Task-irrelevant Locations Research Articles

Entorhinal cortex represents task-relevant remote locations independent of CA1.

Neurons can collectively represent the current sensory experience while an animal is exploring its environment or remote experiences while the animal is immobile. These remote representations can reflect learned associations and be required for learning. Neurons in the medial entorhinal cortex (MEC) reflect the animal's current location during movement, but little is known about what MEC neurons collectively represent during immobility. Here, we recorded thousands of neurons in superficial MEC and dorsal CA1 as mice learned to associate two pairs of rewarded locations. We found that during immobility, the MEC neural population frequently represented positions far from the animal's location, which we defined as 'non-local coding'. Cells with spatial firing fields at remote locations drove non-local coding, even as cells representing the current position remained active. While MEC non-local coding has been reported during sharp-wave ripples in downstream CA1, we observed non-local coding more often outside of ripples. In fact, CA1 activity was less coordinated with MEC during non-local coding. We further observed that non-local coding was pertinent to the task, as MEC preferentially represented remote task-relevant locations at appropriate times, while rarely representing task-irrelevant locations. Together, this work raises the possibility that MEC non-local coding could strengthen associations between locations independently from CA1.

A computational account of transsaccadic attentional allocation based on visual gain fields

Coordination of goal-directed behavior depends on the brain's ability to recover the locations of relevant objects in the world. In humans, the visual system encodes the spatial organization of sensory inputs, but neurons in early visual areas map objects according to their retinal positions, rather than where they are in the world. How the brain computes world-referenced spatial information across eye movements has been widely researched and debated. Here, we tested whether shifts of covert attention are sufficiently precise in space and time to track an object's real-world location across eye movements. We found that observers' attentional selectivity is remarkably precise and is barely perturbed by the execution of saccades. Inspired by recent neurophysiological discoveries, we developed an observer model that rapidly estimates the real-world locations of objects and allocates attention within this reference frame. The model recapitulates the human data and provides a parsimonious explanation for previously reported phenomena in which observers allocate attention to task-irrelevant locations across eye movements. Our findings reveal that visual attention operates in real-world coordinates, which can be computed rapidly at the earliest stages of cortical processing.

Eye-size effects in the dot-probe task: Greater sclera exposure predicts delayed disengagement from fearful faces.

Fearful facial expressions are nonverbal and biologically salient signals of potential threat that automatically hold, capture, and direct observers' attention. They are characterized by enlarged eye whites and dilated pupils, and fearful eyes alone are sufficient to capture attention. The morphological properties of the eye region, such as sclera exposure, are thought to play an important role in nonverbal communication. Specifically, increased sclera exposure associated with fearful expressions has been shown to moderate how observers' shift their attention toward the direction of another's gaze. Yet, the extent to which variability in sclera exposure possibly impacts the capture and hold of attention by fearful faces is untested. To address this, a sample of 249 adults completed a dot-probe task of selective attention with fearful and neutral faces. The results suggested that (1) fearful faces were prioritized over neutral faces (i.e., they captured and held attention), (2) greater sclera exposure at target locations facilitated reaction times, and (3) attention was held by greater sclera exposure of fearful faces at task irrelevant locations resulting in delayed disengagement. Collectively, the results indicate that fearful facial expressions and sclera exposure modulate spatial attention through independent and interactive mechanisms. Sclera exposure appears to be an important facilitator of nonverbal communication and perhaps represents an understudied variable in social cognition more broadly.

Rewarded Stimuli Do Not Capture Attention at Task-Irrelevant Locations

Rewarded Stimuli Do Not Capture Attention at Task-Irrelevant Locations

Spatial task relevance modulates value-driven attentional capture.

Attention tends to be attracted to visual features previously associated with reward. To date, nearly all existing studies examined value-associated stimuli at or near potential target locations, making such locations meaningful to inspect. The present experiments examined whether the attentional priority of a value-associated stimulus depends on its location-wise task relevance. In three experiments we used an RSVP task to compare the attentional demands of a value-associated peripheral distractor to that of a distractor associated with the top-down search goal. At a peripheral location that could never contain the target, a value-associated color did not capture attention. In contrast, at the same location, a distractor in a goal-matching color did capture attention. The results show that value-associated stimuli lose their attentional priority at task-irrelevant locations, in contrast to other types of stimuli that capture attention.

Not looking for any trouble? Purely affective attentional settings do not induce goal-driven attentional capture.

