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1.  Attention for speaking: domain-general control from the anterior cingulate cortex in spoken word production 
Accumulating evidence suggests that some degree of attentional control is required to regulate and monitor processes underlying speaking. Although progress has been made in delineating the neural substrates of the core language processes involved in speaking, substrates associated with regulatory and monitoring processes have remained relatively underspecified. We report the results of an fMRI study examining the neural substrates related to performance in three attention-demanding tasks varying in the amount of linguistic processing: vocal picture naming while ignoring distractors (picture-word interference, PWI); vocal color naming while ignoring distractors (Stroop); and manual object discrimination while ignoring spatial position (Simon task). All three tasks had congruent and incongruent stimuli, while PWI and Stroop also had neutral stimuli. Analyses focusing on common activation across tasks identified a portion of the dorsal anterior cingulate cortex (ACC) that was active in incongruent trials for all three tasks, suggesting that this region subserves a domain-general attentional control function. In the language tasks, this area showed increased activity for incongruent relative to congruent stimuli, consistent with the involvement of domain-general mechanisms of attentional control in word production. The two language tasks also showed activity in anterior-superior temporal gyrus (STG). Activity increased for neutral PWI stimuli (picture and word did not share the same semantic category) relative to incongruent (categorically related) and congruent stimuli. This finding is consistent with the involvement of language-specific areas in word production, possibly related to retrieval of lexical-semantic information from memory. The current results thus suggest that in addition to engaging language-specific areas for core linguistic processes, speaking also engages the ACC, a region that is likely implementing domain-general attentional control.
PMCID: PMC3856851  PMID: 24368899
attentional control; anterior cingulate cortex; superior temporal cortex; picture-word interference; Simon; Stroop; word production
2.  Neural Correlates of Individual Performance Differences in Resolving Perceptual Conflict 
PLoS ONE  2012;7(8):e42849.
Attentional mechanisms are a crucial prerequisite to organize behavior. Most situations may be characterized by a ‘competition’ between salient, but irrelevant stimuli and less salient, relevant stimuli. In such situations top-down and bottom-up mechanisms interact with each other. In the present fMRI study, we examined how interindividual differences in resolving situations of perceptual conflict are reflected in brain networks mediating attentional selection. Doing so, we employed a change detection task in which subjects had to detect luminance changes in the presence and absence of competing distractors. The results show that good performers presented increased activation in the orbitofrontal cortex (BA 11), anterior cingulate (BA 25), inferior parietal lobule (BA 40) and visual areas V2 and V3 but decreased activation in BA 39. This suggests that areas mediating top-down attentional control are stronger activated in this group. Increased activity in visual areas reflects distinct neuronal enhancement relating to selective attentional mechanisms in order to solve the perceptual conflict. Opposed to good performers, brain areas activated by poor performers comprised the left inferior parietal lobule (BA 39) and fronto-parietal and visual regions were continuously deactivated, suggesting that poor performers perceive stronger conflict than good performers. Moreover, the suppression of neural activation in visual areas might indicate a strategy of poor performers to inhibit the processing of the irrelevant non-target feature. These results indicate that high sensitivity in perceptual areas and increased attentional control led to less conflict in stimulus processing and consequently to higher performance in competitive attentional selection.
PMCID: PMC3423441  PMID: 22916169
3.  Effects of Working Memory Load on Visual Selective Attention: Behavioral and Electrophysiological Evidence 
Working memory and attention interact in a way that enables us to focus on relevant items and maintain current goals. The influence of working memory on attention has been noted in several studies using dual task designs. Multitasking increases the demands on working memory and reduces the amount of resources available for cognitive control functions such as resolving stimulus conflict. However, few studies have investigated the temporal activation of the cortex while multitasking. The present study addresses the extent to which working memory load influences early (P1) and late (P300) attention-sensitive event-related potential components using a dual task paradigm. Participants performed an arrow flanker task alone (single task condition) or concurrently with a Sternberg memory task (dual task condition). In the flanker task, participants responded to the direction of a central arrow surrounded by congruent or incongruent arrows. In the dual task condition, participants were presented with a Sternberg task that consisted of either four or seven consonants to remember prior to a short block of flanker trials. Participants were slower and less accurate on incongruent versus congruent trials. Furthermore, accuracy on incongruent trials was reduced in both dual task conditions. Likewise, P300 amplitude to incongruent flanker stimuli decreased when working memory load increased. These findings suggest that interference from incongruent flankers was more difficult to suppress when working memory was taxed. In addition, P1 amplitude was diminished on all flanker trials in the dual task condition. This result indicates that top-down attentional control over early visual processing is diminished by increasing demands on working memory. Both the behavioral and electrophysiological results suggest that working memory is critical in maintaining attentional focus and resolving conflict.
PMCID: PMC3115462  PMID: 21716633
dual task; working memory; ERPs; attention; P1; P300
4.  The Role of Stimulus Salience and Attentional Capture Across the Neural Hierarchy in a Stop-Signal Task 
PLoS ONE  2011;6(10):e26386.
