OUTCOMES AND MEASURES: We compared the following measures between ADHD and non-ADHD groups: dashboard dwell times, fixation variance, entropy, and fixation duration. RESULTS: Findings showed that participants with ADHD were more restricted in their patterns of exploration than control group participants. They spent considerably more time gazing at the dashboard and had longer periods of fixation with lower variability and randomness. CONCLUSIONS AND RELEVANCE: The results support the hypothesis that adolescents with ADHD engage in less active exploration during simulated driving. WHAT THIS ARTICLE ADDS: This study raises concerns regarding the driving competence of people with ADHD and opens up new directions for potential training programs that focus on exploratory gaze control.

Prestimulus Inhibition of Saccades in Adults With and Without Attention-Deficit/Hyperactivity Disorder as an Index of Temporal Expectations.

Knowing when to expect important events to occur is critical for preparing context-appropriate behavior. However, anticipation is inherently complicated to assess because conventional measurements of behavior, such as accuracy and reaction time, are available only after the predicted event has occurred. Anticipatory processes, which occur prior to target onset, are typically measured only retrospectively by these methods. In this study, we utilized a novel approach for assessing temporal expectations through the dynamics of prestimulus saccades. Results showed that saccades of neurotypical participants were inhibited prior to the onset of stimuli that appeared at predictable compared with less predictable times. No such inhibition was found in most participants with attention-deficit/hyperactivity disorder (ADHD), and particularly not in those who experienced difficulties in sustaining attention over time. These findings suggest that individuals with ADHD, especially those with sustained-attention deficits, have diminished ability to benefit from temporal predictability, and this could account for some of their context-inappropriate behaviors.

Spontaneous microsaccades reflect shifts in covert attention.

Microsaccade rate during fixation is modulated by the presentation of a visual stimulus. When the stimulus is an endogenous attention cue, the ensuing microsaccades tend to be directed toward the cue. This finding has been taken as evidence that microsaccades index the locus of spatial attention. But the vast majority of microsaccades that subjects make are not triggered by visual stimuli. Under natural viewing conditions, spontaneous microsaccades occur frequently (2-3 Hz), even in the absence of a stimulus or a task. While spontaneous microsaccades may depend on low-level visual demands, such as retinal fatigue, image fading, or fixation shifts, it is unknown whether their occurrence corresponds to changes in the attentional state. We developed a protocol to measure whether spontaneous microsaccades reflect shifts in spatial attention. Human subjects fixated a cross while microsaccades were detected from streaming eye-position data. Detection of a microsaccade triggered the appearance of a peripheral ring of grating patches, which were followed by an arrow (a postcue) indicating one of them as the target. The target was either congruent or incongruent (opposite) with respect to the direction of the microsaccade (which preceded the stimulus). Subjects reported the tilt of the target (clockwise or counterclockwise relative to vertical). We found that accuracy was higher for congruent than for incongruent trials. We conclude that the direction of spontaneous microsaccades is inherently linked to shifts in spatial attention.

Continuous flash suppression modulates cortical activity in early visual cortex.

A salient visual stimulus can be rendered invisible by presenting it to one eye while flashing a mask to the other eye. This procedure, called continuous flash suppression (CFS), has been proposed as an ideal way of studying awareness as it can make a stimulus imperceptible for extended periods of time. Previous studies have reported robust suppression of cortical activity in higher visual areas during CFS, but the role of primary visual cortex (V1) is still controversial. In this study, we resolve this controversy on the role of V1 in CFS and also begin characterizing the computational processes underlying CFS. Early visual cortical activity was measured with functional magnetic resonance imaging while human subjects viewed stimuli composed of target and mask, presented to the same or different eyes. Functional MRI responses in early visual cortex were smaller when target and mask were in different eyes compared with the same eye, not only for the lowest contrast target rendered invisible by CFS, but also for higher contrast targets, which were visible even when presented to the eye opposite the mask. We infer that CFS is based on modulating the gain of neural responses, akin to reducing target contrast.

On the timing of overt attention deployment: Eye-movement evidence for the priority accumulation framework.

