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attention
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temporal attention
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eye movements
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attention
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attention
spatial attention
feature based attention
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action
eye
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perceptual learning
crowding
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*equal author contributions
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speed accuracy tradeoff
2020
Donovan I, Shen A, Tortarolo C, Barbot A & Carrasco M (2020).
Exogenous attention facilitates perceptual learning in visual acuity to untrained stimulus locations and features.
Journal of Vision.
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.
[attention, spatial attention, feature based attention, perceptual learning]
Denison R, Parker J & Carrasco M (2020).
Modeling pupil responses to rapid sequential events.
Behavior Research Methods.
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.
[modeling; eye movements]
2019
Yashar A, Wu X, Chen J & Carrasco M (2019).
Crowding and binding: Not all feature-dimensions behave in the same way.
Psychological Science.
Humans often fail to identify a target because of nearby flankers. The nature and stages at which this crowding occurs are unclear, and whether crowding operates via a common mechanism across visual dimensions is unknown. Usinga dual-estimation report (N = 42), we quantitatively assessed the processing of features alone and in conjunction with another feature both within and between dimensions. Under crowding, observers misreported colors andorientations (i.e., reported a flanker value instead of the target’s value) but averaged the target’s and flankers’ spatialfrequencies (SFs). Interestingly, whereas orientation and color errors were independent, orientation and SF errorswere interdependent. These qualitative differences of errors across dimensions revealed a tight link between crowdingand feature binding, which is contingent on the type of feature dimension. These results and a computational modelsuggest that crowding and misbinding are due to pooling across a joint coding of orientations and SFs but not of colors.
[modeling; crowding]
Kupers ER, Carrasco M & Winawer J (2019).
Modeling visual performance differences 'around' the visual field: A computational observer approach.
PLOS Computational Biology, 15(5): e1007063.
Visual performance depends on polar angle, even when eccentricity is held constant; on many psychophysical tasks observers perform best when stimuli are presented on the horizontal meridian, worst on the upper vertical, and intermediate on the lower vertical meridian. This variation in performance ‘around’ the visual field can be as pronounced as that of doubling the stimulus eccentricity. The causes of these asymmetries in performance are largely unknown. Some factors in the eye, e.g. cone density, are positively correlated with the reported variations in visual performance with polar angle. However, the question remains whether these correlations can quantitatively explain the perceptual differences observed ‘around’ the visual field. To investigate the extent to which the earliest stages of vision–optical quality and cone density–contribute to performance differences with polar angle, we created a computational observer model. The model uses the open-source software package ISETBIO to simulate an orientation discrimination task for which visual performance differs with polar angle. The model starts from the photons emitted by a display, which pass through simulated human optics with fixational eye movements, followed by cone isomerizations in the retina. Finally, we classify stimulus orientation using a support vector machine to learn a linear classifier on the photon absorptions. To account for the 30% increase in contrast thresholds for upper vertical compared to horizontal meridian, as observed psychophysically on the same task, our computational observer model would require either an increase of ~7 diopters of defocus or a reduction of 500% in cone density. These values far exceed the actual variations as a function of polar angle observed in human eyes. Therefore, we conclude that these factors in the eye only account for a small fraction of differences in visual performance with polar angle. Substantial additional asymmetries must arise in later retinal and/or cortical processing.
[performance fields; modeling]
Fernández A, Li HH & Carrasco M (2019).
How exogenous spatial attention affects visual representation.
Journal of Vision.
Orienting covert spatial attention to a target location enhances visual sensitivity and benefits performance in many visual tasks. How these attention-related improvements in performance affect the underlying visual representation of low-level visual features is not fully understood. Here we focus on characterizing how exogenous spatial attention affects the feature representations of orientation and spatial frequency. We asked observers to detect a vertical grating embedded in noise and performed psychophysical reverse correlation. Doing so allowed us to make comparisons with previous studies that utilized the same task and analysis to assess how endogenous attention and presaccadic modulations affect visual representations. We found that exogenous spatial attention improved performance and enhanced the gain of the target orientation without affecting orientation tuning width. Moreover, we found no change in spatial frequency tuning. We conclude that covert exogenous spatial attention alters performance by strictly boosting gain of orientation-selective filters, much like covert endogenous spatial attention.
[spatial attention; psychophysics, eye-tracking, reverse correlation]
Donovan I, Zhou YJ & Carrasco M (2019).
In search of exogenous feature-based attention.
Attention, Perception & Psychophysics.
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.
[exogenous attention, feature based attention]
Waite SA, Grigorian A, Alexander RG, Macknik SL, Carrasco M, Heeger D & Martinez-Conde S (2019).