Inhibitory motor control is a core function of cognitive control. Evidence from diverse experimental approaches has linked this function to a mostly right-lateralized network of cortical and subcortical areas, wherein a signal from the frontal cortex to the basal ganglia is believed to trigger motor-response cancellation. Recently, however, it has been recognized that in the context of typical motor-control paradigms those processes related to actual response inhibition and those related to the attentional processing of the relevant stimuli are highly interrelated and thus difficult to distinguish. Here, we used fMRI and a modified Stop-signal task to specifically examine the role of perceptual and attentional processes triggered by the different stimuli in such tasks, thus seeking to further distinguish other cognitive processes that may precede or otherwise accompany the implementation of response inhibition. In order to establish which brain areas respond to sensory stimulation differences by rare Stop-stimuli, as well as to the associated attentional capture that these may trigger irrespective of their task-relevance, we compared brain activity evoked by Stop-trials to that evoked by Go-trials in task blocks where Stop-stimuli were to be ignored. In addition, region-of-interest analyses comparing the responses to these task-irrelevant Stop-trials, with those to typical relevant Stop-trials, identified separable activity profiles as a function of the task-relevance of the Stop-signal. While occipital areas were mostly blind to the task-relevance of Stop-stimuli, activity in temporo-parietal areas dissociated between task-irrelevant and task-relevant ones. Activity profiles in frontal areas, in turn, were activated mainly by task-relevant Stop-trials, presumably reflecting a combination of triggered top-down attentional influences and inhibitory motor-control processes.
PMCID: PMC3195690  PMID: 22022611
5.  The effect of non-visual working memory load on top-down modulation of visual processing 
Neuropsychologia  2009;47(7):1637-1646.
While a core function of the working memory (WM) system is the active maintenance of behaviorally relevant sensory representations, it is also critical that distracting stimuli are appropriately ignored. We used functional magnetic resonance imaging to examine the role of domain-general WM resources in the top-down attentional modulation of task-relevant and irrelevant visual representations. In our dual-task paradigm, each trial began with the auditory presentation of six random (high load) or sequentially-ordered (low load) digits. Next, two relevant visual stimuli (e.g., faces), presented amongst two temporally interspersed visual distractors (e.g., scenes), were to be encoded and maintained across a 7-sec delay interval, after which memory for the relevant images and digits was probed. When taxed by high load digit maintenance, participants exhibited impaired performance on the visual WM task and a selective failure to attenuate the neural processing of task-irrelevant scene stimuli. The over-processing of distractor scenes under high load was indexed by elevated encoding activity in a scene-selective region-of-interest relative to low load and passive viewing control conditions, as well as by improved long-term recognition memory for these items. In contrast, the load manipulation did not affect participants' ability to upregulate activity in this region when scenes were task-relevant. These results highlight the critical role of domain-general WM resources in the goal-directed regulation of distractor processing. Moreover, the consequences of increased WM load in young adults closely resemble the effects of cognitive aging on distractor filtering [Gazzaley et al., (2005) Nature Neuroscience 8, 1298-1300], suggesting the possibility of a common underlying mechanism.
PMCID: PMC2701233  PMID: 19397858
selective attention; distraction; suppression; filtering; dual task; fMRI
6.  Modulations of the executive control network by stimulus onset asynchrony in a Stroop task 
BMC Neuroscience  2013;14:79.
Manipulating task difficulty is a useful way of elucidating the functional recruitment of the brain’s executive control network. In a Stroop task, pre-exposing the irrelevant word using varying stimulus onset asynchronies (‘negative’ SOAs) modulates the amount of behavioural interference and facilitation, suggesting disparate mechanisms of cognitive processing in each SOA. The current study employed a Stroop task with three SOAs (−400, -200, 0 ms), using functional magnetic resonance imaging to investigate for the first time the neural effects of SOA manipulation. Of specific interest were 1) how SOA affects the neural representation of interference and facilitation; 2) response priming effects in negative SOAs; and 3) attentional effects of blocked SOA presentation.
The results revealed three regions of the executive control network that were sensitive to SOA during Stroop interference; the 0 ms SOA elicited the greatest activation of these areas but experienced relatively smaller behavioural interference, suggesting that the enhanced recruitment led to more efficient conflict processing. Response priming effects were localized to the right inferior frontal gyrus, which is consistent with the idea that this region performed response inhibition in incongruent conditions to overcome the incorrectly-primed response, as well as more general action updating and response preparation. Finally, the right superior parietal lobe was sensitive to blocked SOA presentation and was most active for the 0 ms SOA, suggesting that this region is involved in attentional control.
SOA exerted both trial-specific and block-wide effects on executive processing, providing a unique paradigm for functional investigations of the cognitive control network.
PMCID: PMC3734141  PMID: 23902451
fMRI; Stroop task; Stimulus onset asynchrony; Cognitive control
7.  Inefficient cognitive control in adult ADHD: evidence from trial-by-trial Stroop test and cued task switching performance 
Contemporary neuropsychological models of ADHD implicate impaired cognitive control as contributing to disorder characteristic behavioral deficiencies and excesses; albeit to varying degrees. While the traditional view of ADHD postulates a core deficiency in cognitive control processes, alternative dual-process models emphasize the dynamic interplay of bottom-up driven factors such as activation, arousal, alerting, motivation, reward and temporal processing with top-down cognitive control. However, neuropsychological models of ADHD are child-based and have yet to undergo extensive empirical scrutiny with respect to their application to individuals with persistent symptoms in adulthood. Furthermore, few studies of adult ADHD samples have investigated two central cognitive control processes: interference control and task-set coordination. The current study employed experimental chronometric Stroop and task switching paradigms to investigate the efficiency of processes involved in interference control and task-set coordination in ADHD adults.
