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The flash-lag effect (FLE) occurs when a flash's position seems to be delayed relative to a continuously moving object, even though both are physically aligned. Although several studies have demonstrated that reduced attention increases FLE magnitude, the precise mechanism underlying these attention-dependent effects remains elusive. In this study, we investigated the influence of visual attention on the FLE by manipulating the level of attention allocated to multiple stimuli moving simultaneously in different locations. Participants were cued to either focus on one moving stimulus or split their attention among two, three, or four moving stimuli presented in different quadrants. We measured trial-wise FLE to explore potential changes in the magnitude of perceived displacement and its trial-to-trial variability under different attention conditions. Our results reveal that FLE magnitudes were significantly greater when attention was divided among multiple stimuli compared with when attention was focused on a single stimulus, suggesting that divided attention considerably augments the perceptual illusion. However, FLE variability, measured as the coefficient of variation, did not differ between conditions, indicating that the consistency of the illusion is unaffected by divided attention. We discuss the interpretations and implications of our findings in the context of widely accepted explanations of the FLE within a dynamic environment.

Updating beliefs after unexpected events is fundamental for an optimal adaptation to the environment. Previous findings suggested a causal involvement of the right temporoparietal junction (rTPJ) in belief updating in an attention task. We combined offline continuous theta-burst stimulation (cTBS) over rTPJ with functional magnetic resonance imaging (fMRI) to investigate local and remote stimulation effects within the attention and salience networks. In a sham-controlled, within-subject crossover design, 25 participants performed an attentional cueing task during fMRI with true or false information about cue predictability. By estimating learning rates from response times, we characterized participants' belief updating. Model-derived cue predictability entered the fMRI analysis as a parametric regressor to identify the neural correlates of updating. rTPJ-cTBS effects showed high interindividual variability. The expected learning rate reduction with false cue predictability information by cTBS was only observed in participants showing higher updating in false than in true blocks after sham. cTBS modulated the neural signatures of belief updating, both in rTPJ and in nodes of the attention and salience networks. The interindividual variability of the behavioral cTBS effect was related to differential activity and rTPJ connectivity of the right anterior insula. These results demonstrate a crucial interaction between ventral attention and salience networks for belief updating., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Predicting upcoming events using past observations is a crucial component of an efficient allocation of attentional resources. Therefore, the deployment of attention is sensitive to different types of cues predicting upcoming events. Here we investigated probabilistic inference abilities in spatial and feature-based attentional, as well as in motor-intentional subsystems, focusing specifically on the age-related changes in these abilities. In two behavioral experiments, younger and older adults (20 younger and 20 older adults for each experiment) performed three versions of a cueing paradigm, where spatial, feature, or motor cues predicted the location, color, or motor response of a target stimulus. The percentage of cue validity (i.e., the probability of the cue being valid) changed over time, thereby creating a volatile environment. A Bayesian hierarchical model was used to estimate trial-wise beliefs concerning the cue validity from reaction times and to derive a subject-specific belief updating parameter ω in each task version. We also manipulated task difficulty: participants performed an easier version of the task in Experiment 1 and a more difficult version in Experiment 2. Results from Experiment 1 suggested a preserved ability of older adults to use the three different cues to generate predictions. However, the increased task demands of Experiment 2 uncovered a difference in belief updating between the two age groups, indicating moderate evidence for a reduction of the ability to update predictions with motor intention cues in older adults. These results point at a distinction of attentional and motor-intentional subsystems, with age-related differences tackling especially the motor-intentional subsystem.