Analysis of Perceptual Expertise in Radiology – Current Knowledge and a New Perspective.
Frontiers in Human Neuroscience, 13:213.
Radiologists rely principally on visual inspection to detect, describe, and classify findings in medical images. As most interpretive errors in radiology are perceptual in nature, understanding the path to radiologic expertise during image analysis is essential to educate future generations of radiologists. We review the perceptual tasks and challenges in radiologic diagnosis, discuss models of radiologic image perception, consider the application of perceptual learning methods in medical training, and suggest a new approach to understanding perceptional expertise. Specific principled enhancements to educational practices in radiology promise to deepen perceptual expertise among radiologists with the goal of improving training and reducing medical error.
[attention, eye movements, clinical, radiology]
Li HH, Pan J & Carrasco M (2019).
Presaccadic attention improves or impairs performance by enhancing sensitivity to higher spatial frequencies.
Scientific Reports, 9:2659.
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.
[presaccadic attention]
Vetter P*, Badde S*, Phelps EA & Carrasco M (2019).
Emotional faces guide the eyes in the absence of awareness.
eLife.
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.
[eye movements, emotion]
Cavanaugh MR, Barbot A, Carrasco M* & Huxlin KR* (2019).
Feature-based attention potentiates recovery of fine direction discrimination in cortically blind patients.
Neuropsychologia.
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.
[feature based attention, cortical blindness]
Carrasco M & Barbot A (2019).
Spatial attention alters visual appearance.
Current Opinion in Psychology.
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.
[attention, spatial attention]
Michel M, Beck D, Block N, Blumenfeld H, Brown R, Carmel D, Carrasco M, Chirimuuta M, Chun M, Cleeremans A, Dehaene S, Fleming SM, Frith C, Haggard P, He B, Heyes C, Goodale MA, Irvine L, Kawato M, Kentridge R, King J-R, Knight RT, Kouider S, Lamme V, Lamy D, Lau H, Laureys S, LeDoux J, Lin Y-T, Liu K, Macknik SL, Martinez-Conde S, Mashour GA, Melloni L, Miracchi L, Mylopoulos M, Naccache L, Owen AM, Passingham RE, Pessoa L, Peters MAK, Rahnev D, Ro T, Rosenthal D, Sasaki Y, Sergent C, Solovey G, Schiff ND, Seth A, Tallon-Baudry C, Tamietto M, Tong F, van Gaal S, Vlassova A, Watanabe T, Weisberg J, Yan K, Yoshida M (2019).
Opportunities and challenges for a maturing science of consciousness.
Nature Human Behavior.
[review; attention, consciousness]
Carrasco M (2019).
Attention alters appearance.
In Blockheads!: Essays on Ned Block's Philosophy of Mind and Consciousness. A. Pautz and D Stoljar (Eds.) Cambridge, MA: MIT Press.
[book chapter; attention]
Amit R, Abeles D, Carrasco M & Yuval-Greenberg S (2019).
Oculomotor inhibition reflects temporal expectations.
Neuroimage.
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.
[EEG; temporal attention, eye movements]
Fernández A, Denison RN & Carrasco M (2019).
Temporal attention improves perception similarly at foveal and parafoveal locations.
Journal of Vision.
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.
[temporal attention]
Denison R, Yuval-Greenberg S & Carrasco M (2019).
Directing voluntary temporal attention increases fixational stability.
Journal of Neuroscience.
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.
[temporal attention, eye movements]
2018
Pham A, Carrasco M & Kiorpes L (2018).
Endogenous attention improves perception in amblyopic macaques.
Journal of Vision.
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.
[spatial attention, special populations, amblyopia, animal study, motion discrimination, contrast]
Cutrone EK, Heeger DJ & Carrasco M (2018).
On spatial attention and its field size on the repulsion effect.
Journal of 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.
[spatial attention, special populations, amblyopia, animal study, motion discrimination, contrast]
Barbot A & Carrasco M (2018).
Emotion and anxiety potentiate the way attention alters visual appearance.
Scientific Reports.
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.
[spatial attention, emotion, apperance, contrast]
Jigo M & Carrasco M (2018).
Attention alters spatial resolution by modulating second-order processing.
Journal of Vision.
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.
[spatial attention, texture segmentation, spatial resolution]
Hilo-Merkovich R, Carrasco M & Yuval-Greenberg S (2018).
Task performance in covert, but not overt, attention correlates with early laterality of visual evoked potentials.
Neuropsychologia.
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.