22 adults diagnosed with persistent ADHD (17 males) and 22 matched healthy control subjects performed a manual trial-by-trial Stroop color-word test and a blocked explicitly cued task switching paradigm. Performance differences between neutral and incongruent trials of the Stroop task measured interference control. Task switching paradigm manipulations allowed for measurement of transient task-set updating, sustained task-set maintenance, preparatory mechanisms and interference control. Control analyses tested for the specificity of group × condition interactions.
Abnormal processing of task-irrelevant stimulus features was evident in ADHD group performance on both tasks. ADHD group interference effects on the task switching paradigm were found to be dependent on the time allotted to prepare for an upcoming task. Group differences in sustained task-set maintenance and transient task-set updating were also found to be dependent on experimental manipulation of task preparation processes. With the exception of Stroop task error rates, all analyses revealed generally slower and less accurate ADHD group response patterns.
The current data obtained with experimental paradigms deliver novel evidence of inefficient interference control and task-set coordination in adults with persistent ADHD. However, all group differences observed in these central cognitive control processes were found to be partially dependent on atypical ADHD group task preparation mechanisms and/or response inconsistency. These deficiences may have contributed not only to inefficient cognitive control, but also generally slower and less accurate ADHD group performance. Given the inability to dissociate these impairments with the current data, it remains inconclusive as to whether ineffecient cognitive control in the clinical sample was due to top-down failure or bottom-up engagement thereof. To clarify this issue, future neuropsychological investigations are encouraged to employ tasks with significantly more trials and direct manipulations of bottom-up mechanisms with larger samples.
PMCID: PMC1988818  PMID: 17708762
8.  Working Memory Load Modulates Distractor Competition in Primary Visual Cortex 
Cerebral Cortex (New York, NY)  2010;21(3):659-665.
When stimuli compete for sensory processing and response selection, coherent goal-guided behavior requires cognitive control so that task-relevant “targets” rather than irrelevant distractors are selected. It has been shown that reduced cognitive control under high working memory load increases distractor competition for selection. It remains unknown, though, whether cognitive control by working memory has an effect on the earliest levels of sensory processing in primary visual cortex. The present study addressed this question by having subjects perform a selective attention task involving classification of meaningful target objects while also ignoring congruent and incongruent distractor images. The level of cognitive control over distractor competition was varied through a concurrent working memory task of either low (1 digit) or high (6 digits) load. Functional magnetic resonance imaging revealed greater distractor competition effects not only on behavior but also on the sensory correlates in primary visual cortex (areas V1–V2) in conditions of high (vs. low) working memory load. In addition, high working memory load resulted in increased congruency-related functional connectivity between anterior cingulate cortex and V1. These results are the first to establish the neural correlates of distractor competition effects in primary visual cortex and the critical role of working memory in their cognitive control.
PMCID: PMC3041013  PMID: 20699229
attention; cognitive control; fMRI; response competition; working memory
9.  Preparatory Effects of Distractor Suppression: Evidence from Visual Cortex 
PLoS ONE  2011;6(12):e27700.
Spatial selective attention is the mechanism that facilitates the selection of relevant information over irrelevant information in the visual field. The current study investigated whether foreknowledge of the presence or absence of distractors surrounding an impending target stimulus results in preparatory changes in visual cortex. We cued the location of the target and the presence or absence of distractors surrounding the target while changes in blood oxygen level dependent (BOLD) signals were measured. In line with prior work, we found that top-down spatial attention resulted in an increased contralateral BOLD response, evoked by the cue throughout early visual cortex (areas V1, V2 and V3). In addition, cues indicating distractor presence evoked a substantial increase in the magnitude of the BOLD signal in visual area V3, but not in V2 or V1. This study shows that prior knowledge concerning the presence of a distractor results in enhanced attentional modulation of visual cortex, in visual areas where neuronal receptive fields are large enough to encompass both targets and distractors. We interpret these findings as evidence that top-down attentional control processes include active preparatory suppression mechanisms for irrelevant, distracting information in the visual scene.
PMCID: PMC3229494  PMID: 22164213
10.  PTSD symptom severity is associated with increased recruitment of top-down attentional control in a trauma-exposed sample☆ 
NeuroImage : Clinical  2014;7:19-27.
Recent neuroimaging work suggests that increased amygdala responses to emotional stimuli and dysfunction within regions mediating top down attentional control (dorsomedial frontal, lateral frontal and parietal cortices) may be associated with the emergence of anxiety disorders, including posttraumatic stress disorder (PTSD). This report examines amygdala responsiveness to emotional stimuli and the recruitment of top down attention systems as a function of task demands in a population of U.S. military service members who had recently returned from combat deployment in Afghanistan/Iraq. Given current interest in dimensional aspects of pathophysiology, it is worthwhile examining patients who, while not meeting full PTSD criteria, show clinically significant functional impairment.
Fifty-seven participants with sub-threshold levels of PTSD symptoms completed the affective Stroop task while undergoing fMRI. Participants with PTSD or depression at baseline were excluded.