Although multiple studies characterized the resting-state functional connectivity (rsFC) of the right temporoparietal junction (rTPJ), little is known about the link between rTPJ rsFC and cognitive functions. Given a putative involvement of rTPJ in both reorienting of attention and the updating of probabilistic beliefs, this study characterized the relationship between rsFC of rTPJ with dorsal and ventral attention systems and these two cognitive processes. Twenty-three healthy young participants performed a modified location-cueing paradigm with true and false prior information about the percentage of cue validity to assess belief updating and attentional reorienting. Resting-state fMRI was recorded before and after the task. Seed-based correlation analysis was employed, and correlations of each behavioral parameter with rsFC before the task, as well as with changes in rsFC after the task, were assessed in an ROI-based approach. Weaker rsFC between rTPJ and right intraparietal sulcus before the task was associated with relatively faster updating of the belief that the cue will be valid after false prior information. Moreover, relatively faster belief updating, as well as faster reorienting, were related to an increase in the interhemispheric rsFC between rTPJ and left TPJ after the task. These findings are in line with task-based connectivity studies on related attentional functions and extend results from stroke patients demonstrating the importance of interhemispheric parietal interactions for behavioral performance. The present results not only highlight the essential role of parietal rsFC for attentional functions but also suggest that cognitive processing during a task changes connectivity patterns in a performance-dependent manner.

The visual system forms predictions about upcoming visual features based on previous visual experiences. Such predictions impact on current perception, so that expected stimuli can be detected faster and with higher accuracy. A key question is how these predictions are formed and on which levels of processing they arise. Particularly, predictions could be formed on early levels of processing, where visual features are represented separately, or might require higher levels of processing, with predictions formed based on full object representations that involve combinations of visual features. In four experiments, the present study investigated whether the visual system forms joint prediction errors or whether expectations about different visual features such as color and orientation are formed independently. The first experiment revealed that task-irrelevant and implicitly learned expectations were formed independently when the features were separately bound to different objects. In a second experiment, no evidence for a mutual influence of both types of task-irrelevant and implicitly formed feature expectations was observed, although both visual features were assigned to the same objects. A third experiment confirmed the findings of the previous experiments for explicitly rather than implicitly formed expectations. Finally, no evidence for a mutual influence of different feature expectations was observed when features were assigned to a single centrally presented object. Overall, the present results do not support the view that object feature binding generates joint feature-based expectancies of different object features. Rather, the results suggest that expectations for color and orientation are processed and resolved independently at the feature level.

The deployment of spatial attention is highly sensitive to stimulus predictability. Despite evidence for strong crossmodal links in spatial attentional systems, it remains to be elucidated how concurrent but divergent predictions for targets in different sensory modalities are integrated. In a series of behavioral studies, we investigated the processing of modality-specific expectancies using a multimodal cueing paradigm in which auditory cues predicted the location of visual or tactile targets with modality-specific cue predictability. The cue predictability for visual and tactile targets was manipulated independently. A Bayesian ideal observer model with a weighting factor was applied to trial-wise individual response speed to investigate how the two probabilistic contexts are integrated. Results showed that the degree of integration depended on the level of predictability and on the divergence of the modality-specific probabilistic contexts (Experiments 1-2). However, when the two probabilistic contexts were matched in their level of predictability and were highly divergent (Experiment 3), higher separate processing was favored, especially when visual targets were processed. These findings suggest that modality-specific predictions are flexibly integrated according to their reliability, supporting the hypothesis of separate modality-specific attentional systems that are however linked to guarantee an efficient deployment of spatial attention across the senses.

