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Jan Theeuwes
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Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2023) 35 (6): 1032–1044.
Published: 01 June 2023
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Although in many cases salient stimuli capture attention involuntarily, it has been proposed recently that under certain conditions, the bottom–up signal generated by such stimuli can be proactively suppressed. In support of this signal suppression hypothesis, ERP studies have demonstrated that salient stimuli that do not capture attention elicit a distractor positivity (P D ), a putative neural index of suppression. At the same time, it is becoming increasingly clear that regularities across preceding search episodes have a large influence on attentional selection. Yet to date, studies in support of the signal suppression hypothesis have largely ignored the role of selection history on the processing of distractors. The current study addressed this issue by examining how electrophysiological markers of attentional selection (N2pc) and suppression (P D ) elicited by targets and distractors, respectively, were modulated when the search target randomly varied instead of being fixed across trials. Results showed that although target selection was unaffected by this manipulation, both in terms of manual response times, as well as in terms of the N2pc component, the P D component was reliably attenuated when the target features varied randomly across trials. This result demonstrates that the distractor P D , which is typically considered the marker of selective distractor processing, cannot unequivocally be attributed to suppression only, as it also, at least in part, reflects the upweighting of target features.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2019) 31 (10): 1535–1548.
Published: 01 October 2019
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Salient yet irrelevant objects often capture our attention and interfere with our daily tasks. Distraction by salient objects can be reduced by suppressing the location where they are likely to appear. The question we addressed here was whether suppression of frequent distractor locations is already implemented beforehand, in anticipation of the stimulus. Using EEG, we recorded cortical activity of human participants searching for a target while ignoring a salient distractor. The distractor was presented more often at one location than at any other location. We found reduced capture for distractors at frequent locations, indicating that participants learned to avoid distraction. Critically, we found evidence for “proactive suppression” as already “prior to display onset,” there was enhanced power in parieto-occipital alpha oscillations contralateral to the frequent distractor location—a signal known to occur in anticipation of irrelevant information. Locked to display onset, ERP analysis showed a distractor suppression-related distractor positivity (P D ) component for this location. Importantly, this P D was found regardless of whether distracting information was presented at the frequent location. In addition, there was an early P D component representing an early attentional index of the frequent distractor location. Our results show anticipatory (proactive) suppression of frequent distractor locations in visual search already starting prior to display onset.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2018) 30 (2): 256–266.
Published: 01 February 2018
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Current theories assume a functional role for covert attention in the maintenance of spatial information in working memory. Consistent with this view, both the locus of attention and positions stored in working memory can be decoded based on the topography of oscillatory alpha-band (8–12 Hz) activity on the scalp. Thus far, however, alpha modulation has been studied in isolation for covert attention and working memory tasks. Here, we applied an inverted spatial encoding model in combination with EEG to study the temporal dynamics of spatially specific alpha activity during a task that required observers to visually select a target location while maintaining another independently varying location in working memory. During the memory delay period, alpha-based spatial tuning functions shifted from the position stored in working memory to the covertly attended position and back again after the attention task was completed. The findings provide further evidence for a common oscillatory mechanism in both the selection and the maintenance of relevant spatial visual information and demonstrate the dynamic trade-off in prioritization between two spatial tasks.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2015) 27 (6): 1215–1237.
Published: 01 June 2015
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It is solidly established that top–down (goal-driven) and bottom–up (stimulus-driven) attention mechanisms depend on distributed cortical networks, including prefrontal and frontoparietal regions. On the other hand, it is less clear whether the BG also contribute to one or the other of these mechanisms, or to both. The current study was principally undertaken to clarify this issue. Parkinson disease (PD), a neurodegenerative disorder primarily affecting the BG, has proven to be an effective model for investigating the contribution of the BG to different brain functions; therefore, we set out to investigate deficits of top–down and bottom–up attention in a selected cohort of PD patients. With this objective in mind, we compared the performance on three computerized tasks of two groups of 12 parkinsonian patients (assessed without any treatment), one otherwise pharmacologically treated and the other also surgically treated, with that of a group of controls. The main behavioral tool for our study was an attentional capture task, which enabled us to tap the competition between top–down and bottom–up mechanisms of visual attention. This task was suitably combined with a choice RT and a simple RT task to isolate any specific deficit of attention from deficits in motor response selection and initiation. In the two groups of patients, we found an equivalent increase of attentional capture but also comparable delays in target selection in the absence of any salient distractor (reflecting impaired top–down mechanisms) and movement initiation compared with controls. In contrast, motor response selection processes appeared to be prolonged only in the operated patients. Our results confirm that the BG are involved in both motor and cognitive domains. Specifically, damage to the BG, as it occurs in PD, leads to a distinct deficit of top–down control of visual attention, and this can account, albeit indirectly, for the enhancement of attentional capture, reflecting weakened ability of top–down mechanisms to antagonize bottom–up control.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2013) 25 (10): 1754–1765.