Greater PTSD symptom severity scores were associated with increased amygdala activation to emotional, particularly positive, stimuli relative to neutral stimuli. Furthermore, greater PTSD symptom severity was associated with increased superior/middle frontal cortex response during task conditions relative to passive viewing conditions. In addition, greater PTSD symptom severity scores were associated with: (i) increased activation in the dorsolateral prefrontal, lateral frontal, inferior parietal cortices and dorsomedial frontal cortex/dorsal anterior cingulate cortex (dmFC/dACC) in response to emotional relative to neutral stimuli; and (ii) increased functional connectivity during emotional trials, particularly positive trials, relative to neutral trials between the right amygdala and dmFC/dACC, left caudate/anterior insula cortex, right lentiform nucleus/caudate, bilateral inferior parietal cortex and left middle temporal cortex.
We suggest that these data may reflect two phenomena associated with increased PTSD symptomatology in combat-exposed, but PTSD negative, armed services members. First, these data indicate increased emotional responsiveness by: (i) the positive relationship between PTSD symptom severity and amygdala responsiveness to emotional relative to neutral stimuli; (ii) greater BOLD response as a function of PTSD symptom severity in regions implicated in emotion (striatum) and representation (occipital and temporal cortices) during emotional relative to neutral conditions; and (iii) increased connectivity between the amygdala and regions implicated in emotion (insula/caudate) and representation (middle temporal cortex) as a function of PTSD symptom severity during emotional relative to neutral trials. Second, these data indicate a greater need for the recruitment of regions implicated in top down attention as indicated by (i) greater BOLD response in superior/middle frontal gyrus as a function of PTSD symptom severity in task relative to view conditions; (ii) greater BOLD response in dmFC/dACC, lateral frontal and inferior parietal cortices as a function of PTSD symptom severity in emotional relative to neutral conditions and (iii) greater functional connectivity between the amygdala and inferior parietal cortex as a function of PTSD symptom severity during emotional relative to neutral conditions.
•Greater PTSD symptoms associated with increased amygdala activation to emotional stimuli•PTSD symptoms associated with greater top down attention response in task and emotion conditions•PTSD symptoms were associated with slower reaction times.•Increased top down attention recruitment may compensate for heightened emotional responses.
PMCID: PMC4299952  PMID: 25610763
Post-traumatic stress disorder; Emotion attention; Amygdala; Top down attention
11.  Neural suppression of irrelevant information underlies optimal working memory performance 
Our ability to focus attention on task-relevant information and ignore distractions is reflected by differential enhancement and suppression of neural activity in sensory cortex (i.e., top-down modulation). Such selective, goal-directed modulation of activity may be intimately related to memory, such that the focus of attention biases the likelihood of successfully maintaining relevant information by limiting interference from irrelevant stimuli. Despite recent studies elucidating the mechanistic overlap between attention and memory, the relationship between top-down modulation of visual processing during working memory (WM) encoding and subsequent recognition performance has not yet been established. Here, we provide neurophysiological evidence in healthy, young adults that top-down modulation of early visual processing (< 200 ms from stimulus onset) is intimately related to subsequent WM performance, such that the likelihood of successfully remembering relevant information is associated with limiting interference from irrelevant stimuli. The consequences of a failure to ignore distractors on recognition performance was replicated for two types of feature-based memory, motion direction and color. Moreover, attention to irrelevant stimuli was reflected neurally during the WM maintenance period as an increased memory load. These results suggest that neural enhancement of relevant information is not the primary determinant of high-level performance, but rather, optimal WM performance is dependent on effectively filtering irrelevant information through neural suppression to prevent overloading a limited memory capacity.
PMCID: PMC2704557  PMID: 19279242
working memory; selective attention; top-down modulation; EEG; vision; behavioral performance
12.  The impact of distractor congruency on stimulus processing in retinotopic visual cortex☆ 
Neuroimage  2013;81(100):158-163.
The brain is frequently confronted with sensory information that elicits conflicting response choices. While much research has addressed the top down control mechanisms associated with detection and resolution of response competition, the effects of response competition on sensory processing in the primary visual cortex remain unclear. To address this question we modified a typical ‘flanker task’ (Eriksen and Eriksen, 1974) so that the effects of response competition on human early retinotopic visual cortex could be assessed. Healthy human participants were scanned using fMRI while making a speeded choice response that classified a target object image into one of two categories (e.g. fruits, animals). An irrelevant distractor image that was either congruent (same image as target), incongruent (image from opposite category as target), or neutral (image from task-irrelevant category, e.g. household items) was also present on each trial, but in a different quadrant of the visual field relative to the target. Retinotopic V1 areas responding to the target stimuli showed increased response to targets in the presence of response-incongruent (compared to response-neutral) distractors. A negative correlation with behavioral response competition effects indicated that an increased primary visual cortical response to targets in the incongruent (vs. neutral) trials is associated with a reduced response competition effect on behavior. These results suggest a novel conflict resolution mechanism in the primary visual cortex.
•fMRI was used to examine effects of distractor congruency on retinotopic cortex.•V1 showed increased response to targets in the presence of incongruent distractors.•V1 effect was negatively correlated with distractor congruency effect on behavior.•A mechanism for conflict resolution involving primary visual cortex is proposed.
PMCID: PMC3734350  PMID: 23648965
fMRI; Attention; Response competition; Flanker; V1; Retinotopic cortex
13.  Deficient prefrontal attentional control in late-life generalized anxiety disorder: an fMRI investigation 
Translational Psychiatry  2011;1(10):e46-.
