The parietal cortex and attentional modulations of activities of the visual cortex

We recorded high density event-related brain potentials (ERPs) from a patient with focal left parietal damage in a covert visual orienting task requiring detection of targets in the attended or unattended hemifield. A positivity peaking at 120 ms (P1) to the left visual field stimuli was enlarged when attended than unattended and was localized to the right extrastirate cortex. However, spatial attention did not influence the ERPs to the right visual field stimuli. The leftward cue elicited an enlarged P1 relative to the rightward cue. The results suggest that human parietal cortex is critical for the attentional modulation of the neural activities in the extrastriate cortex associated with stimuli in the contralateral hemifield.     

Modulation of spatial attention by fear-conditioned stimuli: an event-related fMRI study

Stimuli that signal threat can capture subjects' attention, leading to more efficient detection of, and faster responses to, events occurring in that part of the environment. In the present study we explored the behavioural and anatomical correlates of the modulation of spatial attention by emotion using a fear conditioning paradigm, combined with a covert spatial orienting task. Reaction times for the detection of a peripheral target, which was preceded by brief (50ms) presentations of the visual conditioned stimulus (CS+) in either the same or opposite visual field, showed an interaction between stimulus emotionality and attention shifts. We used event-related functional magnetic resonance imaging (fMRI) to characterise the associated neural responses. Consistent with previous studies, conditioning-induced enhanced responses were observed in the amygdala and extrastriate visual cortex. The modulation of spatial attention by a conditioned stimulus was associated with enhanced activity in regions of frontal and parietal cortices previously implicated in spatial attention, as well as in the lateral orbitofrontal cortex (lOFC).     

Orienting attention in time: behavioural and neuroanatomical distinction between exogenous and endogenous shifts

Temporal orienting of attention is the ability to focus resources at a particular moment in time in order to optimise behaviour, and is associated with activation of left parietal and premotor cortex [Coull, J. T., Nobre, A. C. Where and when to pay attention: the neural systems for directing attention to spatial locations and to time intervals as revealed by both PET and fMRI. Journal of Neuroscience, 1998, 18, 7426-7435]. In the present experiment, we explored the behavioural and anatomical correlates of temporal orienting to foveal visual stimuli, in order to eliminate any spatial attention confounds. We implemented a two-way factorial design in an event-related fMRI study to examine the factors of trial validity (predictability of target by cue), length of delay (cue-target interval), and their interaction. There were two distinct types of invalid trial: those where attention was automatically drawn to a premature target and those where attention was voluntarily shifted to a delayed time-point. Reaction times for valid trials were shorter than those for invalid trials, demonstrating appropriate allocation of attention to temporal cues. All trial-types activated a shared system, including frontoparietal areas bilaterally, showing that this network is consistently associated with attentional orienting and is not specific to spatial tasks. Distinct brain areas were sensitive to cue-target delays and to trial validity. Long cue-target intervals activated areas involved in motor preparation: supplementary motor cortex, basal ganglia and thalamus. Invalid trials, where temporal expectancies were breached, showed enhanced activation of left parietal and frontal areas, and engagement of orbitofrontal cortex bilaterally. Finally, trial validity interacted with length of delay. Appearance of targets prematurely selectively activated visual extrastriate cortex; while postponement of target appearance selectively activated right prefrontal cortex. These findings suggest that distinct brain areas are involved in redirecting attention based upon sensory events (bottom-up, exogenous shifts) and based upon cognitive expectations (top-down, endogenous shifts).     

Orienting auditory spatial attention engages frontal eye fields and medial occipital cortex in congenitally blind humans

