Preparatory allocation of attention and adjustments in conflict processing

Attentional control involves the ability to allocate preparatory attention to improve subsequent stimulus processing and response selection. There is behavioral evidence to support the hypothesis that increased expectancy of stimulus and response conflict may decrease the subsequent experience of conflict during task performance. We used a cued flanker and event-related fMRI design to separate processes involved in preparation from those involved in resolving conflict and to identify the brain systems involved in these processes as well as the association between preparatory activity levels and activity related to subsequent conflict processing. Our results demonstrate that preparatory attentional allocation following a cue to the upcoming level of conflict is mediated by a network involving Dorsolateral Prefrontal Cortex (DLPFC) and the Intraparietal Sulcus (IPS). Informed preparation for conflict processing was associated with decreased Anterior Cingulate Cortex/pre-Supplementary Motor Area (ACC/pre-SMA) and IPS activity during the flanker target presentation, supporting their roles in conflict processing and visuospatial attention during the flanker task. Ventrolateral Prefrontal Cortex/Orbitofrontal Cortex (VLPFC/OFC) was active when specific strategic task rule and outcome information was available.     

Cerebral correlates of alerting, orienting and reorienting of visuospatial attention: an event-related fMRI study

The identification of brain systems contributing to different aspects of visuospatial attention is of both clinical and theoretical interest. Cued target detection tasks provide a simple means to dissociate attentional subcomponents, such as alerting, orienting or reorienting of attention. Event-related functional magnetic resonance imaging (fMRI) was used to study neural correlates of these distinct attentional processes. Volunteers were scanned while performing a centrally cued target detection task. Four different types of trials (no cue, neutral cue, valid cue and invalid cue trials) with targets appearing either in the right or left hemifield were randomly intermixed. Behaviourally, the data provided evidence for alerting, spatial orienting and reorienting of attention. Neurally, the alerting effect was seen in bilaterally increased extrastriatal blood oxygenation level-dependent (BOLD) activity in neutral as compared to no cue trials. Neural correlates of spatial orienting were seen in anterior cingulate cortex, which was more active during valid as compared to neutral cue trials. Neural correlates of reorienting of attention, that is, higher BOLD activity to invalid as compared to validly cued trials were evident in several brain regions including left and right intraparietal sulcus, right temporo-parietal junction and middle frontal gyrus bilaterally. The data suggest that frontal and parietal regions are specifically involved in reorienting rather than orienting attention to a spatial position. Alerting effects were seen in extrastriate regions which suggest that increased phasic alertness results in a top-down modulation of neural activity in visual processing areas.     

BOLD signal in intraparietal sulcus covaries with magnitude of implicitly driven attention shifts

A lot is known about the neural basis of directing attention based on explicit cues. In real life however, attention shifts are rarely directed by explicit cues but rather generated implicitly, for example on the basis of previous experience with a given situation. Here, we aimed at studying attention shifts dependent on recent trial history. While explicitly cued attention shifts involve activity in cortex of the intraparietal sulcus, whether this region is also involved in shifting attention according to recent history is still unknown. We asked observers to detect targets in a stream of visual stimuli with three feature dimensions: color, shape and motion. Critically, target occurrence probability was always higher in one stimulus dimension than in the others, and probabilities switched between dimensions over blocks of trials. After each probability switch, target detection times decreased exponentially for high-probability targets and increased for low-probability targets, compatible with gradual shifts in attention dependent on trial history since the switch. BOLD signal in left prefrontal and intraparietal sulcus regions was higher in the early phase after the switch, while anterior cingulate, cuneus, precuneus, temporal and more anterior frontal regions showed more activation later after the switch. These findings are compatible with the engagement of regions involved in the establishment and maintenance of attentional sets. BOLD signal in left intraparietal sulcus correlated with the size of the performance changes consecutive to the detected targets, suggesting that it reflects the size of attention shifts induced by updating target probabilities over recent trial history.     