Predictions about upcoming events influence how we perceive and respond to our environment. There is increasing evidence that predictions may be generated based upon previous observations following Bayesian principles, but little is known about the underlying cortical mechanisms and their specificity for different cognitive subsystems. The present study aimed at identifying common and distinct neural signatures of predictive processing in the spatial attentional and motor intentional system. Twenty-three female and male healthy human volunteers performed two probabilistic cueing tasks with either spatial or motor cues while lying in the fMRI scanner. In these tasks, the percentage of cue validity changed unpredictably over time. Trialwise estimates of cue predictability were derived from a Bayesian observer model of behavioral responses. These estimates were included as parametric regressors for analyzing the BOLD time series. Parametric effects of cue predictability in valid and invalid trials were considered to reflect belief updating by precision-weighted prediction errors. The brain areas exhibiting predictability-dependent effects dissociated between the spatial attention and motor intention task, with the right temporoparietal cortex being involved during spatial attention and the left angular gyrus and anterior cingulate cortex during motor intention. Connectivity analyses revealed that all three areas showed predictability-dependent coupling with the right hippocampus. These results suggest that precision-weighted prediction errors of stimulus locations and motor responses are encoded in distinct brain regions, but that crosstalk with the hippocampus may be necessary to integrate new trialwise outcomes in both cognitive systems. SIGNIFICANCE STATEMENT The brain is able to infer the environments' statistical structure and responds strongly to expectancy violations. In the spatial attentional domain, it has been shown that parts of the attentional networks are sensitive to the predictability of stimuli. It remains unknown, however, whether these effects are ubiquitous or if they are specific for different cognitive systems. The present study compared the influence of model-derived cue predictability on brain activity in the spatial attentional and motor intentional system. We identified areas with distinct predictability-dependent activation for spatial attention and motor intention, but also common connectivity changes of these regions with the hippocampus. These findings provide novel insights into the generality and specificity of predictive processing signatures in the human brain., (Copyright © 2017 the authors 0270-6474/17/375334-11$15.00/0.)

Humans flexibly attend to features or locations and these processes are influenced by the probability of sensory events. We combined computational modeling of response times with fMRI to compare the functional correlates of (re-)orienting, and the modulation by probabilistic inference in spatial and feature-based attention systems. Twenty-four volunteers performed two task versions with spatial or color cues. Percentage of cue validity changed unpredictably. A hierarchical Bayesian model was used to derive trial-wise estimates of probability-dependent attention, entering the fMRI analysis as parametric regressors. Attentional orienting activated a dorsal frontoparietal network in both tasks, without significant parametric modulation. Spatially invalid trials activated a bilateral frontoparietal network and the precuneus, while invalid feature trials activated the left intraparietal sulcus (IPS). Probability-dependent attention modulated activity in the precuneus, left posterior IPS, middle occipital gyrus, and right temporoparietal junction for spatial attention, and in the left anterior IPS for feature-based and spatial attention. These findings provide novel insights into the generality and specificity of the functional basis of attentional control. They suggest that probabilistic inference can distinctively affect each attentional subsystem, but that there is an overlap in the left IPS, which responds to both spatial and feature-based expectancy violations., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Stroke can result in marked impairments in the processing of information presented in contralesional space. The present prospective study investigated how a contralesional distractor affects ipsilesional perception in patients with a right-hemispheric stroke. In a simple target detection task, the influence of the distractor on reaction times (RTs) was examined in stroke patients and compared to the performance of healthy elderly controls. Distractor interference effects were related to measures of neglect and extinction using a regression analysis. Moreover, the magnitude of the behavioural distractor effect entered a voxel-based lesion-symptom mapping (VLSM) analysis with the hypothesis that parietal and temporoparietal lesions are related to altered distractor processing. While the presence of a distractor in the opposite hemifield slowed down RTs in healthy controls for left and right targets, this effect was only observed for contralesional left targets in the group of right-hemispheric patients. In stark contrast, the presence of a distractor in the contralesional hemifield expedited ipsilesional (i.e., right) target detection. This effect was significantly related to lesions in the anterior middle temporal and temporoparietal cortex, external and internal capsule, as well as the superior longitudinal fascicle (SLF). These results elucidate the nature of the disruption of attentive processing in the contralesional hemifield after right-hemispheric stroke. More specifically, they shed light on the abnormal prioritisation of ipsilesional information: our data suggest that damage to the temporal and temporoparietal cortex and white matter tracts may transform contralesional stimulation into an unspecific saliency signal contributing to facilitated information processing in ipsilesional space., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Modelling psychophysical data using the Theory of Visual Attention (TVA) allows for a quantification of attentional sub-processes, such as the resolution of competition amongst multiple stimuli by top-down control signals for target selection (TVA-parameter α). This fMRI study investigated the neural correlates of α by comparing activity differences and changes of effective connectivity between conditions where a target was accompanied by a distractor or by a second target. Twenty-five participants performed a partial report task inside the MRI scanner. The left angular gyrus (ANG), medial frontal, and posterior cingulate cortex showed higher activity when a target was accompanied by a distractor as opposed to a second target. The reverse contrast yielded activation of a bilateral fronto-parietal network, the anterior insula, anterior cingulate cortex, and left inferior occipital gyrus. A psychophysiological interaction analysis revealed that the connectivity between left ANG and the left and right supramarginal gyrus (SMG), left anterior insula, and right putamen was enhanced in the target-distractor condition in participants with worse attentional top-down control. Dynamic causal modelling suggested that the connection from left ANG to right SMG during distractor presence was modulated by α. Our data show that interindividual differences in attentional processing are reflected in changes of effective connectivity without significant differences in activation strength of network nodes., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Allocation of attentional resources rests on predictions about the likelihood of events. While this effect has been extensively studied in the spatial attention domain where the location of a target stimulus is pre-cued, less is known about the cueing of stimulus features such as the color of a behaviorally relevant target. Moreover, there is disagreement about which types of color cues are effective for biasing attention. Here we investigated the effects of probabilistic context (percentage of cue validity, %CV) for different levels of cue abstraction to elucidate how feature-based search information is processed and used to direct attention. The color of a target was cued by presenting the perceptual color, the color word, or two-letter abbreviations. %CV, i.e., the probability that the cue indicated the color correctly, changed unpredictably between 50, 70, and 90%. Response times (RTs) for valid and invalid trials in each %CV condition were recorded in 60 datasets and analyzed with analyses of variance. The results showed that all cues were associated with comparable RT costs after invalid cueing. The modulation of RT costs by probabilities, however, depended upon level of cue abstraction and time on task: While a strong, immediate impact of %CV was found for two-letter cueing, the effect was solely observed in the second half of the experiment for perceptual and word cues. These results demonstrate that probabilistic feature-based information is processed differently for different levels of cue abstraction. Moreover, the modulatory effect of the environmental statistics differentially depends on the time on task for different feature cues.