[spatial attention]
Carrasco M (2018).
How visual spatial attention alters perception.
Cognitive Processing.
Visual attention is essential for visual perception. Spatial attention allows us to grant priority in processing and selectively process information at a given location. In this paper, I explain how two kinds of spatial attention: covert (allocated to the target location, without accompanying eye movements) and presaccadic (allocated to the location of the upcoming saccade’s target) affect performance and alter appearance. First, I highlight some behavioral and neuroimaging research on covert attention, which alters performance and appearance in many basic visual tasks. Second, I review studies showing that presaccadic attention improves performance and alters appearance at the saccade target location. Further, these modulations change the processing of feature information automatically, even when it is detrimental to the task at hand. We propose that saccade preparation may support transsaccadic integration. Systematically investigating the common and differential characteristics of covert attention and presaccadic attention will continue to further our understanding of the pervasive selective processing of information, which enables us to make sense of our complex visual world.
[review; spatial attention]
Barbot A, Liu S, Kimchi R & Carrasco M (2018).
Attention enhances apparent perceptual organization.
Psychonomic Bulletin & Review.
Perceptual organization and selective attention are two crucial processes that influence how we perceive visual information. The former structures complex visual inputs into coherent units, whereas the later selects relevant information. Attention and perceptual organization can modulate each other, affecting visual processing and performance in various tasks and conditions. Here, we tested whether attention can alter the way multiple elements appear to be perceptually organized. We manipulated covert spatial attention using a rapid serial visual presentation task, and measured perceptual organization of two multielements arrays organized by luminance similarity as rows or columns, at both the attended and unattended locations. We found that the apparent perceptual organization of the multielement arrays is intensified when attended and attenuated when unattended. We ruled out response bias as an alternative explanation. These findings reveal that attention enhances the appearance of perceptual organization, a midlevel vision process, altering the way we perceive our visual environment.
[spatial attention, appearance, perceptual organization]
Denison RN, Adler WT, Carrasco M & Ma WJ (2018).
Humans incorporate attention-dependent uncertainty into perceptual decisions and confidence.
PNAS.
Perceptual decisions are better when they take uncertainty into account. Uncertainty arises not only from the properties of sensory input but also from cognitive sources, such as different levels of attention. However, it is unknown whether humans appropriately adjust for such cognitive sources of uncertainty during perceptual decision-making. Here we show that, in a task in which uncertainty is relevant for performance, human categorization and confidence decisions take into account uncer- tainty related to attention. We manipulated uncertainty in an orientation categorization task from trial to trial using only an attentional cue. The categorization task was designed to disambiguate decision rules that did or did not depend on attention. Using formal model comparison to evaluate decision behavior, we found that category and confidence decision boundaries shifted as a function of attention in an approximately Bayesian fashion. This means that the observer’s attentional state on each trial contributed probabilistically to the decision computation. This responsiveness of an observer’s decisions to attention-dependent uncertainty should improve perceptual decisions in natural vision, in which attention is unevenly distributed across a scene.
[spatial attention, confidence, perceptual decision]
Eckstein MP, Yu C, Sagi D, Carrasco M & Lu Z-L (2018).
Introduction to special issue on perceptual learning.
Vision Research.
[editorial]
Rolfs M, Murray-Smith N & Carrasco M (2018).
Perceptual learning while preparing saccades.
Vision Research.
Traditional perceptual learning protocols rely almost exclusively on long periods of uninterrupted fixation. Taking a first step towards understanding perceptual learning in natural vision, we had observers report the orientation of a briefly flashed stimulus (clockwise or counterclockwise from a reference orientation) presented strictly during saccade preparation at a location offset from the saccade target. For each observer, the saccade direction, stimulus location, and orientation remained the same throughout training. Subsequently, we assessed performance during fixation in three transfer sessions, either at the trained or at an untrained location, and either using an untrained (Experiment 1) or the trained (Experiment 2) stimulus orientation. We modeled the evolution of contrast thresholds (i.e., the stimulus contrast necessary to discriminate its orientation correctly 75% of the time) as an exponential learning curve, and quantified departures from this curve in transfer sessions using two new, complementary measures of transfer costs (i.e., performance decrements after the transition into the Transfer phase). We observed robust perceptual learning and associated transfer costs for untrained locations and orientations. We also assessed if spatial transfer costs were reduced for the remapped location of the presaccadic stimulus—the location the stimulus would have had (but never had) after the saccade. Although the pattern of results at that location differed somewhat from that at the control location, we found no clear evidence for perceptual learning at remapped locations. Using novel, model-based ways to assess learning and transfer costs, our results show that location and feature specificity, hallmarks of perceptual learning, subsist if the target stimulus is presented strictly during saccade preparation throughout training.
