Perceptual skills can be improved through practice on a perceptual task, even in adulthood. Visual perceptual learning is known to be mostly specific to the trained retinal location, which is considered as evidence of neural plasticity in retinotopic early visual cortex. Recent findings demonstrate that transfer of learning to untrained locations can occur under some specific training procedures. Here, we evaluated whether exogenous attention facilitates transfer of perceptual learning to untrained locations, both adjacent to the trained locations (Experiment 1) and distant from them (Experiment 2). The results reveal that attention facilitates transfer of perceptual learning to untrained locations in both experiments, and that this transfer occurs both within and across visual hemifields. These findings show that training with exogenous attention is a powerful regime that is able to overcome the major limitation of location specificity.

Pupil size is an easily accessible, noninvasive online indicator of various perceptual and cognitive processes. Pupil measurements have the potential to reveal continuous processing dynamics throughout an experimental trial, including anticipatory responses. However, the relatively sluggish (∼2 s) response dynamics of pupil dilation make it challenging to connect changes in pupil size to events occurring close together in time. Researchers have used models to link changes in pupil size to specific trial events, but such methods have not been systematically evaluated. Here we developed and evaluated a general linear model (GLM) pipeline that estimates pupillary responses to multiple rapid events within an experimental trial. We evaluated the modeling approach using a sample dataset in which multiple sequential stimuli were presented within 2-s trials. We found: (1) Model fits improved when the pupil impulse response function (puRF) was fit for each observer. PuRFs varied substantially across individuals but were consistent for each individual. (2) Model fits also improved when pupil responses were not assumed to occur simultaneously with their associated trial events, but could have non-zero latencies. For example, pupil responses could anticipate predictable trial events. (3) Parameter recovery confirmed the validity of the fitting procedures, and we quantified the reliability of the parameter estimates for our sample dataset. (4) A cognitive task manipulation modulated pupil response amplitude. We provide our pupil analysis pipeline as open-source software (Pupil Response Estimation Toolbox: PRET) to facilitate the estimation of pupil responses and the evaluation of the estimates in other datasets.

Visual attention prioritizes the processing of sensory information at specific spatial locations (spatial attention; SA) or with specific feature values (feature-based attention; FBA). SA is well characterized in terms of behavior, brain activity, and temporal dynamics—for both top-down (endogenous) and bottom-up (exogenous) spatial orienting. FBA has been thoroughly studied in terms of top-down endogenous orienting, but much less is known about the potential of bottom-up exogenous influences of FBA. Here, in four experiments, we adapted a procedure used in two previous studies that reported exogenous FBA effects, with the goal of replicating and expanding on these findings, especially regarding its temporal dynamics. Unlike the two previous studies, we did not find significant effects of exogenous FBA. This was true (1) whether accuracy or RT was prioritized as the main measure, (2) with precues presented peripherally or centrally, (3) with cue-to-stimulus ISIs of varying durations, (4) with four or eight possible target locations, (5) at different meridians, (6) with either brief or long stimulus presentations, (7) and with either fixation contingent or noncontingent stimulus displays. In the last experiment, a postexperiment participant questionnaire indicated that only a small subset of participants, who mistakenly believed the irrelevant color of the precue indicated which stimulus was the target, exhibited benefits for valid exogenous FBA precues. Overall, we conclude that with the protocol used in the studies reporting exogenous FBA, the exogenous stimulus-driven influence of FBA is elusive at best, and that FBA is primarily a top-down, goal-driven process.

Right before we move our eyes, visual performance and neural responses for the saccade target are enhanced. This effect, presaccadic attention, is considered to prioritize the saccade target and to enhance behavioral performance for the saccade target. Recent evidence has shown that presaccadic attention modulates the processing of feature information. Hitherto, it remains unknown whether presaccadic modulations on feature information are flexible, to improve performance for the task at hand, or automatic, so that they alter the featural representation similarly regardless of the task. Using a masking procedure, here we report that presaccadic attention can either improve or impair performance depending on the spatial frequency content of the visual input. These counterintuitive modulations were significant at a time window right before saccade onset. Furthermore, merely deploying covert attention within the same temporal interval without preparing a saccade did not affect performance. This study reveals that presaccadic attention not only prioritizes the saccade target, but also automatically modifies its featural representation.

The ability to act quickly to a threat is a key skill for survival. Under awareness, threat-related emotional information, such as an angry or fearful face, has not only perceptual advantages but also guides rapid actions such as eye movements. Emotional information that is suppressed from awareness still confers perceptual and attentional benefits. However, it is unknown whether suppressed emotional information can directly guide actions, or whether emotional information has to enter awareness to do so. We suppressed emotional faces from awareness using continuous flash suppression and tracked eye gaze position. Under successful suppression, as indicated by objective and subjective measures, gaze moved towards fearful faces, but away from angry faces. Our findings reveal that: (1) threat-related emotional stimuli can guide eye movements in the absence of visual awareness; (2) threat-related emotional face information guides distinct oculomotor actions depending on the type of threat conveyed by the emotional expression.