The deployment of visuospatial attention and the programming of saccades are governed by the inferred likelihood of events. In the present study, we combined computational modeling of psychophysical data with fMRI to characterize the computational and neural mechanisms underlying this flexible attentional control. Sixteen healthy human subjects performed a modified version of Posner's location-cueing paradigm in which the percentage of cue validity varied in time and the targets required saccadic responses. Trialwise estimates of the certainty (precision) of the prediction that the target would appear at the cued location were derived from a hierarchical Bayesian model fitted to individual trialwise saccadic response speeds. Trial-specific model parameters then entered analyses of fMRI data as parametric regressors. Moreover, dynamic causal modeling (DCM) was performed to identify the most likely functional architecture of the attentional reorienting network and its modulation by (Bayes-optimal) precision-dependent attention. While the frontal eye fields (FEFs), intraparietal sulcus, and temporoparietal junction (TPJ) of both hemispheres showed higher activity on invalid relative to valid trials, reorienting responses in right FEF, TPJ, and the putamen were significantly modulated by precision-dependent attention. Our DCM results suggested that the precision of predictability underlies the attentional modulation of the coupling of TPJ with FEF and the putamen. Our results shed new light on the computational architecture and neuronal network dynamics underlying the context-sensitive deployment of visuospatial attention., Significance Statement: Spatial attention and its neural correlates in the human brain have been studied extensively with the help of fMRI and cueing paradigms in which the location of targets is pre-cued on a trial-by-trial basis. One aspect that has so far been neglected concerns the question of how the brain forms attentional expectancies when no a priori probability information is available but needs to be inferred from observations. This study elucidates the computational and neural mechanisms under which probabilistic inference governs attentional deployment. Our results show that Bayesian belief updating explains changes in cortical connectivity; in that directional influences from the temporoparietal junction on the frontal eye fields and the putamen were modulated by (Bayes-optimal) updates., (Copyright © 2015 Vossel et al.)