It is well established that directing goal-driven attention to a particular stimulus property (e.g., red), or a conceptual category (e.g., toys) can induce powerful involuntary capture by goal-matching stimuli. Here, we tested whether broad affective search goals (e.g., for anything threat-related) could similarly induce a generalized capture to an entire matching affective category. Across four experiments, participants were instructed to search for threat-related images in a Rapid Serial Visual Presentation (RSVP) stream, while ignoring threat-related distractors presented in task-irrelevant locations. Across these experiments we found no evidence of goal-driven attentional capture by threat distractors when participants adopted a general 'threat detection' goal encompassing multiple subcategories of threat (Experiments 1a, 1b). This was true even when there was partial overlap between the threat distractors and the search goal (i.e., subset of the targets matched the distractor; Experiment 2). However, when participants adopted a more specific goal for a single subcategory of threat (e.g., fearful faces), robust goal-driven capture occurred by distractors matching this subcategory (Experiment 3). These findings suggest that while affective criteria can be used in the guidance of attention, attentional settings based on affective properties alone may not induce goal-driven attentional capture. We discuss implications for recent goal-driven accounts of affective attentional biases.

Tuning Attention to Object Categories: Spatially Global Effects of Attention to Faces in Visual Processing.

Feature-based attention is known to enhance visual processing globally across the visual field, even at task-irrelevant locations. Here, we asked whether attention to object categories, in particular faces, shows similar location-independent tuning. Using EEG, we measured the face-selective N170 component of the EEG signal to examine neural responses to faces at task-irrelevant locations while participants attended to faces at another task-relevant location. Across two experiments, we found that visual processing of faces was amplified at task-irrelevant locations when participants attended to faces relative to when participants attended to either buildings or scrambled face parts. The fact that we see this enhancement with the N170 suggests that these attentional effects occur at the earliest stage of face processing. Two additional behavioral experiments showed that it is easier to attend to the same object category across the visual field relative to two distinct categories, consistent with object-based attention spreading globally. Together, these results suggest that attention to high-level object categories shows similar spatially global effects on visual processing as attention to simple, individual, low-level features.

Cognitive development attenuates audiovisual distraction and promotes the selection of task-relevant perceptual saliency during visual search on complex scenes

Searching for a target while avoiding distraction is a core function of selective attention involving both voluntary and reflexive mechanisms. Here, for the first time, we investigated the development of the interplay between voluntary and reflexive mechanisms of selective attention from childhood to early adulthood. We asked 6-, 10-, and 20-year-old participants to search for a target presented in one hemifield of a complex scene, preceded by a task-irrelevant auditory cue on either the target side (valid), the opposite side (invalid), or both sides (neutral). For each scene we computed the number of salient locations (NSL) and the target saliency (TgS). All age groups showed comparable orienting effects (“valid minus neutral” trials), indicating a similar capture of spatial attention by valid cues which was independent of age. However, only adults demonstrated a suppression of the reorienting effect (“invalid minus neutral” trials), indicating late developments in the reallocation of spatial attention toward a target following auditory distraction. The searching performance of the children (both 6- and 10-year-olds), but not of the adults, was predicted by the NSL, indicating an attraction of processing resources to salient but task-irrelevant locations in childhood; conversely, only adults showed greater performance with increased TgS in valid trials, indicating late development in the use of task-related saliency. These findings highlight qualitatively different mechanisms of selective attention operating at different ages, demonstrating important developmental changes in the interplay between voluntary and reflexive mechanisms of selective attention during visual search in complex scenes.

It takes two to tango: Suppression of task-irrelevant features requires (spatial) competition

In a recent electrophysiological study, we reported on global facilitation but local suppression of color stimuli in feature-based attention in human early visual cortex. Subjects attended to one of two centrally located superimposed red/blue random dot kinematograms (RDKs). Task-irrelevant single RDKs in the same colors were presented in the left and right periphery, respectively. Suppression of the to-be-ignored color was only present in the centrally located RDK but not in the one with the same color in the periphery. This result was at odds with the idea of active suppression of task-irrelevant features across the entire visual field. In the present study, we introduced competition in the periphery by superimposing the RDKs at the task-irrelevant location as well. With such competition, we found suppression of the task-irrelevant color in the centrally and peripherally located RDKs. Results clearly demonstrate that suppression of task-irrelevant features at task-irrelevant locations requires (spatial) competitive interactions and is not an inherent neural mechanism in feature-based attention as was found for global facilitation.

Feature-guided attentional capture cannot be prevented by spatial filtering

Feature-based control processes guide attention towards objects with target features in visual search. While these processes are assumed to operate globally across the entire visual field, it remains controversial whether target-matching objects at task-irrelevant locations can be excluded from attentional selection, especially when spatial attention is already narrowly focused elsewhere. We investigated whether probe stimuli at irrelevant lateral locations capture attention when they precede search displays where targets are defined either by a specific feature (colour or orientation) or by a colour/orientation conjunction by measuring N2pc components (an electrophysiological marker of attentional target selection) to these probes. Reliable N2pcs were triggered by probes not only in the feature search tasks but also when participants searched for feature conjunctions, in spite of the fact that conjunction search requires focal spatial attention. Analogous N2pc results were found in the absence of any spatial uncertainty about the location of conjunctively defined targets, which always appeared at fixation. These results show that rapid attentional capture by objects with target-matching features cannot be prevented by top-down spatial filtering mechanisms, and confirm that feature-based attentional guidance processes operate in a spatially global fashion.