[modeling; eye movements, perceptual learning, saccade preparation]
Donovan I & Carrasco M (2018).
Endogenous spatial attention during perceptual learning facilitates location transfer.
Journal of Vision.
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.
[modeling; spatial attention, perceptual learning, orientation discrimination]
2017
Barbot A & Carrasco M (2017).
Attention modifies spatial resolution according to task demands.
Psychological Science.
How does visual attention affect spatial resolution? In texture-segmentation tasks, exogenous (involuntary) attention automatically increases resolution at the attended location, which improves performance where resolution is too low (at the periphery) but impairs performance where resolution is already too high (at central locations). Conversely, endogenous (voluntary) attention improves performance at all eccentricities, which suggests a more flexible mechanism. Here, using selective adaptation to spatial frequency, we investigated the mechanism by which endogenous attention benefits performance in resolution tasks. Participants detected a texture target that could appear at several eccentricities. Adapting to high or low spatial frequencies selectively affected performance in a manner consistent with changes in resolution. Moreover, adapting to high, but not low, frequencies mitigated the attentional benefit at central locations where resolution was too high; this shows that attention can improve performance by decreasing resolution. Altogether, our results indicate that endogenous attention benefits performance by modulating the contribution of high-frequency information in order to flexibly adjust spatial resolution according to task demands.
[spatial attention, texture segmentation, spatial resolution, spatial frequency, adaptation]
Denison RN, Heeger DJ & Carrasco M (2017).
Attention flexibly trades off across points in time.
Psychonomic Bulletin & Review.
Sensory signals continuously enter the brain, raising the question of how perceptual systems handle this constant flow of input. Attention to an anticipated point in time can prioritize visual information at that time. However, how we voluntarily attend across time when there are successive task-relevant stimuli has been barely investigated. We developed a novel experimental protocol that allowed us to assess, for the first time, both the benefits and costs of voluntary temporal attention when perceiving a short sequence of two or three visual targets with predictable timing. We found that when humans directed attention to a cued point in time, their ability to perceive orientation was better at that time but also worse earlier and later. These perceptual tradeoffs across time are analogous to those found across space for spatial attention. We concluded that voluntary attention is limited, and selective, across time.
[temporal attention]
Dankner Y, Shalev L, Carrasco M & Yuval-Greenberg S (2017).
Prestimulus inhibition of saccades in adults with and without ADHD as an index for temporal expectation.
Psychological Science.
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.
[eye movements, microsaccades, special populations]
Mikulak A (2017).
Eye movements reveal temporal expectation deficits in ADHD.
Association for Psychological Science.
[editorial]
Dugué L, Xue AM & Carrasco M (2017).
Distinct perceptual rhythms for feature and conjunction searches.
Journal of Vision.
Feature and conjunction searches are widely used to study attentional deployment. However, the spatiotemporal behavior of attention integration in these tasks remains under debate. Are multiple search stimuli processed in parallel or sequentially? Does sampling of visual information and attentional deployment differ between these two types of search? If so, how? We used an innovative methodology to estimate the distribution of attention on a single-trial basis for feature and conjunction searches. Observers performed feature- and conjunction-search tasks. They had to detect and discriminate a tilted low-spatial-frequency grating among three low-spatial-frequency vertical gratings (feature search) or low-spatial-frequency vertical gratings and high-spatial-frequency tilted gratings (conjunction search). After a variable delay, two probes were flashed at random locations. Performance in reporting the probes was used to infer attentional deployment to those locations. By solving a second-degree equation, we determined the probability of probe report at the most (P1) and least (P2) attended locations on a given trial. Were P1 and P2 equal, we would conclude that attention had been uniformly distributed across all four locations. Otherwise, we would conclude that visual information sampling and attentional deployment had been nonuniformly distributed. Our results show that processing was nonuniformly distributed across the four locations in both searches, and was modulated periodically over time at ;5 Hz for the conjunction search and ;12 Hz for the feature search. We argue that the former corresponds to the periodicity of attentional deployment during the search, whereas the latter corresponds to ongoing sampling of visual information. Because different locations were not simultaneously processed, this study rules out a strict parallel model for both search types.
[spatial attention, visual search]
Dugué L, Merriam EP, Heeger DJ & Carrasco M (2017).
Specific visual subregions of TPJ mediate reorienting of spatial attention.
Cerebral Cortex.