What happens in vision-related cortical areas when congenitally blind (CB) individuals orient attention to spatial locations? Previous neuroimaging of sighted individuals has found overlapping activation in a network of frontoparietal areas including frontal eye fields (FEF), during both overt (with eye movement) and covert (without eye movement) shifts of spatial attention. Since voluntary eye movement planning seems irrelevant in CB, their FEF neurons should be recruited for alternative functions if their attentional role in sighted individuals is only due to eye movement planning. Recent neuroimaging of the blind has also reported activation in medial occipital areas, normally associated with visual processing, during a diverse set of non-visual tasks, but their response to attentional shifts remains poorly understood. Here, we used event-related fMRI to explore FEF and medial occipital areas in CB individuals and sighted controls with eyes closed (SC) performing a covert attention orienting task with endogenous verbal cues and spatialized auditory targets. We found robust stimulus-locked FEF activation of all CB subjects, similar to and stronger than in SC, suggesting that FEF plays a role in endogenous orienting of covert spatial attention even in individuals in whom voluntary eye movements are irrelevant. We also found robust activation in bilateral medial occipital cortex in CB but not in SC subjects. The response decreased below baseline following endogenous verbal cues but increased following auditory targets, suggesting that the medial occipital area in CB does not directly engage during cued orienting of attention but may be recruited for processing of spatialized auditory targets.     

Neural systems for visual orienting and their relationships to spatial working memory

We investigated neural correlates of human visual orienting using event-related functional magnetic resonance imaging (fMRI). When subjects voluntarily directed attention to a peripheral location, we recorded robust and sustained signals uniquely from the intraparietal sulcus (IPs) and superior frontal cortex (near the frontal eye field, FEF). In the ventral IPs and FEF only, the blood oxygen level dependent signal was modulated by the direction of attention. The IPs and FEF also maintained the most sustained level of activation during a 7-sec delay, when subjects maintained attention at the peripheral cued location (working memory). Therefore, the IPs and FEF form a dorsal network that controls the endogenous allocation and maintenance of visuospatial attention. A separate right hemisphere network was activated by the detection of targets at unattended locations. Activation was largely independent of the target's location (visual field). This network included among other regions the right temporo-parietal junction and the inferior frontal gyrus. We propose that this cortical network is important for reorienting to sensory events.     

Slow fluctuations in eye position and resting‐state functional magnetic resonance imaging brain activity during visual fixation

The neuronal circuitry that supports voluntary changes in eye position in tasks that require attention‐driven oculo‐motor control is well known. However, less is known about the neuronal basis for eye control during visual fixation. This, together with the fact that visual fixation is one of the most commonly used baseline conditions in resting‐state functional magnetic resonance imaging (fMRI) studies, prompted us to conduct a study in which we employed resting‐state fMRI and concurrent recordings of eye gaze to investigate the relationship between spontaneous changes in eye position during passive visual fixation and intrinsic brain activity. As a control experiment, we recorded fMRI brain activity related to cued voluntary vertical and horizontal changes in eye position in a block‐related task‐evoked fMRI experiment. Our results for the voluntarily performed changes in eye position elicited brain activity in the bilateral occipitotemporal cortex, supplementary motor cortex and frontal eye fields. In contrast, we show that slow fluctuations in eye position during passive visual fixation are linked to intrinsic brain activity, foremost in midline cortical brain regions located in the posteromedial parietal cortex and the medial prefrontal cortex, brain regions that act as core cortical hubs in the brain's default mode network. Our results suggest that subconscious and sustained changes in behavior are tied to intrinsic brain activity on a moment‐by‐moment basis.     

Attentional reorientation along the meridians of the visual field: Are there different neural mechanisms at play?

Hemispatial neglect, after unilateral lesions to parietal brain areas, is characterized by an inability to respond to unexpected stimuli in contralesional space. As the visual field's horizontal meridian is most severely affected, the brain networks controlling visuospatial processes might be tuned explicitly to this axis. We investigated such a potential directional tuning in the dorsal and ventral frontoparietal attention networks, with a particular focus on attentional reorientation. We used an orientation‐discrimination task where a spatial precue indicated the target position with 80% validity. Healthy participants (n = 29) performed this task in two runs and were required to (re‐)orient attention either only along the horizontal or the vertical meridian, while fMRI and behavioral measures were recorded. By using a general linear model for behavioral and fMRI data, dynamic causal modeling for effective connectivity, and other predictive approaches, we found strong statistical evidence for a reorientation effect for horizontal and vertical runs. However, neither neural nor behavioral measures differed between vertical and horizontal reorienting. Moreover, models from one run successfully predicted the cueing condition in the respective other run. Our results suggest that activations in the dorsal and ventral attention networks represent higher‐order cognitive processes related to spatial attentional (re‐)orientating that are independent of directional tuning and that unilateral attention deficits after brain damage are based on disrupted interactions between higher‐level attention networks and sensory areas.     