Hierarchies of Attention and Experimental Designs: Effects of Spatial and Intermodal Attention Revisited.

When attention is directed to stimuli in a given modality and location, information processing in other irrelevant modalities at this location is affected too. This spread of attention to irrelevant stimuli is often interpreted as superiority of location selection over modality selection. However, this conclusion is based on experimental paradigms in which spatial attention was transient whereas intermodal attention was sustained. Furthermore, whether modality selection affects processing in the task-relevant modality at irrelevant locations remains an open question. Here, we addressed effects of simultaneous spatial and intermodal attention in an EEG study using a balanced design where spatial attention was transient and intermodal attention sustained or vice versa. Effects of spatial attention were not affected by which modality was attended and effects of intermodal attention were not affected by whether the stimuli were at the attended location or not. This suggests not only spread of spatial attention to task-irrelevant modalities but also spread of intermodal attention to task-irrelevant locations. Whether spatial attention was transient or sustained did not alter the effect of spatial attention on visual N1 and Nd1 responses. Prestimulus preparatory occipital alpha band responses were affected by both transient and sustained spatial cueing, whereas late post-stimulus responses were more strongly affected by sustained than by transient spatial attention. Sustained but not transient intermodal attention affected late responses (>200 msec) to visual stimuli. Together, the results undermine the universal superiority of spatial attention and suggest that the mode of attention manipulation is an important factor determining attention effects.

Interactions between space-based and feature-based attention.

Although early research suggested that attention to nonspatial features (i.e., red) was confined to stimuli appearing at an attended spatial location, more recent research has emphasized the global nature of feature-based attention. For example, a distractor sharing a target feature may capture attention even if it occurs at a task-irrelevant location. Such findings have been used to argue that feature-based attention operates independently of spatial attention. However, feature-based attention may nonetheless interact with spatial attention, yielding larger feature-based effects at attended locations than at unattended locations. The present study tested this possibility. In 2 experiments, participants viewed a rapid serial visual presentation (RSVP) stream and identified a target letter defined by its color. Target-colored distractors were presented at various task-irrelevant locations during the RSVP stream. We found that feature-driven attentional capture effects were largest when the target-colored distractor was closer to the attended location. These results demonstrate that spatial attention modulates the strength of feature-based attention capture, calling into question the prior evidence that feature-based attention operates in a global manner that is independent of spatial attention.

The visual P2 is attenuated for attended objects near the hands

Vision is altered when people place their hands near the object they are observing. To investigate the neural processes underlying this effect, we measured electroencephalographic visual-evoked potentials (VEPs) elicited by reversing checkerboards, while participants' hands either surrounded the visual display or rested at their sides. We found the P2 component was attenuated for hand-proximal stimuli, but only when participants attended to the location of the checkerboard. In Experiment 1, participants performed an attention-demanding color-change task that was presented centrally, and the P2 component was attenuated for central, but not peripheral, checkerboards. In Experiment 2, participants performed the attention task in the periphery, and the P2 was attenuated for peripheral, but not central, checkerboards. These results suggest that hand-proximal stimuli benefit from enhanced selective attention at later stages of perceptual processing. The effect only occurs for objects at task-relevant locations, however, even when task-irrelevant locations are physically closer to the hands.

Feature-based enhancement of visual stimuli at task-irrelevant locations

Feature-based enhancement of visual stimuli at task-irrelevant locations

Perceptual load modulates the processing of distractors presented at task-irrelevant locations during the attentional blink

The distribution of attention in both space and time is critical for processing our dynamic environment. Studies of spatial attention suggest that the distribution of attention is decreased when the perceptual load of a task increases, resulting in decreased processing of task-irrelevant distractors. Studies of the attentional blink (AB) suggest that the temporal distribution of attention also influences distractor processing, such that distractor processing increases during the AB relative to outside the AB (Jiang & Chun, 2001). Two experiments are reported in which the extent to which the difficulty of the first target task (T1) modulates the processing of task-irrelevant distractors during the AB was tested. To investigate this issue, both the first and second target tasks (T1 and T2) required identifying a central stimulus that was flanked by low-load or high-load distractors. Consistent with previous studies of the AB, there was evidence of more distractor processing during the AB than outside the AB. Critically, however, the interference caused by distractors presented simultaneously with T2 during the AB was reduced when T1 perceptual load was high relative to when it was low. These results suggest that increasing T1 perceptual load decreases distractor processing during the AB and that perceptual processes influence both the temporal and spatial distribution of attention.