Younger adults with anxiety disorders are known to show an attentional bias toward negative information. Little is known regarding the role of biased attention in anxious older adults, and even less is known about the neural substrates of any such bias. Functional magnetic resonance imaging (fMRI) was used to assess the mechanisms of attentional bias in late life by contrasting predictions of a top-down model emphasizing deficient prefrontal cortex (PFC) control and a bottom-up model emphasizing amygdalar hyperreactivity. In all, 16 older generalized anxiety disorder (GAD) patients (mean age=66 years) and 12 non-anxious controls (NACs; mean age=67 years) completed the emotional Stroop task to assess selective attention to negative words. Task-related fMRI data were concurrently acquired. Consistent with hypotheses, GAD participants were slower to identify the color of negative words relative to neutral, whereas NACs showed the opposite bias, responding more quickly to negative words. During negative words (in comparison with neutral), the NAC group showed PFC activations, coupled with deactivation of task-irrelevant emotional processing regions such as the amygdala and hippocampus. By contrast, GAD participants showed PFC decreases during negative words and no differences in amygdalar activity across word types. Across all participants, greater attentional bias toward negative words was correlated with decreased PFC recruitment. A significant positive correlation between attentional bias and amygdala activation was also present, but this relationship was mediated by PFC activity. These results are consistent with reduced prefrontal attentional control in late-life GAD. Strategies to enhance top-down attentional control may be particularly relevant in late-life GAD treatment.
PMCID: PMC3309492  PMID: 22833192
aging; anxiety; attentional bias; attention; FMRI; prefrontal cortex
14.  Alpha power increases in right parietal cortex reflects focused internal attention 
Neuropsychologia  2014;56(100):393-400.
This study investigated the functional significance of EEG alpha power increases, a finding that is consistently observed in various memory tasks and specifically during divergent thinking. It was previously shown that alpha power is increased when tasks are performed in mind—e.g., when bottom-up processing is prevented. This study aimed to examine the effect of task-immanent differences in bottom-up processing demands by comparing two divergent thinking tasks, one intrinsically relying on bottom-up processing (sensory-intake task) and one that is not (sensory-independence task). In both tasks, stimuli were masked in half of the trials to establish conditions of higher and lower internal processing demands. In line with the hypotheses, internal processing affected performance and led to increases in alpha power only in the sensory-intake task, whereas the sensory-independence task showed high levels of task-related alpha power in both conditions. Interestingly, conditions involving focused internal attention showed a clear lateralization with higher alpha power in parietal regions of the right hemisphere. Considering evidence from fMRI studies, right-parietal alpha power increases may correspond to a deactivation of the right temporoparietal junction, reflecting an inhibition of the ventral attention network. Inhibition of this region is thought to prevent reorienting to irrelevant stimulation during goal-driven, top-down behavior, which may serve the executive function of task shielding during demanding cognitive tasks such as idea generation and mental imagery.
•This study investigated the functional significance of EEG alpha activity.•Right-parietal alpha power increased as a function of internal attention demands.•Alpha power increases during divergent thinking indicates focused attention.•Right-parietal alpha may reflect activity of the ventral attention network.
PMCID: PMC3989020  PMID: 24561034
Attention; Memory; Inhibition; EEG; Alpha; Divergent thinking
15.  Temporal-Spatial Neural Activation Patterns Linked to Perceptual Encoding of Emotional Salience 
PLoS ONE  2014;9(4):e93753.
It is well known that we continuously filter incoming sensory information, selectively allocating attention to what is important while suppressing distracting or irrelevant information. Yet questions remain about spatiotemporal patterns of neural processes underlying attentional biases toward emotionally significant aspects of the world. One index of affectively biased attention is an emotional variant of an attentional blink (AB) paradigm, which reveals enhanced perceptual encoding for emotionally salient over neutral stimuli under conditions of limited executive attention. The present study took advantage of the high spatial and temporal resolution of magnetoencephalography (MEG) to investigate neural activation related to emotional and neutral targets in an AB task. MEG data were collected while participants performed a rapid stimulus visual presentation task in which two target stimuli were embedded in a stream of distractor words. The first target (T1) was a number and the second (T2) either an emotionally salient or neutral word. Behavioural results replicated previous findings of greater accuracy for emotionally salient than neutral T2 words. MEG source analyses showed that activation in orbitofrontal cortex, characterized by greater power in the theta and alpha bands, and dorsolateral prefrontal activation were associated with successful perceptual encoding of emotionally salient relative to neutral words. These effects were observed between 250 and 550 ms, latencies associated with discrimination of perceived from unperceived stimuli. These data suggest that important nodes of both emotional salience and frontoparietal executive systems are associated with the emotional modulation of the attentional blink.
PMCID: PMC3984101  PMID: 24727751
16.  Signal Enhancement and Suppression During Visual-Spatial Selective Attention 
Brain research  2010;1359:155-177.