Cue validity modulates the neural correlates of covert endogenous orienting of attention in parietal and frontal cortex

Parietal brain regions have been implicated in reorienting of visuospatial attention in location-cueing paradigms when misleading advance information is provided in form of a spatially invalid cue. The difference in reaction times to invalidly and validly cued targets is termed the "validity effect" and used as a behavioral measure for attentional reorienting. Behavioral studies suggest that the magnitude of the validity effect depends on the ratio of validly to invalidly cued targets (termed cue validity), i.e., on the amount of top-down information provided. Using fMRI, we investigated the effects of a cue validity manipulation upon the neural mechanisms underlying attentional reorienting using valid and invalid spatial cues in the context of 90% and 60% cue validity, respectively. We hypothesized that increased parietal activation would be elicited when subjects need to reorient their attention in a context of high cue validity. Behaviorally, subjects showed significantly higher validity effects in the high as compared to the low cue validity condition, indicating slower reorienting. The neuroimaging data revealed higher activation of right inferior parietal and right frontal cortex in the 90% than in the 60% cue validity condition. We conclude that the amount of top-down information provided by predictive cues influences the neural correlates of reorienting of visuospatial attention by modulating activation of a right fronto-parietal attentional network.     

Regional brain activation during meditation shows time and practice effects: an exploratory FMRI study

Meditation involves attentional regulation and may lead to increased activity in brain regions associated with attention such as dorsal lateral prefrontal cortex (DLPFC) and anterior cingulate cortex (ACC). Using functional magnetic resonance imaging, we examined whether DLPFC and ACC were activated during meditation. Subjects who meditate were recruited and scanned on a 3.0 Tesla scanner. Subjects meditated for four sessions of 12 min and performed four sessions of a 6 min control task. Individual and group t-maps were generated of overall meditation response versus control response and late meditation response versus early meditation response for each subject and time courses were plotted. For the overall group (n = 13), and using an overall brain analysis, there were no statistically significant regional activations of interest using conservative thresholds. A region of interest analysis of the entire group time courses of DLPFC and ACC were statistically more active throughout meditation in comparison to the control task. Moreover, dividing the cohort into short (n = 8) and long-term (n = 5) practitioners (>10 years) revealed that the time courses of long-term practitioners had significantly more consistent and sustained activation in the DLPFC and the ACC during meditation versus control in comparison to short-term practitioners. The regional brain activations in the more practised subjects may correlate with better sustained attention and attentional error monitoring. In summary, brain regions associated with attention vary over the time of a meditation session and may differ between long- and short-term meditation practitioners.     

不同提示条件下视空间注意的事件相关电位研究

目的：采用“提示-目标”的视觉实验范式,分别以汉字和范围提示不同等级的搜索范围,通过事件相关电位（ERP）技术,研究视觉注意脑机制：方法：16名在校大学生作为有偿被试,刺激序列为背景-提示-靶刺激：背景由三个同心白色圆形线条组成。刺激材料是随机选取的大写英文字母,组成大、中、小3个同心圆圈。指定“T”为靶刺激,大提示时,目标“T”可能会出现在大中小三个圈内;中提示时,“T”出现在中小两个圈内;当小提示时,目标只出现在小圈内。结果：汉字提示的反应时明显长于范围提示：同时与范围提示相比,汉字提示下的靶刺激引起后部P1的增强和N1的抑制以及P2的增强。结论：范围提示能够加快靶刺激识别或视觉搜索的速度,汉字提示时启动的是内源性注意系统,在早期感知阶段,前额叶资源的利用不够完善,证明了汉字提示与范围提示具有不同的加工机制．     

Preparatory activity and connectivity in dorsal anterior cingulate cortex for cognitive control