Most visual-search theories assume that our attention is automatically allocated to the location with the highest priority at any given moment. The Priority Accumulation Framework (PAF) challenges this assumption. It suggests that the priority weight at each location accumulates across sequential events and that evidence for the presence of action-relevant information contributes to determining when attention is deployed to the location with the highest accumulated priority. Here, we tested these hypotheses for overt attention by recording first saccades in a free-viewing spatial-cueing task. We manipulated search difficulty (Experiments 1 and 2) and cue salience (Experiment 2). Standard theories posit that when oculomotor capture by the cue occurs, it is initiated before the search display appears; therefore, these theories predict that the cue's impact on the distribution of first saccades should be independent of search difficulty but influenced by the cue's saliency. By contrast, PAF posits that the cue can bias competition later, after processing of the search display has already started, and therefore predicts that such late impact should increase with both search difficulty and cue salience. The results fully supported PAF's predictions. Our account suggests a distinction between attentional capture and attentional-priority bias that resolves enduring inconsistencies in the attentional-capture literature. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Exposure to temporal variability promotes subsequent adaptation to new temporal regularities.

Noise is intuitively thought to interfere with perceptual learning; However, human and machine learning studies suggest that, in certain contexts, variability may reduce overfitting and improve generalizability. Whereas previous studies have examined the effects of variability in learned stimuli or tasks, it is hitherto unknown what are the effects of variability in the temporal environment. Here, we examined this question in two groups of adult participants (N = 40) presented with visual targets at either random or fixed temporal routines and then tested on the same type of targets at a new nearly-fixed temporal routine. Findings reveal that participants of the random group performed better and adapted quicker following a change in the timing routine, relative to participants of the fixed group. Corroborated with eye-tracking and computational modeling, these findings suggest that prior exposure to temporal variability promotes the formation of new temporal expectations and enhances generalizability in a dynamic environment. We conclude that noise plays an important role in promoting perceptual learning in the temporal domain: rather than interfering with the formation of temporal expectations, noise enhances them. This counterintuitive effect is hypothesized to be achieved through eliminating overfitting and promoting generalizability.

Evidence for content-dependent timing of real-life events during COVID-19 crisis.

How do people estimate the time of past events? A prominent hypothesis suggests that there are multiple timing systems which operate in parallel, depending on circumstances. However, quantitative evidence supporting this hypothesis focused solely on short time-scales (seconds to minutes) and lab-produced events. Furthermore, these studies typically examined the effect of the circumstance and the psychological state of the participant rather than the content of the timed events. Here, we provide, for the first time, support for multiple content-based timing systems when estimating the time of real-life events over long time-scales. The study was conducted during the COVID-19 crisis, which provided a rare opportunity to examine real-life time perception when many were exposed to similar meaningful events. Participants (N = 468) were asked to retrospectively estimate the time that has passed since prominent events, that were either related or unrelated to the pandemic. Results showed an overall time-inflation, which was decreased for events related to the pandemic. This indicates that long-term subjective timing of real-life events exists in multiple systems, which are affected not only by circumstances, but also by content.

Contraction bias in temporal estimation.

When asked to compare the perceptual features of two serially presented objects, participants are often biased to over- or under-estimate the difference in magnitude between the stimuli. Overestimation occurs consistently when a) the two stimuli are relatively small in magnitude and the first stimulus is larger in magnitude than the second; or b) the two stimuli are relatively large in magnitude and the first stimulus is smaller in magnitude than the second; underestimation consistently occurs in the complementary cases. This systematic perceptual bias, known as the contraction bias, was demonstrated for a multitude of perceptual features and in various modalities. Here, we tested whether estimation of time-duration is affected by the contraction bias. In each trial of three experiments (n = 20 each), participants compared the duration of two visually presented stimuli. Findings revealed over- and under-estimation effects as predicted by the contraction bias. Here, we discuss this asymmetry and describe how these findings can be explained via a Bayesian inference framework.

Scalp-recorded induced γ-band responses to auditory stimulation and its correlations with saccadic muscle-activity.

We previously showed that the transient broadband induced gamma-band response in EEG (iGBRtb) appearing around 200-300 ms following a visual stimulus reflects the contraction of extra-ocular muscles involved in the execution of saccades, rather than neural oscillations. Several previous studies reported induced gamma-band responses also following auditory stimulation. It is still an open question whether, similarly to visual paradigms, such auditory paradigms are also sensitive to the saccadic confound. In the current study we address this question using simultaneous eye-tracking and EEG recordings during an auditory oddball paradigm. Subjects were instructed to respond to a rare target defined by sound source location, while fixating on a central screen. Results show that, similar to what was found in visual paradigms, saccadic rate displayed typical temporal dynamics including a post-stimulus decrease followed by an increase. This increase was more moderate, had a longer latency, and was less consistent across subjects than was found in the visual case. Crucially, the temporal dynamics of the induced gamma response were similar to those of saccadic-rate modulation. This suggests that the auditory induced gamma-band responses recorded on the scalp may also be affected by saccadic muscle activity.

Active sensing and overt avoidance: Gaze shifts as a mechanism of predictive avoidance in vision.