Selective attention involves the relative enhancement of relevant versus irrelevant stimuli. However, whether this relative enhancement involves primarily enhancement of attended stimuli, or suppression of irrelevant stimuli, remains controversial. Moreover, if both enhancement and suppression are involved, whether they result from a single mechanism or separate mechanisms during attentional control or selection is not known. In two experiments using a spatial cuing paradigm with task-relevant targets and irrelevant distractors, target and distracter processing was examined as a function of distractor expectancy. Additionally, in the second study the interaction of perceptual load and distractor expectancy was explored. In both experiments, distractors were either validly cued (70%) or invalidly cued (30%) in order to examine the effects of distractor expectancy on attentional control as well as target and distractor processing. The effects of distractor expectancy were assessed using event-related potentials recorded during the cue-to-target period (preparatory attention) and in response to the task-relevant target stimuli (selective stimulus processing). Analyses of distractor-present displays (anticipated versus unanticipated), showed modulations in brain activity during both the preparatory period and during target processing. The pattern of brain responses suggest both facilitation of attended targets and suppression of unattended distractors. These findings provide evidence for a two-process model of visual spatial selective attention, where one mechanism (facilitation) influences relevant stimuli and another (suppression) acts to filter distracting stimuli.
PMCID: PMC2955768  PMID: 20807513
Selective Attention; Event Related Potential; Inhibition; Vision
17.  The role of trauma-related distractors on neural systems for working memory and emotion processing in posttraumatic stress disorder 
Journal of psychiatric research  2008;43(8):809-817.
The relevance of emotional stimuli to threat and survival confers a privileged role in their processing. In PTSD, the ability of trauma-related information to divert attention is especially pronounced. Information unrelated to the trauma may also be highly distracting when it shares perceptual features with trauma material. Our goal was to study how trauma-related environmental cues modulate working memory networks in PTSD. We examined neural activity in participants performing a visual working memory task while distracted by task-irrelevant trauma and non-trauma material. Recent post-9/11 veterans were divided into a PTSD group (n = 22) and a trauma-exposed control group (n = 20) based on the Davidson trauma scale. Using fMRI, we measured hemodynamic change in response to emotional (trauma-related) and neutral distraction presented during the active maintenance period of a delayed-response working memory task. The goal was to examine differences in functional networks associated with working memory (dorsolateral prefrontal cortex and lateral parietal cortex) and emotion processing (amygdala, ventrolateral prefrontal cortex, and fusiform gyrus). The PTSD group showed markedly different neural activity compared to the trauma-exposed control group in response to task-irrelevant visual distractors. Enhanced activity in ventral emotion processing regions was associated with trauma distractors in the PTSD group, whereas activity in brain regions associated with working memory and attention regions was disrupted by distractor stimuli independent of trauma content. Neural evidence for the impact of distraction on working memory is consistent with PTSD symptoms of hypervigilance and general distractibility during goal-directed cognitive processing.
PMCID: PMC2684984  PMID: 19091328
PTSD; fMRI; Working memory; Emotion processing; Cognitive control
18.  Neural Substrates of Attentive Listening Assessed with a Novel Auditory Stroop Task 
A common explanation for the interference effect in the classic visual Stroop test is that reading a word (the more automatic semantic response) must be suppressed in favor of naming the text color (the slower sensory response). Neuroimaging studies also consistently report anterior cingulate/medial frontal, lateral prefrontal, and anterior insular structures as key components of a network for Stroop-conflict processing. It remains unclear, however, whether automatic processing of semantic information can explain the interference effect in other variants of the Stroop test. It also is not known if these frontal regions serve a specific role in visual Stroop conflict, or instead play a more universal role as components of a more generalized, supramodal executive-control network for conflict processing. To address these questions, we developed a novel auditory Stroop test in which the relative dominance of semantic and sensory feature processing is reversed. Listeners were asked to focus either on voice gender (a more automatic sensory discrimination task) or on the gender meaning of the word (a less automatic semantic task) while ignoring the conflicting stimulus feature. An auditory Stroop effect was observed when voice features replaced semantic content as the “to-be-ignored” component of the incongruent stimulus. Also, in sharp contrast to previous Stroop studies, neural responses to incongruent stimuli studied with functional magnetic resonance imaging revealed greater recruitment of conflict loci when selective attention was focused on gender meaning (semantic task) over voice gender (sensory task). Furthermore, in contrast to earlier Stroop studies that implicated dorsomedial cortex in visual conflict processing, interference-related activation in both of our auditory tasks was localized ventrally in medial frontal areas, suggesting a dorsal-to-ventral separation of function in medial frontal cortex that is sensitive to stimulus context.
PMCID: PMC3020403  PMID: 21258643
anterior cingulate; anterior insula; attention; auditory Stroop; conflict processing; fMRI; medial frontal gyrus
19.  Proactive and Reactive Control During Emotional Interference and its Relationship to Trait Anxiety 
Brain research  2012;1481:13-36.
In classic Stroop paradigms, increasing the proportion of control-demanding incongruent trials results in strategic adjustments in behavior and implementation of cognitive control processes. We manipulated expectancy for incongruent trials in an emotional facial Stroop task to investigate the behavioral and neural effects of proportion manipulation in a cognitively demanding task with emotional stimuli. Subjects performed a high expectancy (HE) task (65% incongruent trials) and a low expectancy (LE) task (35% incongruent trials) during functional magnetic resonance imaging (fMRI). As in standard Stroop tasks, behavioral interference was reduced in the emotional facial Stroop HE task compared to the LE task. Functional MRI data revealed a switch in cognitive control strategy, from a reactive, event-related activation of a medial and lateral cognitive control network and right amygdala in the LE task to a proactive, sustained activation of right dorsolateral prefrontal cortex (DLPFC) in the HE task. Higher trait anxiety was associated with impairment (slower response time and decreased accuracy) as well as reduced activity in left ventrolateral prefrontal cortex, anterior insula, and orbitofrontal cortex in the HE task on high conflict trials with task-irrelevant emotional information, suggesting that individual differences in anxiety may be associated with expectancy-related strategic control adjustments, particularly when emotional stimuli must be ignored.