Training chronic, cortically-blind (CB) patients on a coarse [left-right] direction discrimination and integration (CDDI) task recovers performance on this task at trained, blind field locations. However, fine direction difference (FDD) thresholds remain elevated at these locations, limiting the usefulness of recovered vision in daily life. Here, we asked if this FDD impairment can be overcome by training CB subjects with endogenous, feature-based attention (FBA) cues. Ten CB subjects were recruited and trained on CDDI and FDD with an FBA cue or FDD with a neutral cue. After completion of each training protocol, FDD thresholds were re-measured with both neutral and FBA cues at trained, blind-field locations and at corresponding, intact-field locations. In intact portions of the visual field, FDD thresholds were lower when tested with FBA than neutral cues. Training subjects in the blind field on the CDDI task improved FDD performance to the point that a threshold could be measured, but these locations remained impaired relative to the intact field. FDD training with neutral cues resulted in better blind field FDD thresholds than CDDI training, but thresholds remained impaired relative to intact field levels, regardless of testing cue condition. Importantly, training FDD in the blind field with FBA lowered FDD thresholds relative to CDDI training, and allowed the blind field to reach thresholds similar to the intact field, even when FBA trained subjects were tested with a neutral rather than FBA cue. Finally, FDD training appeared to also recover normal integration thresholds at trained, blind-field locations, providing an interesting double dissociation with respect to CDDI training. In summary, mechanisms governing FBA appear to function normally in both intact and impaired regions of the visual field following V1 damage. Our results mark the first time that FDD thresholds in CB fields have been seen to reach intact field levels of performance. Moreover, FBA can be leveraged during visual training to recover normal, fine direction discrimination and integration performance at trained, blind-field locations, potentiating visual recovery of more complex and precise aspects of motion perception in cortically-blinded fields.

It is well established that attention improves performance on many visual tasks. However, for more than 100 years, psychologists, philosophers, and neurophysiologists have debated its phenomenology—whether attention actually changes one’s subjective experience. Here, we show that it is possible to objectively and quantitatively investigate the effects of attention on subjective experience. First, we review evidence showing that attention alters the appearance of many static and dynamic basic visual dimensions, which mediate changes in appearance of higher-level perceptual aspects. Then, we summarize current views on how attention alters appearance. These findings have implications for our understanding of perception and attention, illustrating that attention affects not only how we perform in visual tasks, but actually alters our experience of the visual world.

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.

Temporal attention, the prioritization of information at a specific point in time, improves visual performance, but it is unknown whether it does so to the same extent across the visual field. This knowledge is necessary to establish whether temporal attention compensates for heterogeneities in discriminability and speed of processing across the visual field. Discriminability and rate of information accrual depend on eccentricity as well as on polar angle, a characteristic known as performance fields. Spatial attention improves speed of processing more at locations at which discriminability is lower and information accrual is slower, but it improves discriminability to the same extent across isoeccentric locations. Here we asked whether temporal attention benefits discriminability in a similar or differential way across the visual field. Observers were asked to report the orientation of one of two targets presented at different points in time at the same spatial location (fovea, right horizontal meridian, or upper vertical meridian, blocked). Temporal attention improved discriminability and shortened reaction times at the foveal and each parafoveal location similarly. These results provide evidence that temporal attention is similarly effective at multiple locations in the visual field. Consequently, at the tested locations, performance fields are preserved with temporal orienting of attention.

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.

Amblyopia, a developmental disorder of vision, affects many aspects of spatial vision as well as motion perception and some cognitive skills. Current models of amblyopic vision based on known neurophysiological deficiencies have yet to provide an understanding of the wide range of amblyopic perceptual losses. Visual spatial attention is known to enhance performance in a variety of detection and discrimination tasks in visually typical humans and nonhuman primates. We investigated whether and how voluntary spatial attention affected psychophysical performance in amblyopic macaques. Full-contrast response functions for motion direction discrimination were measured for each eye of six monkeys: five amblyopic and one control. We assessed whether the effect of a valid spatial cue on performance corresponded to a change in contrast gain, a leftward shift of the function, or response gain, an upward scaling of the function. Our results showed that macaque amblyopes benefit from a valid spatial cue. Performance with amblyopic eyes viewing showed enhancement of both contrast and response gain whereas fellow and control eyes’ performance showed only contrast gain. Reaction time analysis showed no speed accuracy trade- off in any case. The valid spatial cue improved contrast sensitivity for the amblyopic eye, effectively eliminating the amblyopic contrast sensitivity deficit. These results suggest that engaging endogenous spatial attention may confer substantial benefit to amblyopic vision.

We investigated the attentional repulsion effect—stimuli appear displaced further away from attended locations—in three experiments: one with exogenous (involuntary) attention, and two with endogenous (voluntary) attention with different attention-field sizes. It has been proposed that differences in attention-field size can account for qualitative differences in neural responses elicited by attended stimuli. We used psychophysical comparative judgments and manipulated either exogenous attention via peripheral cues or endogenous attention via central cues and a demanding rapid serial visual presentation task. We manipulated the attention field size of endogenous attention by presenting streams of letters at two specific locations or at two of many possible locations during each block. We found a robust attentional repulsion effect in all three experiments: with endogenous and exogenous attention and with both attention-field sizes. These findings advance our understanding of the influence of spatial attention on the perception of visual space and help relate this repulsion effect to possible neurophysiological correlates.