Deconstructing the architecture of dorsal and ventral attention systems with dynamic causal modeling

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.     

Retinotopic effects during spatial audio-visual integration

The successful integration of visual and auditory stimuli requires information about whether visual and auditory signals originate from corresponding places in the external world. Here we report crossmodal effects of spatially congruent and incongruent audio-visual (AV) stimulation. Visual and auditory stimuli were presented from one of four horizontal locations in external space. Seven healthy human subjects had to assess the spatial fit of a visual stimulus (i.e. a gray-scaled picture of a cartoon dog) and a simultaneously presented auditory stimulus (i.e. a barking sound). Functional magnetic resonance imaging (fMRI) revealed two distinct networks of cortical regions that processed preferentially either spatially congruent or spatially incongruent AV stimuli. Whereas earlier visual areas responded preferentially to incongruent AV stimulation, higher visual areas of the temporal and parietal cortex (left inferior temporal gyrus [ITG], right posterior superior temporal gyrus/sulcus [pSTG/STS], left intra-parietal sulcus [IPS]) and frontal regions (left pre-central gyrus [PreCG], left dorsolateral pre-frontal cortex [DLPFC]) responded preferentially to congruent AV stimulation. A position-resolved analysis revealed three robust cortical representations for each of the four visual stimulus locations in retinotopic visual regions corresponding to the representation of the horizontal meridian in area V1 and at the dorsal and ventral borders between areas V2 and V3. While these regions of interest (ROIs) did not show any significant effect of spatial congruency, we found subregions within ROIs in the right hemisphere that showed an incongruency effect (i.e. an increased fMRI signal during spatially incongruent compared to congruent AV stimulation). We interpret this finding as a correlate of spatially distributed recurrent feedback during mismatch processing: whenever a spatial mismatch is detected in multisensory regions (such as the IPS), processing resources are re-directed to low-level visual areas.     

Auditory and visual attention assessed with PET

Brain mechanisms involved in the maintenance of attention to auditory and visual stimuli at different spatial locations were assessed using positron emission tomography with [¹⁵O]water to measure regional cerebral blood flow (rCBF) changes in 13 normal volunteers. Simultaneous auditory [dichotically presented consonant-vowel-consonants (CVCs)] and visual stimuli (vertically oriented, CVCs presented to the left and right of fixation) were presented on every trial. In different conditions subjects attended for targets in a specified stimulus channel (left or right ears or left or right visual fields) while maintaining fixation on a central x. Attending left or right for auditory stimuli increased rCBF in primary auditory cortex in Heschl's gyrus and in temporal lobe auditory association cortices in both hemispheres. Attending left or right for visual stimuli did not change rCBF in primary visual cortex, and only attention to the right significantly increased rCBF in contralateral occipital cortex. Visual attention caused significant rCBF changes in a widespread network that included frontal, parietal, and temporal cortical regions as well as the cerebellum, whereas rCBF changes due to auditory attention were largely localized in the temporal lobes. The results suggest that spatially directed attention is mediated by different mechanisms in the auditory and visual modalities. Hum. Brain Mapping 5:422–436, 1997. © 1997 Wiley-Liss, Inc.     

Covert reorienting and inhibition of return: an event-related fMRI study

Using event-related fMRI, we analyzed the functional neuroanatomy of covert reorienting and inhibition of return (IOR). Covert reorienting to a target appearing within 250 msec after an invalid contralateral location cue elicited increased activation in the left fronto-polar cortex (LFPC), right anterior and left posterior middle frontal gyrus, and right cerebellum, areas that have previously been associated with attentional processes, specifically attentional change. In contrast, IOR, which leads to prolonged response times to targets that appear at the cued location at a stimulus-onset-asynchrony (SOA)>250 msec, was accompanied by increased activation in brain areas involved in oculomotor programming, such as the right medial frontal gyrus (supplementary eye field; SEF) and the right inferior precentral sulcus (frontal eye field; FEF), supporting the oculomotor bias theory of IOR. Pre-SEF and pre-FEF areas were involved both in covert reorienting and IOR. The supramarginal gyri were bilaterally involved in IOR, with the right supramarginal gyrus additionally involved in covert reorienting.     