The exact mechanisms whereby the cholinergic neurotransmitter system contributes to attentional processing remain poorly understood. Here, we applied computational modeling to psychophysical data (obtained from a spatial attention task) under a psychopharmacological challenge with the cholinesterase inhibitor galantamine (Reminyl). This allowed us to characterize the cholinergic modulation of selective attention formally, in terms of hierarchical Bayesian inference. In a placebo-controlled, within-subject, crossover design, 16 healthy human subjects performed a modified version of Posner's location-cueing task in which the proportion of validly and invalidly cued targets (percentage of cue validity, % CV) changed over time. Saccadic response speeds were used to estimate the parameters of a hierarchical Bayesian model to test whether cholinergic stimulation affected the trial-wise updating of probabilistic beliefs that underlie the allocation of attention or whether galantamine changed the mapping from those beliefs to subsequent eye movements. Behaviorally, galantamine led to a greater influence of probabilistic context (% CV) on response speed than placebo. Crucially, computational modeling suggested this effect was due to an increase in the rate of belief updating about cue validity (as opposed to the increased sensitivity of behavioral responses to those beliefs). We discuss these findings with respect to cholinergic effects on hierarchical cortical processing and in relation to the encoding of expected uncertainty or precision., (Copyright © 2014 the authors 0270-6474/14/3415735-08$15.00/0.)

Inferring the environment's statistical structure and adapting behavior accordingly is a fundamental modus operandi of the brain. A simple form of this faculty based on spatial attentional orienting can be studied with Posner's location-cueing paradigm in which a cue indicates the target location with a known probability. The present study focuses on a more complex version of this task, where probabilistic context (percentage of cue validity) changes unpredictably over time, thereby creating a volatile environment. Saccadic response speed (RS) was recorded in 15 subjects and used to estimate subject-specific parameters of a Bayesian learning scheme modeling the subjects' trial-by-trial updates of beliefs. Different response models-specifying how computational states translate into observable behavior-were compared using Bayesian model selection. Saccadic RS was most plausibly explained as a function of the precision of the belief about the causes of sensory input. This finding is in accordance with current Bayesian theories of brain function, and specifically with the proposal that spatial attention is mediated by a precision-dependent gain modulation of sensory input. Our results provide empirical support for precision-dependent changes in beliefs about saccade target locations and motivate future neuroimaging and neuropharmacological studies of how Bayesian inference may determine spatial attention.

The idea of two separate attention networks in the human brain for the voluntary deployment of attention and the reorientation to unexpected events, respectively, has inspired an enormous amount of research over the past years. In this review, we will reconcile these theoretical ideas on the dorsal and ventral attentional system with recent empirical findings from human neuroimaging experiments and studies in stroke patients. We will highlight how novel methods-such as the analysis of effective connectivity or the combination of neurostimulation with functional magnetic resonance imaging-have contributed to our understanding of the functionality and interaction of the two systems. We conclude that neither of the two networks controls attentional processes in isolation and that the flexible interaction between both systems enables the dynamic control of attention in relation to top-down goals and bottom-up sensory stimulation. We discuss which brain regions potentially govern this interaction according to current task demands.