The ability to swiftly detect and prioritize the processing of relevant information around us is critical for the way we interact with our environment. Selective attention is a key mechanism that serves this purpose, improving performance in numerous visual tasks. Reflexively attending to sudden information helps detect impeding threat or danger, a possible reason why emotion modulates the way selective attention affects perception. For instance, the sudden appearance of a fearful face potentiates the effects of exogenous (involuntary, stimulus-driven) attention on performance. Internal states such as trait anxiety can also modulate the impact of attention on early visual processing. However, attention does not only improve performance; it also alters the way visual information appears to us, e.g. by enhancing perceived contrast. Here we show that emotion potentiates the effects of exogenous attention on both performance and perceived contrast. Moreover, we found that trait anxiety mediates these effects, with stronger influences of attention and emotion in anxious observers. Finally, changes in performance and appearance correlated with each other, likely reflecting common attentional modulations. Altogether, our findings show that emotion and anxiety interact with selective attention to truly alter how we see.

Endogenous and exogenous visuospatial attention both alter spatial resolution, but they operate via distinct mechanisms. In texture segmentation tasks, exogenous attention inflexibly increases resolution even when detrimental for the task at hand and does so by modulating second-order processing. Endogenous attention is more flexible and modulates resolution to benefit performance according to task demands, but it is unknown whether it also operates at the second-order level. To answer this question, we measured performance on a second-order texture segmentation task while independently manipulating endogenous and exogenous attention. Observers discriminated a second- order texture target at several eccentricities. We found that endogenous attention improved performance uniformly across eccentricity, suggesting a flexible mechanism that can increase or decrease resolution based on task demands. In contrast, exogenous attention improved performance in the periphery but impaired it at central retinal locations, consistent with an inflexible resolution enhancement. Our results reveal that endogenous and exogenous attention both alter spatial resolution by differentially modulating second-order processing.

Attention affects visual perception at target locations via the amplification of stimuli signal strength, perceptual performance and perceived contrast. Behavioral and neural correlates of attention can be observed when attention is both covertly and overtly oriented (with or without accompanying eye movements). Previous studies have demonstrated that at the grand-average level, lateralization of Event Related Potentials (ERP) is associated with attentional facilitation at cued, relative to uncued locations. Yet, the correspondence between ERP lateralization and behavior has not been established at the single-subject level. Specifically, it is an open question whether inter-individual differences in the neural manifestation of attentional orienting can predict differences in perception. Here, we addressed this question by examining the correlation between ERP lateralization and visual sensitivity at attended locations. Participants were presented with a cue indicating where a low-contrast grating patch target will appear, following a delay of varying durations. During this delay, while participants were waiting for the target to appear, a task-irrelevant checkerboard probe was presented briefly and bilaterally. ERP was measured relative to the onset of this probe. In separate blocks, participants were requested to report detection of a low-contrast target either by making a fast eye-movement toward the target (overt orienting), or by pressing a button (covert orienting). Results show that in the covert orienting condition, ERP lateralization of individual participants was positively correlated with their mean visual sensitivity for the target. But, no such correlation was found in the overt orienting condition. We conclude that ERP lateralization of individual participants can predict their performance on a covert, but not an overt, target detection task.

Attention allows us to select relevant sensory information for preferential processing. Behaviourally, it improves performance in various visual tasks. One prominent effect of attention is the modulation of performance in tasks that involve the visual system’s spatial resolution. Physiologically, attention modulates neuronal responses and alters the profile and position of receptive fields near the attended location. Here, we develop a hypothesis linking the behavioural and electrophysiological evidence. The proposed framework seeks to explain how these receptive field changes enhance the visual system’s effective spatial resolution and how the same mechanisms may also underlie attentional effects on the representation of spatial information.

At attended locations emotion and attention interact to benefit contrast sensitivity, a basic visual dimension. Whether there are associated costs at unattended locations is unknown. Furthermore, emotion and attention affect response time, and anxiety modulates these effects. We investigated how trait-anxiety influences the interaction of emotion and attention on contrast sensitivity. On each trial, non-predictive pre-cues (neutral or fearful faces) directed exogenous attention to four contrast- varying, tilted stimuli (Gabor patches). Attention was cued toward the target (valid), a distracter (invalid), or distributed over all locations. Observers discriminated target orientation, and completed self-report measures of anxiety. Effects of fearful expressions were mediated by trait anxiety. Only high-trait-anxious individuals showed decreased target contrast sensitivity after attention was diverted to a distracter by a fearful cue, and anxiety score correlated with degree of impairment across participants. This indicates that increasing anxiety exacerbates threat-related attentional costs to visual perception, hampering processing at non-threat-related locations.

Rapid manipulation of the attention field (i.e. the location and spread of visual spatial attention) is a critical aspect of human cognition, and previous research on spatial attention in individuals with autism spectrum disorders (ASD) has produced inconsistent results. In a series of three psychophysical experiments, we evaluated claims in the literature that individuals with ASD exhibit a deficit in voluntarily controlling the deployment and size of the spatial attention field. We measured the spatial distribution of performance accuracies and reaction times to quantify the sizes and locations of the attention field, with and without spatial uncertainty (i.e. the lack of predictability concerning the spatial position of the upcoming stimulus). We found that high-functioning adults with autism exhibited slower reaction times overall with spatial uncertainty, but the effects of attention on performance accuracies and reaction times were indistinguishable between individuals with autism and typically developing individuals in all three experiments. These results provide evidence of intact endogenous spatial attention function in high-functioning adults with ASD, suggesting that atypical endogenous attention cannot be a latent characteristic of autism in general.