The temporo-parietal junction (TPJ) has been associated with various cognitive and social functions, and is critical for attentional reorienting. Attention affects early visual processing. Neuroimaging studies dealing with such processes have thus far concentrated on striate and extrastriate areas. Here, we investigated whether attention orienting or reorienting modulate activity in visually driven TPJ subregions. For each observer we identified 3 visually responsive subregions within TPJ: 2 bilateral (vTPJant and vTPJpost) and 1 right lateralized (vTPJcent). Cortical activity in these subregions was measured using fMRI while observers performed a 2-alternative forced-choice orientation discrimination task. Covert spatial endogenous (voluntary) or exogenous (involuntary) attention was manipulated using either a central or a peripheral cue with task, stimuli and observers constant. Both endogenous and exogenous attention increased activity for invalidly cued trials in right vTPJpost; only endogenous attention increased activity for invalidly cued trials in left vTPJpost and in right vTPJcent; and neither type of attention modulated either right or left vTPJant. These results demonstrate that vTPJpost and vTPJcent mediate the reorientation of covert attention to task relevant stimuli, thus playing a critical role in visual attention. These findings reveal a differential reorienting cortical response after observers’ attention has been oriented to a given location voluntarily or involuntarily.
[fMRI; spatial attention, reorienting]
Yashar A, White AL, Fang W & Carrasco M (2017).
Feature singletons attract spatial attention independently of feature priming.
Journal of Vision.
People perform better in visual search when the target feature repeats across trials (intertrial feature priming [IFP]). Here, we investigated whether repetition of a feature singleton’s color modulates stimulus-driven shifts of spatial attention by presenting a probe stimulus immediately after each singleton display. The task alternated every two trials between a probe discrimination task and a singleton search task. We measured both stimulus-driven spatial attention (via the distance between the probe and singleton) and IFP (via repetition of the singleton’s color). Color repetition facilitated search performance (IFP effect) when the set size was small. When the probe appeared at the singleton’s location, performance was better than at the opposite location (stimulus-driven attention effect). The magnitude of this attention effect increased with the singleton’s set size (which increases its saliency) but did not depend on whether the singleton’s color repeated across trials, even when the previous singleton had been attended as a search target. Thus, our findings show that repetition of a salient singleton’s color affects performance when the singleton is task relevant and voluntarily attended (as in search trials). However, color repetition does not affect performance when the singleton becomes irrelevant to the current task, even though the singleton does capture attention (as in probe trials). Therefore, color repetition per se does not make a singleton more salient for stimulus- driven attention. Rather, we suggest that IFP requires voluntary selection of color singletons in each consecutive trial.
[spatial attention, visual search, feature singletons]
Li H-H, Rankin J, Rinzel J, Carrasco M & Heeger DJ (2017).
Attention model of binocular rivalry.
PNAS.
When the corresponding retinal locations in the two eyes are presented with incompatible images, a stable percept gives way to perceptual alternations in which the two images compete for perceptual dominance. As perceptual experience evolves dynamically under constant external inputs, binocular rivalry has been used for studying intrinsic cortical computations and for understanding how the brain regulates competing inputs. Converging behavioral and EEG results have shown that binocular rivalry and attention are intertwined: binocular rivalry ceases when attention is diverted away from the rivalry stimuli. In addition, the competing image in one eye suppresses the target in the other eye through a pattern of gain changes similar to those induced by attention. These results require a revision of the current computational theories of binocular rivalry, in which the role of attention is ignored. Here, we provide a computational model of binocular rivalry. In the model, competition between two images in rivalry is driven by both attentional modulation and mutual inhibition, which have distinct selectivity (feature vs. eye of origin) and dynamics (relatively slow vs. relatively fast). The proposed model explains a wide range of phenomena reported in rivalry, including the three hallmarks: (i) binocular rivalry requires attention; (ii) various perceptual states emerge when the two images are swapped between the eyes multiple times per second; (iii) the dominance duration as a function of input strength follows Levelt’s propositions. With a bifurcation analysis, we identified the parameter space in which the model’s behavior was consistent with experimental results.
[modeling; spatial attention, binocular rivalry]
Poletti M, Rucci M & Carrasco M (2017).
Selective attention within the foveola.
Nature Neuroscience.
Efficient control of attentional resources and high-acuity vision are both fundamental for survival. Shifts in visual attention are known to covertly enhance processing at locations away from the center of gaze, where visual resolution is low. It is unknown, however, whether selective spatial attention operates where the observer is already looking—that is, within the high-acuity foveola, the small yet disproportionally important rod-free region of the retina. Using new methods for precisely controlling retinal stimulation, here we show that covert attention flexibly improves and speeds up both detection and discrimination at loci only a fraction of a degree apart within the foveola. These findings reveal a surprisingly precise control of attention and its involvement in fine spatial vision. They show that the commonly studied covert shifts of attention away from the fovea are the expression of a global mechanism that exerts its action across the entire visual field.