Neural substrates differentiating global/local processing of bilateral visual inputs

We investigated neural substrates of global/local processing of bilateral hierarchical stimuli using functional magnetic resonance imaging (fMRI). Subjects were presented with two compound letters that were displayed simultaneously in the left and right visual fields, respectively. In a steady-state, block-design paradigm, hemodynamic responses were recorded while subjects detected infrequent global or local targets presented in one hemifield in separate epochs of trials. While behavioural responses were more accurate and faster to global than local targets, attention to the global level of bilateral visual inputs induced stronger activations in the left and right temporal cortex relative to attention to the local level. However, attention to the local level generated stronger activations in bilateral superior parietal cortex compared with attention to the global level. The results suggest that distinct neural substrates in the temporal and parietal cortices are preferentially engaged in the global and local processing of bilateral visual inputs, respectively.     

Neural correlates of crossmodal visual-tactile extinction and of tactile awareness revealed by fMRI in a right-hemisphere stroke patient

We used fMRI to study the neural correlates of crossmodal, visual-tactile extinction in a single case (patient GK). GK has chronic extinction after a lesion centred on right inferior parietal cortex, and has previously been investigated extensively in purely visual fMRI studies [e.g. Rees, G., Wojciulik, E., Clarke, K., Husain, M., Frith, C., & Driver, J. (2000). Unconscious activation of visual cortex in the damaged right hemisphere of a parietal patient with extinction. Brain, 123(Pt 8), 1624-1633; Rees, G., Kreiman, G., & Koch, C. (2002). Neural correlates of consciousness in humans. Nature Reviews Neuroscience, 3(4), 261-270]. With concurrent stimulation of the right visual field plus left index finger, GK showed crossmodal extinction of left touch on approximately half of such trials here (reflecting impaired sensitivity, i.e. lowered d-prime), albeit becoming aware of left touch on the other half. fMRI revealed activation of contralateral primary somatosensory cortex on crossmodal trials when touch was extinguished from awareness, suggesting unconscious residual processing there. When GK became aware of the left touch, additional activation was found in surviving right parietal cortex, and in frontal regions; moreover, functional coupling was enhanced with a region of frontal cortex implicated in awareness by previous work. Finally, on trials where crossmodal extinction arose, surviving right parietal cortex showed stronger functional coupling with the left visual and right somatosensory regions driven by the competing stimuli, indicating that crossmodal extinction arises when inputs to separate modalities interact competitively via multimodal cortex.     

Attention orienting dysfunction during salient novel stimulus processing in schizophrenia

Schizophrenia is characterised by marked disturbances of attention and information processing. Patients experience difficulty focusing on relevant cues and avoiding distraction by irrelevant stimuli. Event-related potential recordings indicate an amplitude reduction in the P3a component elicited by involuntary orienting to task-irrelevant, infrequent novel stimuli presented during auditory oddball detection in patients with schizophrenia. The goal of the present study was to elucidate the functional abnormality underlying the disturbed orienting to novel stimuli in schizophrenia. Twenty-eight stable, partially remitted, medicated patients with schizophrenia and 28 healthy control participants completed a novelty oddball variant during event-related fMRI. Relative to healthy participants, patients with schizophrenia were characterised by underactivity during novel stimulus processing in the right amygdala-hippocampus, within paralimbic cortex in the rostral anterior cingulate and posterior cingulate cortices and the right frontal operculum, and in association cortex at the right temporo-parietal-occipital junction, bilateral intraparietal sulcus, and bilateral dorsal frontal cortex. Subcortically, relative hypoactivation during novelty processing was apparent in the cerebellum, thalamus, and basal ganglia. These results suggest that patients less efficiently reorient processing resources away from the ongoing task of detecting and responding to the task-relevant target stimuli. In addition, trend results suggest that patients experienced increased distraction by novel stimuli.     