The right temporo-parietal junction (TPJ) is widely considered as part of a network that reorients attention to task-relevant, but currently unattended stimuli (Corbetta and Shulman, 2002). Despite the prevalence of this theory in cognitive neuroscience, there is little direct evidence for the principal hypothesis that TPJ sends an early reorientation signal that "circuit breaks" attentional processing in regions of the dorsal attentional network (e.g., the frontal eye fields) or is completely right lateralized during attentional processing. In this review, we examine both functional neuroimaging work on TPJ in the attentional literature as well as anatomical findings. We first critically evaluate the idea that TPJ reorients attention and is right lateralized; we then suggest that TPJ signals might rather reflect post-perceptual processes involved in contextual updating and adjustments of top-down expectations; and then finally discuss how these ideas relate to the electrophysiological (P300) literature, and to TPJ findings in other cognitive and social domains. We conclude that while much work is needed to define the computational functions of regions encapsulated as TPJ, there is now substantial evidence that it is not specialized for stimulus-driven attentional reorienting., (Copyright © 2013 The Authors. Published by Elsevier Ltd.. All rights reserved.)

The right intraparietal sulcus (rIPS) is a key region for the endogenous control of selective visual attention in the human brain. Previous studies suggest that the rIPS is especially involved in top-down control and spatial distribution of attention across both visual hemifields. We further explored these attentional functions using transcranial direct current stimulation (tDCS) of the rIPS to modulate behavioral performance in a partial report task. Performance was analyzed according to the theory of visual attention (TVA) (Bundesen, 1990), which provides a computational framework to investigate different parameters of visuo-attentional processing such as top-down control, attentional weighting, capacity of visual short term memory, and processing speed. We investigated the effects of different tDCS current strengths (1 mA and 2 mA) in two experiments: 1 mA tDCS (anodal, cathodal, sham) did not affect any of the TVA parameters, but cathodal 2 mA stimulation significantly enhanced top-down control as evidenced by a reduction of the α parameter of TVA, regardless of hemifield. This differential impact on the top-down control component of attentional processing suggests that the horizontal rIPS is mainly involved in attentional selection as none of the spatial or resource variables of TVA were altered. Furthermore, the data add evidence to previous work highlighting (1) the importance of using appropriate current strength in stimulation protocols, and (2) that the often reported inhibitory effect of cathodal stimulation in e.g., motor tasks might not extend to cognitive paradigms.

How the human brain reconstructs the three-dimensional (3D) world from two-dimensional (2D) retinal images has received a great deal of interest as has how we shift attention in 2D space. In contrast, it remains poorly understood how visuospatial attention is shifted in depth. In this fMRI study, by constructing a virtual 3D environment in the MR scanner and by presenting targets either close to or far from the participants in an adapted version of the Posner spatial-cueing paradigm, we investigated the behavioral and neural mechanisms underlying visuospatial orienting/reorienting in depth. At the behavioral level, although covering the same spatial distance, attentional reorienting to objects unexpectedly appearing closer to the observer and in the unattended hemispace was faster than reorienting to unexpected objects farther away. At the neural level, we found that in addition to the classical attentional reorienting system in the right temporoparietal junction, two additional brain networks were differentially involved in aspects of attentional reorienting in depth. First, bilateral premotor cortex reoriented visuospatial attention specifically along the third dimension of visual space (i.e., from close to far or vice versa), compared with attentional reorienting within the same depth plane. Second, a network of areas reminiscent of the human "default-mode network," including posterior cingulate cortex, orbital prefrontal cortex, and left angular gyrus, was involved in the neural interaction between depth and attentional orienting, by boosting attentional reorienting to unexpected objects appearing both closer to the observer and in the unattended hemispace.