[spatial attention, eye movements, microsaccades]
Roberts M, Ashinoff BK, Castellanos FX & Carrasco M (2017).
When attention is intact in adults with ADHD.
Psychonomic Bulletin & Review.
Is covert visuospatial attention—selective processing of information in the absence of eye movements—preserved in adults with attention-deficit/hyperactivity disorder (ADHD)? Previous findings are inconclusive due to inconsistent terminology and suboptimal methodology. To settle this question, we used well-established spatial cueing protocols to investigate the perceptual effects of voluntary and involuntary attention on an orientation discrimination task for a group of adults with ADHD and their neurotypical age-matched and gender-matched controls. In both groups, voluntary attention significantly improved accuracy and decreased reaction times at the relevant location, but impaired accuracy and slowed reaction times at irrelevant locations, relative to a distributed attention condition. Likewise, involuntary attention improved accuracy and speeded responses. Critically, the magnitudes of all these orienting and reorienting attention effects were indistinguishable between groups. Thus, these counterintuitive findings indicate that spatial covert attention remains functionally intact in adults with ADHD.
[spatial attention, special populations]
2013
Anton-Erxleben K & Carrasco M (2013).
Attentional enhancement of spatial resolution: Linking behavioral and neurophysiological evidence.
Nature Reviews Neuroscience.
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.
[review; attention, spatial resolution]
Ferneyhough E, Kim MK, Phelps EA & Carrasco M (2013).
Anxiety modulates the effects of motion and attention on early vision.
Cognition & Emotion.
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.
[spatial attention, emotion, contrast sensitivity, orientation discrimination]
Grubb MA, Behrmann M, Egan R, Minshew NJ, Carrasco M & Heeger DJ (2013).
Endogenous spatial attention: Evidence for intact functioning in adults with autism.
Autism Research.
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.
[spatial attention, special populations]
White AL, Rolfs M & Carrasco M (2013).
Adaptive deployment of spatial and feature-based attention before saccades.
Vision Research.
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.
[spatial attention, feature based attention, presaccadic attention, eye movements, motion perception]
Carrasco M, Eckstein M, Krauzlis R & Verghese (2013).
Attentional modulation: Target selection, active search and cognitive processing.
Vision Research.
[editorial; attention]
Rolfs M, Lawrence BM & Carrasco M (2013).
Reach preparation enhances visual performance and appearance.
Philosophical Transactions of the Royal Society B.
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.
[reach, appearance, orientation discrimination, contrast]
Pomplun M, Garaas TW & Carrasco M (2013).
The effects of task difficulty on visual search strategy in virtual 3D displays.
Journal of Vision.
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 xy 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.
[eye movements, visual search]
White AL, Lunau R & Carrasco M (2013).
The attentional effects of single cues and color singletons on visual sensitivity.
Journal of Experimental Psychology: Human Perception and Performance.
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.
[spatial attention, contrast sensitivity]
overview
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topic
attention
temporal
attention
spatial attention
feature based attention
presaccadic attention
action
eye
movements
reach
perceptual learning
crowding
performance fields
micro-
saccades
texture
segmentation
emotion
binocular
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cortical blindness
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speed accuracy tradeoff
2020
Donovan I, Shen A, Tortarolo C, Barbot A & Carrasco M (2020).
Exogenous attention facilitates perceptual learning in visual acuity to untrained stimulus locations and features.
Journal of Vision.
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.
[attention, spatial attention, feature based attention, perceptual learning]
Denison R, Parker J & Carrasco M (2020).
Modeling pupil responses to rapid sequential events.
Behavior Research Methods.
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.
[modeling; eye movements]
2019
Yashar A, Wu X, Chen J & Carrasco M (2019).
Crowding and binding: Not all feature-dimensions behave in the same way.
Psychological Science.
Humans often fail to identify a target because of nearby flankers. The nature and stages at which this crowding occurs are unclear, and whether crowding operates via a common mechanism across visual dimensions is unknown. Usinga dual-estimation report (N = 42), we quantitatively assessed the processing of features alone and in conjunction with another feature both within and between dimensions. Under crowding, observers misreported colors andorientations (i.e., reported a flanker value instead of the target’s value) but averaged the target’s and flankers’ spatialfrequencies (SFs). Interestingly, whereas orientation and color errors were independent, orientation and SF errorswere interdependent. These qualitative differences of errors across dimensions revealed a tight link between crowdingand feature binding, which is contingent on the type of feature dimension. These results and a computational modelsuggest that crowding and misbinding are due to pooling across a joint coding of orientations and SFs but not of colors.