PMCID: PMC3541031  PMID: 22960116
amygdala; anterior cingulate cortex; conflict monitoring; fMRI; prefrontal cortex; sustained and transient control
20.  Occipital Alpha Activity during Stimulus Processing Gates the Information Flow to Object-Selective Cortex 
PLoS Biology  2014;12(10):e1001965.
A simultaneous EEG-fMRI study demonstrates that alpha-band activity in early visual cortex is associated with gating visual information to downstream regions, boosting attended information and suppressing distraction.
Given the limited processing capabilities of the sensory system, it is essential that attended information is gated to downstream areas, whereas unattended information is blocked. While it has been proposed that alpha band (8–13 Hz) activity serves to route information to downstream regions by inhibiting neuronal processing in task-irrelevant regions, this hypothesis remains untested. Here we investigate how neuronal oscillations detected by electroencephalography in visual areas during working memory encoding serve to gate information reflected in the simultaneously recorded blood-oxygenation-level-dependent (BOLD) signals recorded by functional magnetic resonance imaging in downstream ventral regions. We used a paradigm in which 16 participants were presented with faces and landscapes in the right and left hemifields; one hemifield was attended and the other unattended. We observed that decreased alpha power contralateral to the attended object predicted the BOLD signal representing the attended object in ventral object-selective regions. Furthermore, increased alpha power ipsilateral to the attended object predicted a decrease in the BOLD signal representing the unattended object. We also found that the BOLD signal in the dorsal attention network inversely correlated with visual alpha power. This is the first demonstration, to our knowledge, that oscillations in the alpha band are implicated in the gating of information from the visual cortex to the ventral stream, as reflected in the representationally specific BOLD signal. This link of sensory alpha to downstream activity provides a neurophysiological substrate for the mechanism of selective attention during stimulus processing, which not only boosts the attended information but also suppresses distraction. Although previous studies have shown a relation between the BOLD signal from the dorsal attention network and the alpha band at rest, we demonstrate such a relation during a visuospatial task, indicating that the dorsal attention network exercises top-down control of visual alpha activity.
Author Summary
In complex environments, our sensory systems are bombarded with information. Only a fraction of this information is processed, whereas most is ignored. As such, our brain must rely on powerful mechanisms to filter the relevant information. It has been proposed that alpha band oscillations (8–13 Hz) gate task-relevant visual information to downstream areas and supress irrelevant visual information. We tested this hypothesis in a study that combined electroencephalography (EEG) and functional MRI (fMRI) recordings. From the EEG, we directly measured alpha band oscillations in early visual regions. Using fMRI, we quantified neuronal activity in downstream regions. The participants performed a spatial working memory task that required them to encode pictures of objects presented in the left field of view while ignoring objects in the right field (or vice versa). We found that suppression of alpha band activity in visual areas opened the gate for relevant visual information to be routed to downstream regions. Conversely, an increase in alpha oscillations suppressed visual information that was irrelevant to the task. These findings suggest that alpha band oscillations are directly involved in boosting attended information and suppressing distraction in the ventral visual stream.
PMCID: PMC4205112  PMID: 25333286
21.  Cardiorespiratory Fitness and Attentional Control in the Aging Brain 
A growing body of literature provides evidence for the prophylactic influence of cardiorespiratory fitness on cognitive decline in older adults. This study examined the association between cardiorespiratory fitness and recruitment of the neural circuits involved in an attentional control task in a group of healthy older adults. Employing a version of the Stroop task, we examined whether higher levels of cardiorespiratory fitness were associated with an increase in activation in cortical regions responsible for imposing attentional control along with an up-regulation of activity in sensory brain regions that process task-relevant representations. Higher fitness levels were associated with better behavioral performance and an increase in the recruitment of prefrontal and parietal cortices in the most challenging condition, thus providing evidence that cardiorespiratory fitness is associated with an increase in the recruitment of the anterior processing regions. There was a top-down modulation of extrastriate visual areas that process both task-relevant and task-irrelevant attributes relative to the baseline. However, fitness was not associated with differential activation in the posterior processing regions, suggesting that fitness enhances attentional function by primarily influencing the neural circuitry of anterior cortical regions. This study provides novel evidence of a differential association of fitness with anterior and posterior brain regions, shedding further light onto the neural changes accompanying cardiorespiratory fitness.