Dorsal anterior cingulate cortex (dACC) is composed of functionally distinct subregions that may contribute to the top-down control of response selection and preparation. Multiple motor areas have been identified in dACC, including an anterior zone implicated in conflict monitoring and a caudal zone involved in movement execution. This study tested the involvement of a third cingulate area, the posterior zone of dACC, in the top-down control of response selection and preparation. Sixteen healthy young adults were scanned with event-related functional magnetic resonance imaging while performing a cued go/no-go task that was designed to minimize response conflicts. The activation and functional connectivity of dACC were tested with standard convolution models and psychophysiological interaction analyses, respectively. Ready cues that informed the direction of the impending response triggered preparatory neural activity in the posterior zone of dACC and strengthened functional connectivity with the anterior and caudal zones of dACC, as well as perigenual anterior cingulate cortex, frontal operculum, dorsolateral prefrontal cortex, sensory association cortices, and extra-pyramidal motor areas. The preparatory cues activated dACC above and beyond the general arousing effects common to cues despite negligible conflict in the go/no-go task. The integration of cognitive, sensorimotor, and incentive signals in dACC places the region in an ideal position to select and prepare appropriate behavioral responses to achieve higher-level goals.     

Impaired disengagement from threatening cues of impending pain in a crossmodal cueing paradigm.

This paper reports an experimental investigation of attentional engagement to and disengagement from cues of impending pain. Pain-free volunteers performed a cueing task in which they were instructed to detect somatosensory and tone targets. Target stimuli were preceded by visual cues informing participants of the modality of the impending stimuli. Participants were randomly assigned to a pain group (n = 54) or to a control group (n = 53). Somatosensory targets consisted of painful electrocutaneous stimuli in the pain group and non-painful vibrotactile targets in the control group. Analyses revealed a similar amount of attentional engagement to both cues signalling somatosensory targets, irrespective of their threat value. However, participants had significantly more difficulty in disengaging attention from a threatening cue of impending pain compared to a cue signalling the non-painful vibrotactile target. Our findings provide further evidence that pain cues demand attention, particularly resulting in impaired disengagement.     

Practice-related effects demonstrate complementary roles of anterior cingulate and prefrontal cortices in attentional control

The purpose of this study was to test the hypothesis that the dorsolateral prefrontal cortex (DLFPC), not the anterior cingulate cortex (ACC), plays the predominant role in implementing top-down attentional control. To do so, we used fMRI to examine practice-related changes in neural activity during a variant of the Stroop task. The results indicated that the DLPFC's activity decreased gradually as the need for control was reduced (as indexed by behavioral measures), while the ACC's activity dropped off rapidly. Such a pattern is consistent with the DLPFC taking a leading role in implementing top-down attentional control and the ACC being involved in other aspects of attentional control, such as response-related processes. In addition, with practice, there was a reduction in activity within cortical systems handling the processing of task-irrelevant information capable of interfering with task performance. This finding suggests that with practice the brain is capable of identifying and strategically inhibiting such processing.     

Heterogeneity of Cingulate Contributions to Spatial Attention

Functional magnetic resonance imaging was used to investigate activation patterns within the cingulate region during tasks based on spatial attention. Subjects were asked to detect targets which appeared either at the site indicated by a cue or on the opposite side. A “cue effect” was identified by the presence of shorter reaction times to validly than invalidly cued targets, showing that an anticipatory bias had been generated in the direction of the cue. Target detection accuracy was consistently above 90% although cue effects and reaction times displayed substantial variations, from one task session to another. Activation within the anterior cingulate region was seen in 16 of the 26 sessions but showed no correlation with reaction time. Posterior cingulate activation was seen in only 6 of the 26 sessions. However, a random effects analysis showed that the task-related signal change in this region was strongly correlated with the speed of target detection. A post hoc analysis indicated that this correlation was significant only when cue effects were present. No other part of the cerebral cortex displayed significant correlations with reaction times or cue effects. These results suggest that the cingulate component of the attentional network has at least two functionally segregated sectors, an anterior one in BA 24/32 and a posterior cingulo-retrosplenial one in BA 23/29/30. The posterior sector appears to be associated with the speed of detecting spatial targets, especially when attention is under the influence of a cue-induced anticipatory bias. The anterior cingulate focus did not display such a relationship in our tasks and is likely to mediate other aspects of attentional deployment such as performance monitoring, response selection or target identification.     