[modeling; crowding]
Kupers ER, Carrasco M & Winawer J (2019).
Modeling visual performance differences 'around' the visual field: A computational observer approach.
PLOS Computational Biology, 15(5): e1007063.
Visual performance depends on polar angle, even when eccentricity is held constant; on many psychophysical tasks observers perform best when stimuli are presented on the horizontal meridian, worst on the upper vertical, and intermediate on the lower vertical meridian. This variation in performance ‘around’ the visual field can be as pronounced as that of doubling the stimulus eccentricity. The causes of these asymmetries in performance are largely unknown. Some factors in the eye, e.g. cone density, are positively correlated with the reported variations in visual performance with polar angle. However, the question remains whether these correlations can quantitatively explain the perceptual differences observed ‘around’ the visual field. To investigate the extent to which the earliest stages of vision–optical quality and cone density–contribute to performance differences with polar angle, we created a computational observer model. The model uses the open-source software package ISETBIO to simulate an orientation discrimination task for which visual performance differs with polar angle. The model starts from the photons emitted by a display, which pass through simulated human optics with fixational eye movements, followed by cone isomerizations in the retina. Finally, we classify stimulus orientation using a support vector machine to learn a linear classifier on the photon absorptions. To account for the 30% increase in contrast thresholds for upper vertical compared to horizontal meridian, as observed psychophysically on the same task, our computational observer model would require either an increase of ~7 diopters of defocus or a reduction of 500% in cone density. These values far exceed the actual variations as a function of polar angle observed in human eyes. Therefore, we conclude that these factors in the eye only account for a small fraction of differences in visual performance with polar angle. Substantial additional asymmetries must arise in later retinal and/or cortical processing.
[performance fields; modeling]
Fernández A, Li HH & Carrasco M (2019).
How exogenous spatial attention affects visual representation.
Journal of Vision.
Orienting covert spatial attention to a target location enhances visual sensitivity and benefits performance in many visual tasks. How these attention-related improvements in performance affect the underlying visual representation of low-level visual features is not fully understood. Here we focus on characterizing how exogenous spatial attention affects the feature representations of orientation and spatial frequency. We asked observers to detect a vertical grating embedded in noise and performed psychophysical reverse correlation. Doing so allowed us to make comparisons with previous studies that utilized the same task and analysis to assess how endogenous attention and presaccadic modulations affect visual representations. We found that exogenous spatial attention improved performance and enhanced the gain of the target orientation without affecting orientation tuning width. Moreover, we found no change in spatial frequency tuning. We conclude that covert exogenous spatial attention alters performance by strictly boosting gain of orientation-selective filters, much like covert endogenous spatial attention.
[spatial attention; psychophysics, eye-tracking, reverse correlation]
Donovan I, Zhou YJ & Carrasco M (2019).
In search of exogenous feature-based attention.
Attention, Perception & Psychophysics.
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.
[exogenous attention, feature based attention]
Waite SA, Grigorian A, Alexander RG, Macknik SL, Carrasco M, Heeger D & Martinez-Conde S (2019).
Analysis of Perceptual Expertise in Radiology – Current Knowledge and a New Perspective.
Frontiers in Human Neuroscience, 13:213.
Radiologists rely principally on visual inspection to detect, describe, and classify findings in medical images. As most interpretive errors in radiology are perceptual in nature, understanding the path to radiologic expertise during image analysis is essential to educate future generations of radiologists. We review the perceptual tasks and challenges in radiologic diagnosis, discuss models of radiologic image perception, consider the application of perceptual learning methods in medical training, and suggest a new approach to understanding perceptional expertise. Specific principled enhancements to educational practices in radiology promise to deepen perceptual expertise among radiologists with the goal of improving training and reducing medical error.
[attention, eye movements, clinical, radiology]
Li HH, Pan J & Carrasco M (2019).
Presaccadic attention improves or impairs performance by enhancing sensitivity to higher spatial frequencies.
Scientific Reports, 9:2659.
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.
[presaccadic attention]
Vetter P*, Badde S*, Phelps EA & Carrasco M (2019).
Emotional faces guide the eyes in the absence of awareness.
eLife.
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.
[eye movements, emotion]
Cavanaugh MR, Barbot A, Carrasco M* & Huxlin KR* (2019).