What you see depends not only on where you are looking but also on where you will look next. The pre-saccadic attention shift is an automatic enhancement of visual sensitivity at the target of the next saccade. We investigated whether and how perceptual factors independent of the oculomotor plan modulate pre-saccadic attention within and across trials. Observers made saccades to one (the target) of six patches of moving dots and discriminated a brief luminance pulse (the probe) that appeared at an unpredictable location. Sensitivity to the probe was always higher at the target’s location (spatial attention), and this attention effect was stronger if the previous probe appeared at the previous target’s location. Furthermore, sensitivity was higher for probes moving in directions similar to the target’s direction (feature- based attention), but only when the previous probe moved in the same direction as the previous target. Therefore, implicit cognitive processes permeate pre-saccadic attention, so that–contingent on recent experience–it flexibly distributes resources to potentially relevant locations and features.

We investigated the impact of the preparation of reach movements on visual perception by simultaneously quantifying both an objective measure of visual sensitivity and the subjective experience of apparent contrast. Using a two-by-two alternative forced choice task, observers compared the orientation (clockwise or counterclockwise) and the contrast (higher or lower) of a Standard Gabor and a Test Gabor, the latter of which was presented during reach preparation, at the reach target location or the opposite location. Discrimination performance was better overall at the reach target than at the opposite location. Perceived contrast increased continuously at the target relative to the opposite location during reach preparation, that is, after the onset of the cue indicating the reach target. The finding that performance and appearance do not evolve in parallel during reach preparation points to a distinction with saccade preparation, for which we have shown previously there is a parallel temporal evolution of performance and appearance. Yet akin to saccade preparation, this study reveals that overall reach preparation enhances both visual performance and appearance.

Analyzing the factors that determine our choice of visual search strategy may shed light on visual behavior in everyday situations. Previous results suggest that increasing task difficulty leads to more systematic search paths. Here we analyze observers’ eye movements in an ‘‘easy’’ conjunction search task and a ‘‘difficult’’ shape search task to study visual search strategies in stereoscopic search displays with virtual depth induced by binocular disparity. Standard eye-movement variables, such as fixation duration and initial saccade latency, as well as new measures proposed here, such as saccadic step size, relative saccadic selectivity, and x􏰇y target distance, revealed systematic effects on search dynamics in the horizontal-vertical plane throughout the search process. We found that in the ‘‘easy’’ task, observers start with the processing of display items in the display center immediately after stimulus onset and subsequently move their gaze outwards, guided by extrafoveally perceived stimulus color. In contrast, the ‘‘difficult’’ task induced an initial gaze shift to the upper- left display corner, followed by a systematic left-right and top-down search process. The only consistent depth effect was a trend of initial saccades in the easy task with smallest displays to the items closest to the observer. The results demonstrate the utility of eye- movement analysis for understanding search strategies and provide a first step toward studying search strategies in actual 3D scenarios.

Sudden changes in the visual periphery can automatically draw attention to their locations. For example, the brief flash of a single object (a “cue”) rapidly enhances contrast sensitivity for subsequent stimuli in its vicinity. Feature singletons (e.g., a red circle among green circles) can also capture attention in a variety of tasks. Here, we evaluate whether a peripheral cue that enhances contrast sensitivity when it appears alone has a similar effect when it appears as a color singleton, with the same stimuli and task. In four experiments we asked observers to report the orientation of a target Gabor stimulus, which was preceded by an uninformative cue array consisting either of a single disk or of 16 disks containing a color or luminance singleton. Accuracy was higher and contrast thresholds lower when the single cue appeared at or near the target’s location, compared with farther away. The color singleton also modulated performance but to a lesser degree and only when it appeared exactly at the target’s location. Thus, this is the first study to demonstrate that cueing by color singletons, like single cues, can enhance sensory signals at an early stage of processing.

Covert attention and perceptual learning enhance perceptual performance. The relation between these two mechanisms is largely unknown. Previously, we showed that manipulating involuntary, exogenous spatial attention during training improved performance at trained and untrained locations, thus overcoming the typical location specificity. Notably, attention-induced transfer only occurred for high stimulus contrasts, at the upper asymptote of the psychometric function (i.e., via response gain). Here, we investigated whether and how voluntary, endogenous attention, the top-down and goal-based type of covert visual attention, influences perceptual learning. Twenty-six participants trained in an orientation discrimination task at two locations: half of participants received valid endogenous spatial precues (attention group), while the other half received neutral precues (neutral group). Before and after training, all participants were tested with neutral precues at two trained and two untrained locations. Within each session, stimulus contrast varied on a trial basis from very low (2%) to very high (64%). Performance was fit by a Weibull psychometric function separately for each day and location. Performance improved for both groups at the trained location, and unlike training with exogenous attention, at the threshold level (i.e., via contrast gain). The neutral group exhibited location specificity: Thresholds decreased at the trained locations, but not at the untrained locations. In contrast, participants in the attention group showed significant location transfer: Thresholds decreased to the same extent at both trained and untrained locations. These results indicate that, similar to exogenous spatial attention, endogenous spatial attention induces location transfer, but influences contrast gain instead of response gain.