Published: 01 October 2013
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The mechanisms that underlie the integration of visual and goal-related signals for the production of saccades remain poorly understood. Here, we examined how spatial proximity of competing stimuli shapes goal-directed responses in the superior colliculus (SC), a midbrain structure closely associated with the control of visual attention and eye movements. Monkeys were trained to perform an oculomotor-capture task [Theeuwes, J., Kramer, A. F., Hahn, S., Irwin, D. E., & Zelinsky, G. J. Influence of attentional capture on oculomotor control. Journal of Experimental Psychology. Human Perception and Performance, 25, 1595–1608, 1999], in which a target singleton was revealed via an isoluminant color change in all but one item. On a portion of the trials, an additional salient item abruptly appeared near or far from the target. We quantified how spatial proximity between the abrupt-onset and the target shaped the goal-directed response. We found that the appearance of an abrupt-onset near the target induced a transient decrease in goal-directed discharge of SC visuomotor neurons. Although this was indicative of spatial competition, it was immediately followed by a rebound in presaccadic activation, which facilitated the saccadic response (i.e., it induced shorter saccadic RT). A similar suppression also occurred at most nontarget locations even in the absence of the abrupt-onset. This is indicative of a mechanism that enabled monkeys to quickly discount stimuli that shared the common nontarget feature. These results reveal a pattern of excitation/inhibition across the SC visuomotor map that acted to facilitate optimal behavior—the short duration suppression minimized the probability of capture by salient distractors, whereas a subsequent boost in accumulation rate ensured a fast goal-directed response. Such nonlinear dynamics should be incorporated into future biologically plausible models of saccade behavior.
Journal Articles
A Retinotopic Attentional Trace after Saccadic Eye Movements: Evidence from Event-related Potentials
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2013) 25 (9): 1563–1577.
Published: 01 September 2013
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Saccadic eye movements are a major source of disruption to visual stability, yet we experience little of this disruption. We can keep track of the same object across multiple saccades. It is generally assumed that visual stability is due to the process of remapping, in which retinotopically organized maps are updated to compensate for the retinal shifts caused by eye movements. Recent behavioral and ERP evidence suggests that visual attention is also remapped, but that it may still leave a residual retinotopic trace immediately after a saccade. The current study was designed to further examine electrophysiological evidence for such a retinotopic trace by recording ERPs elicited by stimuli that were presented immediately after a saccade (80 msec SOA). Participants were required to maintain attention at a specific location (and to memorize this location) while making a saccadic eye movement. Immediately after the saccade, a visual stimulus was briefly presented at either the attended location (the same spatiotopic location), a location that matched the attended location retinotopically (the same retinotopic location), or one of two control locations. ERP data revealed an enhanced P1 amplitude for the stimulus presented at the retinotopically matched location, but a significant attenuation for probes presented at the original attended location. These results are consistent with the hypothesis that visuospatial attention lingers in retinotopic coordinates immediately following gaze shifts.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2012) 24 (3): 707–717.
Published: 01 March 2012
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During natural viewing, the trajectories of saccadic eye movements often deviate dramatically from a straight-line path between objects. In human studies, saccades have been shown to deviate toward or away from salient visual distractors depending on visual- and goal-related parameters, but the neurophysiological basis for this is not well understood. Some studies suggest that deviation toward is associated with competition between simultaneously active sites within the intermediate layers of the superior colliculus (SC), a midbrain structure that integrates sensory and goal-related signals for the production of saccades. In contrast, deviation away is hypothesized to reflect a higher-level process, whereby the neural site associated with the distractor isactively suppressed via a form of endogenous, top–down inhibition. We tested this hypothesis by measuring presaccadic distractor-evoked activation of SC visuomotor neurons while monkeys performed a simple task configured specifically to induce a high degree of saccades that deviate away. In the SC, cognitive processes such as top–down expectation are represented as variation in the sustained, low-frequency presaccadic discharge. We reasoned that any inhibition at the distractor-related locus associated with saccade deviation should affect the excitability of the neuron, thereby affecting the discharge rate. We found that, although the task produced robust deviation away, there was no evidence of a relationship between saccade deviation and distractor-evoked activation outside a short perisaccadic window that began no earlier than 22 msec before saccade onset. This indicates that deviation away is not adequately explained by a form of sustained, top–down inhibition at the distractor-related locus in the SC. The results are discussed in relation to the primary sources of inhibition associated with saccadic control.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2010) 22 (4): 761–774.