How efficient is the orienting of spatial attention to pain? An experimental investigation

This study investigated how efficient spatial attention was oriented to pain in 2 experiments. Participants detected whether painful (pain group) or nonpainful (control group) somatosensory stimuli were delivered to the left or right hand. Each stimulus was preceded by a visual cue presented near to the stimulated hand (valid trial), the opposite hand (invalid trial), or centrally between hands. To examine both exogenous and endogenous orienting of attention, the spatial predictability of somatosensory targets was manipulated. In the first experiment, visual cues were nonpredictive for the location of the pain stimulus, as a result of which, orienting was purely exogenous, i.e., resulting from the occurrence of the visual cue at the location of somatosensory input. In the second experiment, visual cues were spatially predictive, as a result of which, endogenous control was added, i.e., attention driven by expectations of where the somatosensory target will occur. The results showed that only in experiment 1 was spatial attention oriented more efficiently to painful compared with nonpainful somatosensory stimulation. This effect was due to faster responses on valid relative to baseline trials (engagement), rather than slower responses on invalid relative to baseline trials (disengagement), and was significantly correlated with self-reported bodily threat. In experiment 2, prioritization of the pain location was probably overridden by task strategies because it was advantageous for participants' task performance to attend to the cued location irrespective of whether stimulation was painful or not. Implications of these findings for theories of hypervigilance and attentional management of pain are discussed.     

Left posterior parietal cortex participates in both task preparation and episodic retrieval

Optimal memory retrieval depends not only on the fidelity of stored information, but also on the attentional state of the subject. Factors such as mental preparedness to engage in stimulus processing can facilitate or hinder memory retrieval. The current study used functional magnetic resonance imaging (fMRI) to distinguish preparatory brain activity before episodic and semantic retrieval tasks from activity associated with retrieval itself. A catch-trial imaging paradigm permitted separation of neural responses to preparatory task cues and memory probes. Episodic and semantic task preparation engaged a common set of brain regions, including the bilateral intraparietal sulcus (IPS), left fusiform gyrus (FG), and the pre-supplementary motor area (pre-SMA). In the subsequent retrieval phase, the left IPS was among a set of frontoparietal regions that responded differently to old and new stimuli. In contrast, the right IPS responded to preparatory cues with little modulation during memory retrieval. The findings support a strong left-lateralization of retrieval success effects in left parietal cortex, and further indicate that left IPS performs operations that are common to both task preparation and memory retrieval. Such operations may be related to attentional control, monitoring of stimulus relevance, or retrieval.     

Preparatory deployment of attention to motion activates higher-order motion-processing brain regions

We used event-related fMRI to test the hypothesis that preparatory attention modulations occur in higher-order motion-processing regions when subjects deploy attention to internally driven representations in a complex motion-processing task. Using a cued attention-to-motion task, we found preparatory increases in fMRI activity in visual motion regions in the absence of visual motion stimulation. The cue, a brief enlargement of the fixation cross, directed subjects to prepare for a complex motion discrimination task. This preparation activated higher-order and lower-order motion regions. The motion regions activated included temporal regions consistent with V5/MT+, occipital regions consistent with V3+, parietal-occipital junction regions, ventral and dorsal intraparietal sulcus, superior temporal sulcus (STS), posterior insular cortex (PIC), and a region of BA 39/40 superior to V5/MT+ involving the angular gyrus and supramarginal gyrus (A-SM). Consistent with our hypothesis that these motion sensory activations are under top-down control, we also found activation of an extensive frontal network during the cue period, including anterior cingulate and multiple prefrontal regions. These results support the hypothesis that anticipatory deployment of attention to internally driven representations is achieved via top-down modulation of activity in task-relevant processing areas.     