Attentional orientation to a spatial cue and reorientation-after invalid cueing-are mediated by two distinct networks in the human brain. A bilateral dorsal frontoparietal network, comprising the intraparietal sulcus (IPS) and the frontal eye fields (FEF), controls the voluntary deployment of attention and may modulate visual cortex in preparation for upcoming stimulation. In contrast, reorienting attention to invalidly cued targets engages a right-lateralized ventral frontoparietal network comprising the temporoparietal junction (TPJ) and ventral frontal cortex. The present fMRI study investigated the functional architecture of these two attentional systems by characterizing effective connectivity during lateralized orienting and reorienting of attention, respectively. Subjects performed a modified version of Posner's location-cueing paradigm. Dynamic causal modeling (DCM) of regional responses in the dorsal and ventral network, identified in a conventional (SPM) whole-brain analysis, was used to compare different functional architectures. Bayesian model selection showed that top-down connections from left and right IPS to left and right visual cortex, respectively, were modulated by the direction of attention. Moreover, model evidence was highest for a model with directed influences from bilateral IPS to FEF, and reciprocal coupling between right and left FEF. Invalid cueing enhanced forward connections from visual areas to right TPJ, and directed influences from right TPJ to right IPS and IFG (inferior frontal gyrus). These findings shed further light on the functional organization of the dorsal and ventral attentional network and support a context-sensitive lateralization in the top-down (backward) mediation of attentional orienting and the bottom-up (forward) effects of invalid cueing.

Nicotine enhances attentional functions. Since chronic nicotine exposure through smoking induces neuroadaptive changes in the brain at a structural and molecular level, the present functional MRI (fMRI) study aimed at investigating the neural mechanisms underlying visuospatial and sustained attention in smokers and non-smokers. Visuospatial attention was assessed with a location-cueing paradigm, while sustained attention was measured by changes in response speed over time. During invalid trials, neural activity within the basal forebrain was selectively enhanced in smokers and higher basal forebrain activity was associated with increased parietal cortex activation. Moreover, higher levels of expired carbon monoxide in smokers before scanning were associated with higher parietal cortex activation and faster responses to invalidly cued targets. Smokers showed a slowing of responses and additionally recruited an area within the right supramarginal gyrus with increasing time on task. Activity decreases over time were observed in visual areas in smokers. The data provide evidence for altered attentional functions in smokers as compared with non-smokers, which were partly modulated by residual nicotine levels and were observed at a behavioural level for sustained and at a neural level for spatial and sustained attention.

Besides the fact that RTs in cognitive tasks are affected by the specific demands of a trial, the context in which this trial occurs codetermines the speed of the response. For instance, invalid spatial cues generally prolong RTs to targets in the location-cueing paradigm, whereas the magnitude of these RT costs additionally varies as a function of the preceding trial types so that RTs for invalid trials may be increased when preceded by valid rather than invalid trials. In the present fMRI study, we investigated trial sequence effects in a combined oddball and location-cueing paradigm. In particular, we tested whether RTs and neural activity to infrequent invalid or deviant targets varied as a function of the number of preceding valid standard trials. As expected, RTs in invalid and deviant trials were significantly slower when more valid standard trials had been presented beforehand. This behavioral effect was reflected in the neural activity of the right inferior/middle frontal gyrus where the amplitude of the hemodynamic response in invalid and deviant trials was positively related to the number of preceding valid standard trials. In contrast, decreased activity (i.e., a negative parametric modulation effect) was observed when more valid standard trials were successively presented. Further positive parametric effects for the number of preceding valid standard trials were observed in the left caudate nucleus and lingual gyrus. The data suggest that inferior frontal cortex extracts both event regularities and irregularities in event streams.

Spatial neglect as a multifaceted syndrome may consist of perceptual/attentional as well as motor/intentional components. The present study investigated the lesion anatomy underlying perceptual and response bias using a manual response Landmark task (Bisiach, Ricci, Lualdi, & Colombo, 1998) in 68 patients with right-hemispheric stroke. The two differential aspects of the neglect syndrome were assessed by measuring response tendencies resulting from underestimations of the length of left line segments and from hypometric movement execution towards contralesional space, respectively. Perceptual and response bias were orthogonal components of task performance in the Landmark task. Perceptual as well as response bias both explained variance in the performance of standard paper-and-pencil neglect tests. While lesions within middle frontal, inferior parietal and parieto-occipital brain regions were related to perceptual bias, subcortical lesions within the caudate were related to response bias in the Landmark task. Our data suggest that perceptual/attentional and motor/intentional aspects of neglect are independent components of the syndrome with differential neural underpinnings in fronto-parietal and subcortical brain regions., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