Perceptual skills can be improved through practice on a perceptual task, even in adulthood. Visual perceptual learning is known to be mostly specific to the trained retinal location, which is considered as evidence of neural plasticity in retinotopic early visual cortex. Recent findings demonstrate that transfer of learning to untrained locations can occur under some specific training procedures. Here, we evaluated whether exogenous attention facilitates transfer of perceptual learning to untrained locations, both adjacent to the trained locations (Experiment 1) and distant from them (Experiment 2). The results reveal that attention facilitates transfer of perceptual learning to untrained locations in both experiments, and that this transfer occurs both within and across visual hemifields. These findings show that training with exogenous attention is a powerful regime that is able to overcome the major limitation of location specificity.

Pupil size is an easily accessible, noninvasive online indicator of various perceptual and cognitive processes. Pupil measurements have the potential to reveal continuous processing dynamics throughout an experimental trial, including anticipatory responses. However, the relatively sluggish (∼2 s) response dynamics of pupil dilation make it challenging to connect changes in pupil size to events occurring close together in time. Researchers have used models to link changes in pupil size to specific trial events, but such methods have not been systematically evaluated. Here we developed and evaluated a general linear model (GLM) pipeline that estimates pupillary responses to multiple rapid events within an experimental trial. We evaluated the modeling approach using a sample dataset in which multiple sequential stimuli were presented within 2-s trials. We found: (1) Model fits improved when the pupil impulse response function (puRF) was fit for each observer. PuRFs varied substantially across individuals but were consistent for each individual. (2) Model fits also improved when pupil responses were not assumed to occur simultaneously with their associated trial events, but could have non-zero latencies. For example, pupil responses could anticipate predictable trial events. (3) Parameter recovery confirmed the validity of the fitting procedures, and we quantified the reliability of the parameter estimates for our sample dataset. (4) A cognitive task manipulation modulated pupil response amplitude. We provide our pupil analysis pipeline as open-source software (Pupil Response Estimation Toolbox: PRET) to facilitate the estimation of pupil responses and the evaluation of the estimates in other datasets.

Visual attention prioritizes the processing of sensory information at specific spatial locations (spatial attention; SA) or with specific feature values (feature-based attention; FBA). SA is well characterized in terms of behavior, brain activity, and temporal dynamics—for both top-down (endogenous) and bottom-up (exogenous) spatial orienting. FBA has been thoroughly studied in terms of top-down endogenous orienting, but much less is known about the potential of bottom-up exogenous influences of FBA. Here, in four experiments, we adapted a procedure used in two previous studies that reported exogenous FBA effects, with the goal of replicating and expanding on these findings, especially regarding its temporal dynamics. Unlike the two previous studies, we did not find significant effects of exogenous FBA. This was true (1) whether accuracy or RT was prioritized as the main measure, (2) with precues presented peripherally or centrally, (3) with cue-to-stimulus ISIs of varying durations, (4) with four or eight possible target locations, (5) at different meridians, (6) with either brief or long stimulus presentations, (7) and with either fixation contingent or noncontingent stimulus displays. In the last experiment, a postexperiment participant questionnaire indicated that only a small subset of participants, who mistakenly believed the irrelevant color of the precue indicated which stimulus was the target, exhibited benefits for valid exogenous FBA precues. Overall, we conclude that with the protocol used in the studies reporting exogenous FBA, the exogenous stimulus-driven influence of FBA is elusive at best, and that FBA is primarily a top-down, goal-driven process.

Right before we move our eyes, visual performance and neural responses for the saccade target are enhanced. This effect, presaccadic attention, is considered to prioritize the saccade target and to enhance behavioral performance for the saccade target. Recent evidence has shown that presaccadic attention modulates the processing of feature information. Hitherto, it remains unknown whether presaccadic modulations on feature information are flexible, to improve performance for the task at hand, or automatic, so that they alter the featural representation similarly regardless of the task. Using a masking procedure, here we report that presaccadic attention can either improve or impair performance depending on the spatial frequency content of the visual input. These counterintuitive modulations were significant at a time window right before saccade onset. Furthermore, merely deploying covert attention within the same temporal interval without preparing a saccade did not affect performance. This study reveals that presaccadic attention not only prioritizes the saccade target, but also automatically modifies its featural representation.

The ability to act quickly to a threat is a key skill for survival. Under awareness, threat-related emotional information, such as an angry or fearful face, has not only perceptual advantages but also guides rapid actions such as eye movements. Emotional information that is suppressed from awareness still confers perceptual and attentional benefits. However, it is unknown whether suppressed emotional information can directly guide actions, or whether emotional information has to enter awareness to do so. We suppressed emotional faces from awareness using continuous flash suppression and tracked eye gaze position. Under successful suppression, as indicated by objective and subjective measures, gaze moved towards fearful faces, but away from angry faces. Our findings reveal that: (1) threat-related emotional stimuli can guide eye movements in the absence of visual awareness; (2) threat-related emotional face information guides distinct oculomotor actions depending on the type of threat conveyed by the emotional expression.