Feature-based attention potentiates recovery of fine direction discrimination in cortically blind patients.
Neuropsychologia.
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.
[feature based attention, cortical blindness]
Carrasco M & Barbot A (2019).
Spatial attention alters visual appearance.
Current Opinion in Psychology.
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.
Michel M, Beck D, Block N, Blumenfeld H, Brown R, Carmel D, Carrasco M, Chirimuuta M, Chun M, Cleeremans A, Dehaene S, Fleming SM, Frith C, Haggard P, He B, Heyes C, Goodale MA, Irvine L, Kawato M, Kentridge R, King J-R, Knight RT, Kouider S, Lamme V, Lamy D, Lau H, Laureys S, LeDoux J, Lin Y-T, Liu K, Macknik SL, Martinez-Conde S, Mashour GA, Melloni L, Miracchi L, Mylopoulos M, Naccache L, Owen AM, Passingham RE, Pessoa L, Peters MAK, Rahnev D, Ro T, Rosenthal D, Sasaki Y, Sergent C, Solovey G, Schiff ND, Seth A, Tallon-Baudry C, Tamietto M, Tong F, van Gaal S, Vlassova A, Watanabe T, Weisberg J, Yan K, Yoshida M (2019).
Opportunities and challenges for a maturing science of consciousness.
Nature Human Behavior.
Carrasco M (2019).
Attention alters appearance.
In Blockheads!: Essays on Ned Block's Philosophy of Mind and Consciousness. A. Pautz and D Stoljar (Eds.) Cambridge, MA: MIT Press.
Amit R, Abeles D, Carrasco M & Yuval-Greenberg S (2019).
Oculomotor inhibition reflects temporal expectations.
Neuroimage.
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.
Fernández A, Denison RN & Carrasco M (2019).
Temporal attention improves perception similarly at foveal and parafoveal locations.
Journal of Vision.
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.
Denison R, Yuval-Greenberg S & Carrasco M (2019).
Directing voluntary temporal attention increases fixational stability.
Journal of Neuroscience.
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.
2018
Pham A, Carrasco M & Kiorpes L (2018).
Endogenous attention improves perception in amblyopic macaques.
Journal of Vision.
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.
[spatial attention, special populations, amblyopia, animal study, motion discrimination, contrast]
Cutrone EK, Heeger DJ & Carrasco M (2018).
On spatial attention and its field size on the repulsion effect.
Journal of 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.
[spatial attention, special populations, amblyopia, animal study, motion discrimination, contrast]
Barbot A & Carrasco M (2018).
Emotion and anxiety potentiate the way attention alters visual appearance.
Scientific Reports.
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.
[spatial attention, emotion, apperance, contrast]
Jigo M & Carrasco M (2018).
Attention alters spatial resolution by modulating second-order processing.
Journal of Vision.
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.
[spatial attention, texture segmentation, spatial resolution]
Hilo-Merkovich R, Carrasco M & Yuval-Greenberg S (2018).
Task performance in covert, but not overt, attention correlates with early laterality of visual evoked potentials.
Neuropsychologia.
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.
[spatial attention]
2013
Anton-Erxleben K & Carrasco M (2013).
Attentional enhancement of spatial resolution: Linking behavioral and neurophysiological evidence.
Nature Reviews Neuroscience.
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.
Ferneyhough E, Kim MK, Phelps EA & Carrasco M (2013).
Anxiety modulates the effects of motion and attention on early vision.
Cognition & Emotion.
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.
Grubb MA, Behrmann M, Egan R, Minshew NJ, Carrasco M & Heeger DJ (2013).
Endogenous spatial attention: Evidence for intact functioning in adults with autism.
Autism Research.
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.
White AL, Rolfs M & Carrasco M (2013).
Adaptive deployment of spatial and feature-based attention before saccades.
Vision Research.
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.
Carrasco M, Eckstein M, Krauzlis R & Verghese (2013).
Attentional modulation: Target selection, active search and cognitive processing.
Vision Research.
Rolfs M, Lawrence BM & Carrasco M (2013).
Reach preparation enhances visual performance and appearance.
Philosophical Transactions of the Royal Society B.
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.
Pomplun M, Garaas TW & Carrasco M (2013).
The effects of task difficulty on visual search strategy in virtual 3D displays.
Journal of Vision.
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 xy 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.
White AL, Lunau R & Carrasco M (2013).
The attentional effects of single cues and color singletons on visual sensitivity.
Journal of Experimental Psychology: Human Perception and Performance.
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.
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