Sensory organs are not only involved in passively transmitting sensory input, but are also involved in actively seeking it. Some sensory organs move dynamically to allow highly prioritized input to be detected by their most sensitive parts. Such 'active sensing' systems engage in pursuing relevant input, relying on attentional prioritizations. However, pursuing input may not always be advantageous. Task-irrelevant input may be distracting and interfere with task performance. We hypothesize that an efficient 'active sensing' mechanism should be able to not only pursue relevant input but also to predict irrelevant input and avoid it. Moreover, we hypothesize that this mechanism should be evident even when the task is non-visual and all visual information acts as a distractor. In this study, we demonstrate the existence of a predictive 'overt avoidance' mechanism in vision. In two experiments, participants were asked to perform a continuous mental-arithmetic task while occasionally being presented with task-irrelevant crowded displays limited to one quadrant of a screen. The locations of these visual stimuli were constant within a block but varied between blocks. Results show that gaze was consistently shifted away from the predicted location of distraction, even prior to its appearance, confirming the existence of a predictive 'overt avoidance' mechanism in vision. Based on these findings, we propose a conceptual model to explain how an 'active sensing' system, hardwired to explore, can overcome this drive when presented with distracting information. According to the model, distraction is handled through a dual mechanism of suppression and avoidance processes that are causally linked. This framework demonstrates how perception and motion work together to approach relevant information while avoiding irrelevant distraction.

Prestimulus inhibition of eye movements reflects temporal expectation rather than time estimation.

Eye movements are inhibited prior to the occurrence of temporally predictable events. This 'oculomotor inhibition effect' has been demonstrated with various tasks and modalities. Specifically, it was shown that when intervals between cue and target are fixed, saccade rate prior to the target is lower than when they are varied. However, it is still an open question whether this effect is linked to temporal expectation to the predictable target, or to the duration estimation of the interval preceding it. Here, we examined this question in 20 participants while they performed an implicit temporal expectation and an explicit time estimation task. In each trial, following cue onset, two consecutive grating patches were presented, each preceded by an interval. Temporal expectation was manipulated by setting the first interval duration to be either fixed or varied within each block. Participants were requested to compare either the durations of the two intervals (time estimation), or the tilts of the two grating patches (temporal expectation). Saccade rate, measured prior to the first grating, was lower in the fixed relative to the varied condition of both tasks. This suggests that the inhibition effect is elicited by target predictability and indicates that it is linked to temporal expectation, rather than to time estimation processes. Additionally, this finding suggests that the oculomotor inhibition is independent of motor readiness, as it was elicited even when no response was required. We conclude that the prestimulus oculomotor inhibition effect can be used as a marker of temporal expectation, and discuss its potential underlying mechanisms.

Gender bias in academia: A lifetime problem that needs solutions.

Despite increased awareness of the lack of gender equity in academia and a growing number of initiatives to address issues of diversity, change is slow, and inequalities remain. A major source of inequity is gender bias, which has a substantial negative impact on the careers, work-life balance, and mental health of underrepresented groups in science. Here, we argue that gender bias is not a single problem but manifests as a collection of distinct issues that impact researchers' lives. We disentangle these facets and propose concrete solutions that can be adopted by individuals, academic institutions, and society.

Saccadic spike potentials in gamma-band EEG: characterization, detection and suppression.

Analysis of high-frequency (gamma-band) neural activity by means of non-invasive EEG is gaining increasing interest. However, we have recently shown that a saccade-related spike potential (SP) seriously confounds the analysis of EEG induced gamma-band responses (iGBR), as the SP eludes traditional EEG artifact rejection methods. Here we provide a comprehensive profile of the SP and evaluate methods for its detection and suppression, aiming to unveil true cerebral gamma-band activity. The SP appears consistently as a sharp biphasic deflection of about 22 ms starting at the saccade onset, with a frequency band of approximately 20-90 Hz. On the average, larger saccades elicit higher SP amplitudes. The SP amplitude gradually changes from the extra-ocular channels towards posterior sites with the steepest gradients around the eyes, indicating its ocular source. Although the amplitude and the sign of the SP depend on the choice of reference channel, the potential gradients remain the same and non-zero for all references. The scalp topography is modulated almost exclusively by the direction of saccades, with steeper gradients ipsilateral to the saccade target. We discuss how the above characteristics impede attempts to remove these SPs from the EEG by common temporal filtering, choice of different references, or rejection of contaminated trials. We examine the extent to which SPs can be reliably detected without an eye tracker, assess the degree to which scalp current density derivation attenuates the effect of the SP, and propose a tailored ICA procedure for minimizing the effect of the SP.