Training chronic, cortically-blind (CB) patients on a coarse [left-right] direction discrimination and integration (CDDI) task recovers performance on this task at trained, blind field locations. However, fine direction difference (FDD) thresholds remain elevated at these locations, limiting the usefulness of recovered vision in daily life. Here, we asked if this FDD impairment can be overcome by training CB subjects with endogenous, feature-based attention (FBA) cues. Ten CB subjects were recruited and trained on CDDI and FDD with an FBA cue or FDD with a neutral cue. After completion of each training protocol, FDD thresholds were re-measured with both neutral and FBA cues at trained, blind-field locations and at corresponding, intact-field locations. In intact portions of the visual field, FDD thresholds were lower when tested with FBA than neutral cues. Training subjects in the blind field on the CDDI task improved FDD performance to the point that a threshold could be measured, but these locations remained impaired relative to the intact field. FDD training with neutral cues resulted in better blind field FDD thresholds than CDDI training, but thresholds remained impaired relative to intact field levels, regardless of testing cue condition. Importantly, training FDD in the blind field with FBA lowered FDD thresholds relative to CDDI training, and allowed the blind field to reach thresholds similar to the intact field, even when FBA trained subjects were tested with a neutral rather than FBA cue. Finally, FDD training appeared to also recover normal integration thresholds at trained, blind-field locations, providing an interesting double dissociation with respect to CDDI training. In summary, mechanisms governing FBA appear to function normally in both intact and impaired regions of the visual field following V1 damage. Our results mark the first time that FDD thresholds in CB fields have been seen to reach intact field levels of performance. Moreover, FBA can be leveraged during visual training to recover normal, fine direction discrimination and integration performance at trained, blind-field locations, potentiating visual recovery of more complex and precise aspects of motion perception in cortically-blinded fields.

It is well established that attention improves performance on many visual tasks. However, for more than 100 years, psychologists, philosophers, and neurophysiologists have debated its phenomenology—whether attention actually changes one’s subjective experience. Here, we show that it is possible to objectively and quantitatively investigate the effects of attention on subjective experience. First, we review evidence showing that attention alters the appearance of many static and dynamic basic visual dimensions, which mediate changes in appearance of higher-level perceptual aspects. Then, we summarize current views on how attention alters appearance. These findings have implications for our understanding of perception and attention, illustrating that attention affects not only how we perform in visual tasks, but actually alters our experience of the visual world.

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.

Temporal attention, the prioritization of information at a specific point in time, improves visual performance, but it is unknown whether it does so to the same extent across the visual field. This knowledge is necessary to establish whether temporal attention compensates for heterogeneities in discriminability and speed of processing across the visual field. Discriminability and rate of information accrual depend on eccentricity as well as on polar angle, a characteristic known as performance fields. Spatial attention improves speed of processing more at locations at which discriminability is lower and information accrual is slower, but it improves discriminability to the same extent across isoeccentric locations. Here we asked whether temporal attention benefits discriminability in a similar or differential way across the visual field. Observers were asked to report the orientation of one of two targets presented at different points in time at the same spatial location (fovea, right horizontal meridian, or upper vertical meridian, blocked). Temporal attention improved discriminability and shortened reaction times at the foveal and each parafoveal location similarly. These results provide evidence that temporal attention is similarly effective at multiple locations in the visual field. Consequently, at the tested locations, performance fields are preserved with temporal orienting of attention.

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.

Amblyopia, a developmental disorder of vision, affects many aspects of spatial vision as well as motion perception and some cognitive skills. Current models of amblyopic vision based on known neurophysiological deficiencies have yet to provide an understanding of the wide range of amblyopic perceptual losses. Visual spatial attention is known to enhance performance in a variety of detection and discrimination tasks in visually typical humans and nonhuman primates. We investigated whether and how voluntary spatial attention affected psychophysical performance in amblyopic macaques. Full-contrast response functions for motion direction discrimination were measured for each eye of six monkeys: five amblyopic and one control. We assessed whether the effect of a valid spatial cue on performance corresponded to a change in contrast gain, a leftward shift of the function, or response gain, an upward scaling of the function. Our results showed that macaque amblyopes benefit from a valid spatial cue. Performance with amblyopic eyes viewing showed enhancement of both contrast and response gain whereas fellow and control eyes’ performance showed only contrast gain. Reaction time analysis showed no speed accuracy trade- off in any case. The valid spatial cue improved contrast sensitivity for the amblyopic eye, effectively eliminating the amblyopic contrast sensitivity deficit. These results suggest that engaging endogenous spatial attention may confer substantial benefit to amblyopic vision.

We investigated the attentional repulsion effect—stimuli appear displaced further away from attended locations—in three experiments: one with exogenous (involuntary) attention, and two with endogenous (voluntary) attention with different attention-field sizes. It has been proposed that differences in attention-field size can account for qualitative differences in neural responses elicited by attended stimuli. We used psychophysical comparative judgments and manipulated either exogenous attention via peripheral cues or endogenous attention via central cues and a demanding rapid serial visual presentation task. We manipulated the attention field size of endogenous attention by presenting streams of letters at two specific locations or at two of many possible locations during each block. We found a robust attentional repulsion effect in all three experiments: with endogenous and exogenous attention and with both attention-field sizes. These findings advance our understanding of the influence of spatial attention on the perception of visual space and help relate this repulsion effect to possible neurophysiological correlates.