Published: 01 April 2010
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It is still debated to what degree top–down and bottom–up driven attentional control processes are subserved by shared or by separate mechanisms. Interactions between these attentional control forms were investigated using a rapid event-related fMRI design, using an attentional search task. Following a prestimulus mask, target stimuli (consisting of a letter C or a mirror image of the C, enclosed in a diamond outline) were presented either at one unique location among three nontarget items (consisting of a random letter, enclosed in a circle outline; 50% probability), or at all four possible target locations (also 50% probability). On half the trials, irrelevant color singletons were presented, consisting of a color change of one of the four prestimulus masks, just prior to target appearance. Participants were required to search for a target letter inside the diamond and report its orientation. Results indicate that, in addition to a common network of parietal areas, medial frontal cortex is uniquely involved in top–down orienting, whereas bottom–up control is mainly subserved by a network of occipital and parietal areas. Additionally, we found that participants who were better able to suppress orienting to the color singleton showed middle frontal gyrus activation, and that the degree of top–down control correlated with insular activity. We conclude that, in addition to a common set of parietal areas, separate brain areas are involved in top–down and bottom–up driven attentional control, and that frontal areas play a role in the suppression of attentional capture by an irrelevant color singleton.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2008) 20 (12): 2285–2297.
Published: 01 December 2008
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Previous research has shown that task-irrelevant onsets trigger an eye movement in their direction. Such oculomotor capture is often impervious to conscious awareness. The present study used event-related brain potentials to examine how such oculomotor errors are detected, evaluated, and compensated for and whether awareness of an error played a role at any of these stages of processing. The results show that the early processes of error detection and correction (as represented by the error-related negativity and the parietal N1) are not directly affected by subjective awareness of making an error. Instead, they seem to be modulated by the degree of temporal overlap between the programming of the correct and erroneous saccade. We found that only a later component (the error-related positivity [Pe]) is modulated by awareness of making an erroneous eye movement. We propose that awareness of oculomotor capture primarily depends on this later process.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2008) 20 (11): 2025–2036.
Published: 01 November 2008
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There is evidence that some visual information in blind regions may still be processed in patients with hemifield defects after cerebral lesions (“blindsight”). We tested the hypothesis that, in the absence of retinogeniculostriate processing, residual retinotectal processing may still be detected as modifications of saccades to seen targets by irrelevant distractors in the blind hemifield. Six patients were presented with distractors in the blind and intact portions of their visual field and participants were instructed to make eye movements to targets in the intact field. Eye movements were recorded to determine if blind-field distractors caused deviation in saccadic trajectories. No deviation was found in one patient with an optic chiasm lesion, which affect both retinotectal and retinogeniculostriate pathways. In five patients with lesions of the optic radiations or the striate cortex, the results were mixed, with two of the five patients showing significant deviations of saccadic trajectory away from the “blind” distractor. In a second experiment, two of the five patients were tested with the target and the distractor more closely aligned. Both patients showed a “global effect,” in that saccades deviated toward the distractor, but the effect was stronger in the patient who also showed significant trajectory deviation in the first experiment. Although our study confirms that distractor effects on saccadic trajectory can occur in patients with damage to the retinogeniculostriate visual pathway but preserved retinotectal projections, there remain questions regarding what additional factors are required for these effects to manifest themselves in a given patient.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2006) 18 (4): 604–613.
Published: 01 April 2006
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We investigated the ability of salient yet task-irrelevant stimuli to capture attention in two visual search experiments. Participants were presented with circular search arrays that contained a highly salient distractor singleton defined by color and a less salient target singleton defined by form. A component of the event-related potential called the N2pc was used to track the allocation of attention to lateralized positions in the arrays. In Experiment 1, a lateralized distractor elicited an N2pc when a concurrent target was presented on the vertical meridian and thus could not elicit lateralized components such as the N2pc. A similar distractor-elicited N2pc was found in Experiment 2, which was conducted to rule out certain voluntary search strategies. Additionally, in Experiment 2 both the distractor and the target elicited the N2pc component when the two stimuli were presented on opposite sides of the search array. Critically, the distractor-elicited N2pc preceded the target-elicited N2pc on these trials. These results demonstrate that participants shifted attention to the target only after shifting attention to the more salient but task-irrelevant distractor. This pattern of results is in line with theories of attention in which stimulus-driven control plays an integral role.