Perceptual load modulates the processing of distractors presented at task-irrelevant locations during the attentional blink

The distribution of attention in both space and time is critical for processing our dynamic environment. Studies of spatial attention suggest that the distribution of attention is decreased when the perceptual load of a task increases, resulting in decreased processing of task-irrelevant distractors. Studies of the attentional blink (AB) suggest that the temporal distribution of attention also influences distractor processing, such that distractor processing increases during the AB relative to outside the AB (Jiang & Chun, 2001). Two experiments are reported in which the extent to which the difficulty of the first target task (T1) modulates the processing of task-irrelevant distractors during the AB was tested. To investigate this issue, both the first and second target tasks (T1 and T2) required identifying a central stimulus that was flanked by low-load or high-load distractors. Consistent with previous studies of the AB, there was evidence of more distractor processing during the AB than outside the AB. Critically, however, the interference caused by distractors presented simultaneously with T2 during the AB was reduced when T1 perceptual load was high relative to when it was low. These results suggest that increasing T1 perceptual load decreases distractor processing during the AB and that perceptual processes influence both the temporal and spatial distribution of attention.

Attention cannot spare task-irrelevant locations on an attended object

Attention cannot spare task-irrelevant locations on an attended object

Temporal precedence of emotion over attention modulations in the lateral amygdala: Intracranial ERP evidence from a patient with temporal lobe epilepsy

Previous fMRI studies have reported mixed evidence for the influence of selective attention on amygdala responses to emotional stimuli, with some studies showing "automatic" emotional effects to threat-related stimuli without attention (or even without awareness), but other studies showing a gating of amygdala activity by selective attention with no response to unattended stimuli. We recorded intracranial local field potentials from the intact left lateral amygdala in a human patient prior to surgery for epilepsy and tested, with a millisecond time resolution, for neural responses to fearful faces appearing at either task-relevant or task-irrelevant locations. Our results revealed an early emotional effect in the amygdala arising prior to, and independently of, attentional modulation. However, at a later latency, we found a significant modulation of the differential emotional response when attention was directed toward or away from fearful faces. These results suggest separate influences of emotion and attention on amygdala activation and may help reconcile previous discrepancies concerning the relative responsiveness of the human amygdala to emotional and attentional factors.

Exploring the modulation of attentional capture by attentional control settings using performance and illusory line motion

Modulation of attentional capture by attentional control settings was explored using performance and phenomenology. Trials began with four figure-8s presented above, below, left, and right of fixation. Any figure-8 (or none) brightened uninformatively (cue) before presentation of either a digit target (2 or 5 made from a figure-8 by deletion) calling for a speeded identification, or a line connecting adjacent figure-8s calling for a motion judgement. Such lines are seen drawn away from an adjacent cue. Digit targets appeared only on the horizontal or vertical axes, encouraging voluntary attention to two (target-relevant) of the four figure-8s. Whereas voluntary attention reduced attentional capture from cues at taskirrelevant locations (when lines were unlikely to be presented near the task-irrelevant locations), it had no effect on motion judgements, suggesting that cue-elicited exogenous attention affects perceptual arrival times while voluntary attention prioritizes locations for further processing without affecting arrival times.

Cholinergic enhancement modulates neural correlates of selective attention and emotional processing

Neocortical cholinergic afferents are proposed to influence both selective attention and emotional processing. In a study of healthy adults we used event-related fMRI while orthogonally manipulating attention and emotionality to examine regions showing effects of cholinergic modulation by the anticholinesterase physostigmine. Either face or house pictures appeared at task-relevant locations, with the alternative picture type at irrelevant locations. Faces had either neutral or fearful expressions. Physostigmine increased relative activity within the anterior fusiform gyrus for faces at attended, versus unattended, locations, but decreased relative activity within the posterolateral occipital cortex for houses in attended, versus unattended, locations. A similar pattern of regional differences in the effect of physostigmine on cue-evoked responses was also present in the absence of stimuli. Cholinergic enhancement augmented the relative neuronal response within the middle fusiform gyrus to fearful faces, whether at attended or unattended locations. By contrast, physostigmine influenced responses in the orbitofrontal, intraparietal and cingulate cortices to fearful faces when faces occupied task-irrelevant locations. These findings suggest that acetylcholine may modulate both selective attention and emotional processes through independent, region-specific effects within the extrastriate cortex. Furthermore, cholinergic inputs to the frontoparietal cortex may influence the allocation of attention to emotional information.