PMCID: PMC3024830  PMID: 21267428
cardiorespiratory fitness; Stroop task; cognitive and attentional control
22.  The anodal tDCS over the left posterior parietal cortex enhances attention toward a focus word in a sentence 
The posterior parietal cortex (PPC) has two attentional functions: top-down attentional control and stimulus-driven attentional processing. Using the focused version of the reading span test (RST), in which the target word to be remembered is the critical word for comprehending a sentence (focused word) or a non-focused word, we examined the effect of tDCS on resolution of distractor interference by the focused word in the non-focus condition (top-down attentional control) and on augmented/shrunk attentional capture by the focused word in both the focus and non-focus conditions (stimulus-driven attentional processing). Participants were divided into two groups: anodal tDCS (atDCS) and cathodal tDCS (ctDCS). Online stimulation was given while participants performed the RST. A post-hoc recognition task was also administered in which three kinds of words were presented: target words in the RST, distractor words in the RST, and novel words. atDCS augmented the effect of the focused word by increasing differences in performance between the focus and non-focus conditions. Such an effect was not observed in the ctDCS group. As for the recognition task, atDCS again produced the augmented effect of the focused words in the distractor recognition. On the other hand, ctDCS brought less recognition of non-focused target words in comparison to sham. The results indicate that atDCS promotes stimulus-driven attentional processing, possibly by affecting neural firing in the inferior parietal regions. In contrast, ctDCS appears to prevent retrieval of less important information from episodic memory, which may require top-down attentional processing.
PMCID: PMC4260498  PMID: 25538609
posterior parietal cortex; tDCS; attention; working memory; reading span test
23.  Neural bases of selective attention in action video game players 
Vision Research  2011;61:132-143.
Over the past few years, the very act of playing action video games has been shown to enhance several different aspects of visual selective attention. Yet little is known about the neural mechanisms that mediate such attentional benefits. A review of the aspects of attention enhanced in action game players suggests there are changes in the mechanisms that control attention allocation and its efficiency (Hubert-Wallander et al., 2010). The present study used brain imaging to test this hypothesis by comparing attentional network recruitment and distractor processing in action gamers versus non-gamers as attentional demands increased. Moving distractors were found to elicit lesser activation of the visual motion-sensitive area (MT/MST) in gamers as compared to non-gamers, suggestive of a better early filtering of irrelevant information in gamers. As expected, a fronto-parietal network of areas showed greater recruitment as attentional demands increased in non-gamers. In contrast, gamers barely engaged this network as attentional demands increased. This reduced activity in the fronto-parietal network that is hypothesized to control the flexible allocation of top-down attention is compatible with the proposal that action game players may allocate attentional resources more automatically, possibly allowing more efficient early filtering of irrelevant information.
PMCID: PMC3260403  PMID: 21864560
action video games; brain plasticity; visual attention; fMRI; perceptual load; fronto-parietal network
24.  Neural mechanisms mediating contingent capture of attention by affective stimuli 
Journal of cognitive neuroscience  2012;24(5):1113-1126.
Attention is attracted exogenously by physically salient stimuli, but this effect can be dampened by endogenous attention settings, a phenomenon called “contingent capture”. Emotionally salient stimuli are also thought to exert a strong exogenous influence on attention, especially in anxious individuals, but whether and how top-down attention can ameliorate bottom-up capture by affective stimuli is currently unknown. Here, we paired a novel spatial cueing task with functional magnetic resonance imaging (fMRI) in order to investigate contingent capture as a function of the affective salience of bottom-up cues (face stimuli) and individual differences in trait anxiety. In the absence of top-down cues, exogenous stimuli validly cueing targets facilitated attention in low anxious participants, regardless of affective salience. However, while high anxious participants exhibited similar facilitation following neutral exogenous cues, this facilitation was completely absent following affectively negative exogenous cues. Critically, these effects were contingent on endogenous attentional settings, such that explicit top-down cues presented prior to the appearance of exogenous stimuli removed anxious individuals’ sensitivity to affectively salient stimuli. FMRI analyses revealed a network of brain regions underlying this variability in affective contingent capture across individuals, including the fusiform face area (FFA), posterior ventrolateral frontal cortex, and supplementary motor area. Importantly, activation in the posterior ventrolateral frontal cortex and the supplementary motor area fully mediated the effects observed in FFA, demonstrating a critical role for these frontal regions in mediating attentional orienting and interference resolution processes when engaged by affectively salient stimuli.
PMCID: PMC3654100  PMID: 22360642
Attention; Posner cueing paradigm; ventral attention network; contingent capture; emotion; fear
25.  More than Meets the Eye: Age Differences in the Capture and Suppression of Oculomotor Action 
Salient visual stimuli capture attention and trigger an eye-movement toward its location reflexively, regardless of an observer’s intentions. Here we aim to investigate the effect of aging (1) on the extent to which salient yet task-irrelevant stimuli capture saccades, and (2) on the ability to selectively suppress such oculomotor responses. Young and older adults were asked to direct their eyes to a target appearing in a stimulus array. Analysis of overall performance shows that saccades to the target object were disrupted by the appearance of a task-irrelevant abrupt-onset distractor when the location of this distractor did not coincide with that of the target object. Conditional capture function analyses revealed that, compared to young adults, older adults were more susceptible to oculomotor capture, and exhibited deficient selective suppression of the responses captured by task-irrelevant distractors. These effects were uncorrelated, suggesting two independent sources off age-related decline. Thus, with advancing age, salient visual distractors become more distracting; in part because they trigger reflexive eye-movements more potently; in part because of failing top-down control over such reflexes. The fact that these process-specific age effects remained concealed in overall oculomotor performance analyses emphasizes the utility of looking beyond the surface; indeed, there may be more than meets the eye.
PMCID: PMC3201057  PMID: 22046165
oculomotor capture; inhibitory control; saccades; aging; distributional analysis

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