The ability to swiftly detect and prioritize the processing of relevant information around us is critical for the way we interact with our environment. Selective attention is a key mechanism that serves this purpose, improving performance in numerous visual tasks. Reflexively attending to sudden information helps detect impeding threat or danger, a possible reason why emotion modulates the way selective attention affects perception. For instance, the sudden appearance of a fearful face potentiates the effects of exogenous (involuntary, stimulus-driven) attention on performance. Internal states such as trait anxiety can also modulate the impact of attention on early visual processing. However, attention does not only improve performance; it also alters the way visual information appears to us, e.g. by enhancing perceived contrast. Here we show that emotion potentiates the effects of exogenous attention on both performance and perceived contrast. Moreover, we found that trait anxiety mediates these effects, with stronger influences of attention and emotion in anxious observers. Finally, changes in performance and appearance correlated with each other, likely reflecting common attentional modulations. Altogether, our findings show that emotion and anxiety interact with selective attention to truly alter how we see.

Endogenous and exogenous visuospatial attention both alter spatial resolution, but they operate via distinct mechanisms. In texture segmentation tasks, exogenous attention inflexibly increases resolution even when detrimental for the task at hand and does so by modulating second-order processing. Endogenous attention is more flexible and modulates resolution to benefit performance according to task demands, but it is unknown whether it also operates at the second-order level. To answer this question, we measured performance on a second-order texture segmentation task while independently manipulating endogenous and exogenous attention. Observers discriminated a second- order texture target at several eccentricities. We found that endogenous attention improved performance uniformly across eccentricity, suggesting a flexible mechanism that can increase or decrease resolution based on task demands. In contrast, exogenous attention improved performance in the periphery but impaired it at central retinal locations, consistent with an inflexible resolution enhancement. Our results reveal that endogenous and exogenous attention both alter spatial resolution by differentially modulating second-order processing.

Attention affects visual perception at target locations via the amplification of stimuli signal strength, perceptual performance and perceived contrast. Behavioral and neural correlates of attention can be observed when attention is both covertly and overtly oriented (with or without accompanying eye movements). Previous studies have demonstrated that at the grand-average level, lateralization of Event Related Potentials (ERP) is associated with attentional facilitation at cued, relative to uncued locations. Yet, the correspondence between ERP lateralization and behavior has not been established at the single-subject level. Specifically, it is an open question whether inter-individual differences in the neural manifestation of attentional orienting can predict differences in perception. Here, we addressed this question by examining the correlation between ERP lateralization and visual sensitivity at attended locations. Participants were presented with a cue indicating where a low-contrast grating patch target will appear, following a delay of varying durations. During this delay, while participants were waiting for the target to appear, a task-irrelevant checkerboard probe was presented briefly and bilaterally. ERP was measured relative to the onset of this probe. In separate blocks, participants were requested to report detection of a low-contrast target either by making a fast eye-movement toward the target (overt orienting), or by pressing a button (covert orienting). Results show that in the covert orienting condition, ERP lateralization of individual participants was positively correlated with their mean visual sensitivity for the target. But, no such correlation was found in the overt orienting condition. We conclude that ERP lateralization of individual participants can predict their performance on a covert, but not an overt, target detection task.

Attention allows us to select relevant sensory information for preferential processing. Behaviourally, it improves performance in various visual tasks. One prominent effect of attention is the modulation of performance in tasks that involve the visual system’s spatial resolution. Physiologically, attention modulates neuronal responses and alters the profile and position of receptive fields near the attended location. Here, we develop a hypothesis linking the behavioural and electrophysiological evidence. The proposed framework seeks to explain how these receptive field changes enhance the visual system’s effective spatial resolution and how the same mechanisms may also underlie attentional effects on the representation of spatial information.

At attended locations emotion and attention interact to benefit contrast sensitivity, a basic visual dimension. Whether there are associated costs at unattended locations is unknown. Furthermore, emotion and attention affect response time, and anxiety modulates these effects. We investigated how trait-anxiety influences the interaction of emotion and attention on contrast sensitivity. On each trial, non-predictive pre-cues (neutral or fearful faces) directed exogenous attention to four contrast- varying, tilted stimuli (Gabor patches). Attention was cued toward the target (valid), a distracter (invalid), or distributed over all locations. Observers discriminated target orientation, and completed self-report measures of anxiety. Effects of fearful expressions were mediated by trait anxiety. Only high-trait-anxious individuals showed decreased target contrast sensitivity after attention was diverted to a distracter by a fearful cue, and anxiety score correlated with degree of impairment across participants. This indicates that increasing anxiety exacerbates threat-related attentional costs to visual perception, hampering processing at non-threat-related locations.

Rapid manipulation of the attention field (i.e. the location and spread of visual spatial attention) is a critical aspect of human cognition, and previous research on spatial attention in individuals with autism spectrum disorders (ASD) has produced inconsistent results. In a series of three psychophysical experiments, we evaluated claims in the literature that individuals with ASD exhibit a deficit in voluntarily controlling the deployment and size of the spatial attention field. We measured the spatial distribution of performance accuracies and reaction times to quantify the sizes and locations of the attention field, with and without spatial uncertainty (i.e. the lack of predictability concerning the spatial position of the upcoming stimulus). We found that high-functioning adults with autism exhibited slower reaction times overall with spatial uncertainty, but the effects of attention on performance accuracies and reaction times were indistinguishable between individuals with autism and typically developing individuals in all three experiments. These results provide evidence of intact endogenous spatial attention function in high-functioning adults with ASD, suggesting that atypical endogenous attention cannot be a latent characteristic of autism in general.