Hypnosis decouples cognitive control from conflict monitoring processes of the frontal lobe

Hypnosis can profoundly alter sensory awareness and cognitive processing. While the cognitive and behavioral phenomena associated with hypnosis have long been thought to relate to attentional processes, the neural mechanisms underlying susceptibility to hypnotic induction and the hypnotic condition are poorly understood. Here, we tested the proposal that highly hypnotizable individuals are particularly adept at focusing attention at baseline, but that their attentional control is compromised following hypnosis due to a decoupling between conflict monitoring and cognitive control processes of the frontal lobe. Employing event-related fMRI and EEG coherence measures, we compared conflict-related neural activity in the anterior cingulate cortex (ACC) and control-related activity in the lateral frontal cortex (LFC) during Stroop task performance between participants of low and high hypnotic susceptibility, at baseline and after hypnotic induction. The fMRI data revealed that conflict-related ACC activity interacted with hypnosis and hypnotic susceptibility, in that highly susceptible participants displayed increased conflict-related neural activity in the hypnosis condition compared to baseline, as well as with respect to subjects with low susceptibility. Cognitive-control-related LFC activity, on the other hand, did not differ between groups and conditions. These data were complemented by a decrease in functional connectivity (EEG gamma band coherence) between frontal midline and left lateral scalp sites in highly susceptible subjects after hypnosis. These results suggest that individual differences in hypnotic susceptibility are linked with the efficiency of the frontal attention system, and that the hypnotized condition is characterized by a functional dissociation of conflict monitoring and cognitive control processes.     

Atypical neural networks for social orienting in autism spectrum disorders

Autism spectrum disorders (ASD) are characterized by significant social impairments, including deficits in orienting attention following social cues. Behavioral studies investigating social orienting in ASD, however, have yielded mixed results, as the use of naturalistic paradigms typically reveals clear deficits whereas computerized laboratory experiments often report normative behavior. The present study is the first to examine the neural mechanisms underlying social orienting in ASD in order to provide new insight into the social attention impairments that characterize this disorder. Using fMRI, we examined the neural correlates of social orienting in children and adolescents with ASD and in a matched sample of typically developing (TD) controls while they performed a spatial cueing paradigm with social (eye gaze) and nonsocial (arrow) cues. Cues were either directional (indicating left or right) or neutral (indicating no direction), and directional cues were uninformative of the upcoming target location in order to engage automatic processes by minimizing expectations. Behavioral results demonstrated intact orienting effects for social and nonsocial cues, with no differences between groups. The imaging results, however, revealed clear group differences in brain activity. When attention was directed by social cues compared to nonsocial cues, the TD group showed increased activity in frontoparietal attention networks, visual processing regions, and the striatum, whereas the ASD group only showed increased activity in the superior parietal lobule. Significant group × cue type interactions confirmed greater responsivity in task-relevant networks for social cues than nonsocial cues in TD as compared to ASD, despite similar behavioral performance. These results indicate that, in the autistic brain, social cues are not assigned the same privileged status as they are in the typically developing brain. These findings provide the first empirical evidence that the neural circuitry involved in social orienting is disrupted in ASD and highlight that normative behavioral performance in a laboratory setting may reflect compensatory mechanisms rather than intact social attention.     

Cooperation of the anterior cingulate cortex and dorsolateral prefrontal cortex for attention shifting

Attention shifting in the working memory system plays an important role in goal-oriented behavior, such as reading, reasoning, and driving, because it involves several cognitive processes. This study identified brain activity leading to individual differences in attention shifting for dual-task performance by using the group comparison approach. A large-scale pilot study was initially conducted to select suitable good and poor performers. The fMRI experiment consisted of a dual-task condition and two single-task conditions. Under the dual-task condition, participants verified the status of letters while concurrently retaining arrow orientations. The behavioral results indicated that accuracy in arrow recognition was better in the good performers than in the poor performers under the dual-task condition but not under the single-task condition. Dual-task performance showed a positive correlation with mean signal change in the right anterior cingulate cortex (ACC) and right dorsolateral prefrontal cortex (DLPFC). Structural equation modeling indicated that effective connectivity between the right ACC and right DLPFC was present in the good performers but not in the poor performers, although activations of the task-dependent posterior regions were modulated by the right ACC and right DLPFC. We conclude that individual differences in attention shifting heavily depend on the functional efficiency of the cingulo-prefrontal network.     