The deficit to reorient attention from ipsilesional to contralesional space is one key feature of the spatial neglect syndrome. As previous studies suggest that reorienting of visuospatial attention is modulated by cholinergic neurotransmission, we investigated whether cholinergic stimulation with nicotine (Nicorette 2 mg, Pharmacia/Pfizer, Helsingborg, Sweden) facilitates attentional reorienting in spatial neglect patients. Nine nonsmoking patients with stable neglect symptoms were investigated in a within-subject cross-over design. We used a location-cueing paradigm and analysed reaction time (RT) differences between validly and invalidly cued, as well as between neutrally cued and uncued targets as a function of hemifield and drug. Moreover, since the nicotine effect is mediated by parietal brain areas in healthy subjects, we tested whether lesion location influences the pharmacological effect. Nicotine speeded RTs in valid and invalid trials nonspecifically, without modulating the validity effect in the location-cueing task in the whole group of patients. Lesion-symptom mapping revealed a relationship between lesion site and the pharmacological effect on reorienting to contralesional space in right parietal and temporal brain regions. We conclude that in patients with chronic spatial neglect the performance in the location-cueing paradigm can be modulated by a cholinergic stimulant provided that the lesion spares right parietal and temporal cortex.

Within the parietal cortex, the temporo-parietal junction (TPJ) and the intraparietal sulcus (IPS) seem to be involved in both spatial and nonspatial functions: Both areas are activated when misleading information is provided by invalid spatial cues in Posner's location-cueing paradigm, but also when infrequent deviant stimuli are presented within a series of standard events. In the present study, we used functional magnetic resonance imaging to investigate the distinct and shared brain responses to (i) invalidly cued targets requiring attentional reorienting, and (ii) to target stimuli deviating in color and orientation leading to an oddball-like distraction effect. Both unexpected location and feature changes were accompanied by a significant slowing of manual reaction times. Bilateral TPJ and right superior parietal lobe (SPL) activation was observed in response to invalidly as compared to validly cued targets. In contrast, the bilateral inferior occipito-temporal cortex, the left inferior parietal cortex, right frontal areas, and the cerebellum showed stronger activation in response to deviant than to standard targets. Common activations were observed in the right angular gyrus along the IPS and in the right inferior frontal gyrus. We conclude that the superior parietal and temporo-parietal activations observed here as well as previously in location-cueing paradigms do not merely reflect the detection and processing of unexpected stimuli. Furthermore, our data suggest that the right IPS and the inferior frontal gyrus are involved in attentional selection and distractor processing of both spatial and nonspatial features.

The cholinergic neurotransmitter system has been proposed to be involved in the processing of probabilistic top-down information provided by endogenous cues in location-cueing paradigms. It has been shown that the behavioral and neural effects of a nicotinic cholinergic stimulation resemble the effects obtained by manipulating the validity of the spatial cues: enhancing cortical nicotine levels and decreasing cue validity both reduce the reaction time difference between invalidly and validly cued targets (ie, the 'validity effect') as well as neural activity related to attentional reorienting in parietal brain regions. In the present study, we investigated whether the behavioral and neural effects of nicotine in location-cueing paradigms are dependent upon different a priori cue validities. Twenty-four subjects were investigated in a double-blind placebo-controlled between-subject design with functional magnetic resonance imaging. Nicotine was administered to non-smoking volunteers via polacrilex gums (Nicorette, 2 mg) before performing a location-cueing paradigm with valid and invalid cues in the context of 90 and 60% cue validity in the MR scanner. Nicotine significantly reduced the validity effect in the 90% but not in the 60% cue validity condition. Fronto-parietal and cingulate regions showed stronger nicotinic reductions of reorienting-related neural activity in the high than in the low cue validity condition. Our data reveal an interaction effect between the pharmacological and cognitive modulation of attentional reorienting, which is evident at both a behavioral as well as the neuronal level.

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.