What you see depends not only on where you are looking but also on where you will look next. The pre-saccadic attention shift is an automatic enhancement of visual sensitivity at the target of the next saccade. We investigated whether and how perceptual factors independent of the oculomotor plan modulate pre-saccadic attention within and across trials. Observers made saccades to one (the target) of six patches of moving dots and discriminated a brief luminance pulse (the probe) that appeared at an unpredictable location. Sensitivity to the probe was always higher at the target’s location (spatial attention), and this attention effect was stronger if the previous probe appeared at the previous target’s location. Furthermore, sensitivity was higher for probes moving in directions similar to the target’s direction (feature- based attention), but only when the previous probe moved in the same direction as the previous target. Therefore, implicit cognitive processes permeate pre-saccadic attention, so that–contingent on recent experience–it flexibly distributes resources to potentially relevant locations and features.

We investigated the impact of the preparation of reach movements on visual perception by simultaneously quantifying both an objective measure of visual sensitivity and the subjective experience of apparent contrast. Using a two-by-two alternative forced choice task, observers compared the orientation (clockwise or counterclockwise) and the contrast (higher or lower) of a Standard Gabor and a Test Gabor, the latter of which was presented during reach preparation, at the reach target location or the opposite location. Discrimination performance was better overall at the reach target than at the opposite location. Perceived contrast increased continuously at the target relative to the opposite location during reach preparation, that is, after the onset of the cue indicating the reach target. The finding that performance and appearance do not evolve in parallel during reach preparation points to a distinction with saccade preparation, for which we have shown previously there is a parallel temporal evolution of performance and appearance. Yet akin to saccade preparation, this study reveals that overall reach preparation enhances both visual performance and appearance.

Analyzing the factors that determine our choice of visual search strategy may shed light on visual behavior in everyday situations. Previous results suggest that increasing task difficulty leads to more systematic search paths. Here we analyze observers’ eye movements in an ‘‘easy’’ conjunction search task and a ‘‘difficult’’ shape search task to study visual search strategies in stereoscopic search displays with virtual depth induced by binocular disparity. Standard eye-movement variables, such as fixation duration and initial saccade latency, as well as new measures proposed here, such as saccadic step size, relative saccadic selectivity, and x􏰇y target distance, revealed systematic effects on search dynamics in the horizontal-vertical plane throughout the search process. We found that in the ‘‘easy’’ task, observers start with the processing of display items in the display center immediately after stimulus onset and subsequently move their gaze outwards, guided by extrafoveally perceived stimulus color. In contrast, the ‘‘difficult’’ task induced an initial gaze shift to the upper- left display corner, followed by a systematic left-right and top-down search process. The only consistent depth effect was a trend of initial saccades in the easy task with smallest displays to the items closest to the observer. The results demonstrate the utility of eye- movement analysis for understanding search strategies and provide a first step toward studying search strategies in actual 3D scenarios.

Sudden changes in the visual periphery can automatically draw attention to their locations. For example, the brief flash of a single object (a “cue”) rapidly enhances contrast sensitivity for subsequent stimuli in its vicinity. Feature singletons (e.g., a red circle among green circles) can also capture attention in a variety of tasks. Here, we evaluate whether a peripheral cue that enhances contrast sensitivity when it appears alone has a similar effect when it appears as a color singleton, with the same stimuli and task. In four experiments we asked observers to report the orientation of a target Gabor stimulus, which was preceded by an uninformative cue array consisting either of a single disk or of 16 disks containing a color or luminance singleton. Accuracy was higher and contrast thresholds lower when the single cue appeared at or near the target’s location, compared with farther away. The color singleton also modulated performance but to a lesser degree and only when it appeared exactly at the target’s location. Thus, this is the first study to demonstrate that cueing by color singletons, like single cues, can enhance sensory signals at an early stage of processing.

Covert attention and perceptual learning enhance perceptual performance. The relation between these two mechanisms is largely unknown. Previously, we showed that manipulating involuntary, exogenous spatial attention during training improved performance at trained and untrained locations, thus overcoming the typical location specificity. Notably, attention-induced transfer only occurred for high stimulus contrasts, at the upper asymptote of the psychometric function (i.e., via response gain). Here, we investigated whether and how voluntary, endogenous attention, the top-down and goal-based type of covert visual attention, influences perceptual learning. Twenty-six participants trained in an orientation discrimination task at two locations: half of participants received valid endogenous spatial precues (attention group), while the other half received neutral precues (neutral group). Before and after training, all participants were tested with neutral precues at two trained and two untrained locations. Within each session, stimulus contrast varied on a trial basis from very low (2%) to very high (64%). Performance was fit by a Weibull psychometric function separately for each day and location. Performance improved for both groups at the trained location, and unlike training with exogenous attention, at the threshold level (i.e., via contrast gain). The neutral group exhibited location specificity: Thresholds decreased at the trained locations, but not at the untrained locations. In contrast, participants in the attention group showed significant location transfer: Thresholds decreased to the same extent at both trained and untrained locations. These results indicate that, similar to exogenous spatial attention, endogenous spatial attention induces location transfer, but influences contrast gain instead of response gain.