A parietal-frontal network studied by somatosensory oddball MEG responses, and its cross-modal consistency

Previous studies using functional magnetic resonance imaging (fMRI) and event-related potentials (ERPs) of the brain have found that a distributed parietal-frontal neuronal network is activated in normals during both auditory and visual oddball tasks. The common cortical regions in this network are inferior parietal lobule (IPL)/supramarginal gyrus (SMG), anterior cingulate cortex (ACC), and dorsolateral prefrontal cortex (DLPFC). It is not clear whether the same network is activated by oddball tasks during somatosensory stimulation. The present study addressed this question by testing healthy adults as they performed a novel median-nerve oddball paradigm while undergoing magnetoencephalography (MEG). An automated multiple dipole analysis technique, the Multi-Start Spatio-Temporal (MSST) algorithm, localized multiple neuronal generators, and identified their time-courses. IPL/SMG, ACC, and DLPFC were reliably localized in the MEG median-nerve oddball responses, with IPL/SMG activation significantly preceding ACC and DLPFC activation. Thus, the same parietal-frontal neuronal network that shows activation during auditory and visual oddball tests is activated in a median-nerve oddball paradigm. Regions uniquely related to somatosensory oddball responses (e.g., primary and secondary somatosensory, dorsal premotor, primary motor, and supplementary motor areas) were also localized. Since the parietal-frontal network supports attentional allocation during performance of the task, this study may provide a novel method, as well as normative baseline data, for examining attention-related deficits in the somatosensory system of patients with neurological or psychiatric disorders.     

The role of the right anterior prefrontal cortex in episodic retrieval

Regional brain activity was measured with H(2) (15)O PET while participants attempted to complete word-stem and word-fragment retrieval cues with previously studied words. The retrieval cue manipulation was employed to gain control over the monitoring operations associated with evaluating the episodic status of alternative cue completions. These operations were more constrained for fragments, which had fewer possible completions than each corresponding stem. In one condition (zero target), during the scanning interval none of the cues could be completed with studied items, whereas in another condition (high target), 80% of cues belonged to studied items. Relative to baseline tasks, right anterior prefrontal activity was greater for stems than for fragments in the zero target condition. The target density manipulation did not modulate right anterior prefrontal activity, but was associated with increased activity in right dorsolateral prefrontal cortex. These findings are consistent with the proposal that the right anterior prefrontal cortex supports monitoring operations during episodic retrieval tasks. In addition, the findings add to evidence suggesting that the dorsolateral and anterior right prefrontal cortex make functionally distinct contributions to episodic retrieval.     

Neural mechanisms of top-down control during spatial and feature attention

Theories of visual selective attention posit that both spatial location and nonspatial stimulus features (e.g., color) are elementary dimensions on which top-down attentional control mechanisms can selectively influence visual processing. Neuropsychological and neuroimaging studies have demonstrated that regions of superior frontal and parietal cortex are critically involved in the control of visual-spatial attention. This frontoparietal control network has also been found to be activated when attention is oriented to nonspatial stimulus features (e.g., motion). To test the generality of the frontoparietal network in attentional control, we directly compared spatial and nonspatial attention in a cuing paradigm. Event-related fMRI methods permitted the isolation of attentional control activity during orienting to a location or to a nonspatial stimulus feature (color). Portions of the frontoparietal network were commonly activated to the spatial and nonspatial cues. However, direct statistical comparisons of cue-related activity revealed subregions of the frontoparietal network that were significantly more active during spatial than nonspatial orienting when all other stimulus, task, and attentional factors were equated. No regions of the frontal-parietal network were more active for nonspatial cues in comparison to spatial cues. These findings support models suggesting that subregions of the frontal-parietal network are highly specific for controlling spatial selective attention.