The cortical representation of foveal stimuli: evidence from quadrantanopia and TMS-induced suppression

To address the extent to which the visual foveal representation is split, we examined a 29-year-old patient with a lower right quadrantanopia following surgical removal of the left occipital cortex above the calcarine sulcus and compared her performance with subjects receiving transcranial magnetic stimulation (TMS) over the occipital lobes. In a letter/digit classification task, the patient responded accurately to targets presented in the upper visual field, for all horizontal eccentricities. In the lower visual field, she failed to discriminate letters from digits when targets were presented in the right, but not the left visual field (RVF and LVF, respectively). This pattern was also true for the foveal targets, with poor performance to foveal-RVF (0.5 degrees to the right of fixation) but not foveal-LVF (0.5 degrees to the left of fixation) targets. Similar patterns of normal performance to LVF but not RVF or foveal-RVF targets were observed in a group of nine normal observers when TMS was applied over their left occipital cortex. Complementary impairments to LVF and foveal-LVF target classification were induced with TMS over the right occipital cortex. Thus, we have induced an hemianopic pattern in normal observers contralateral to the magnetically stimulated hemisphere. This correspondence between real and TMS-induced visual field defects is further evidence, in neurologically intact subjects, that the cortical representation of the fovea is split between the two hemispheres along the vertical meridian.     

Visual attention deficits in Alzheimer's disease: relationship to HMPAO SPECT cortical hypoperfusion

Patients with Alzheimer's disease (AD) display a multiplicity of cognitive deficits in domains such as memory, language, and attention, all of which can be clearly linked to the underlying neuropathological alterations. The typical degenerative changes occur early on in the disease in the temporal-parietal lobes, with other brain regions, such as the frontal cortex, becoming more affected as the disease progresses. In light of the importance of the parietal cortex in mediating visuospatial attentional processing, in the present study, we investigated a deficit in covert orienting of visual attention and its relationship to cortical hypoperfusion in AD. We characterized the visual attentional profile of 21 AD patients, relative to that of 26 matched normal individuals, and then assessed the correspondence between behavior and hypoperfusion, as measured by regional cerebral blood flow using SPECT. Relative to controls, the AD group demonstrated a unilateral attentional deficit, with disproportionate slowing in reorienting attention to targets in the left compared to the right hemispace, especially following an invalid peripheral cue. Furthermore, even in the presence of bilateral pathology typical of AD, there was a positive correlation between this unilateral attentional disorder and the magnitude of the right superior parietal lobe hypoperfusion. The association of the altered attentional processing profile (i.e., greater difficulty disengaging attention from right-sided stimuli) with right-hemisphere-predominant hypoperfusion not only confirms the critical role of the right parietal lobe in covert attentional orienting but, more importantly, identifies a potential locus of the behavioral alterations in visuospatial processing in AD.     

Altered brain functional connectivity in relation to perception of scrutiny in social anxiety disorder

Although the fear of being scrutinized by others in a social context is a key symptom in social anxiety disorder (SAD), the neural processes underlying the perception of scrutiny have not previously been studied by functional magnetic resonance imaging (fMRI). We used fMRI to map brain activation during a perception-of-scrutiny task in 20 SAD patients and 20 controls. A multi-dimensional analytic approach was used. Scrutiny perception was mediated by activation of the medial frontal cortex, insula-operculum region and cerebellum, and the additional recruitment of visual areas and the thalamus in patients. Between-group comparison demonstrated significantly enhanced brain activation in patients in the primary visual cortex and cerebellum. Functional connectivity mapping demonstrated an abnormal connectivity between regions underlying general arousal and attention. SAD patients showed significantly greater task-induced functional connectivity in the thalamo-cortical and the fronto-striatal circuits. A statistically significant increase in task-induced functional connectivity between the anterior cingulate cortex and scrutiny-perception-related regions was observed in the SAD patients, suggesting the existence of enhanced behavior-inhibitory control. The presented data indicate that scrutiny perception in SAD enhances brain activity in arousal-attention systems, suggesting that fMRI may be a useful tool to explore such a behavioral dimension.     

Sex and performance level effects on brain activation during a verbal fluency task: A functional magnetic resonance imaging study

Neuroimaging studies investigating the neural correlates of verbal fluency (VF) focused on sex differences without taking into account behavioural variation. Nevertheless, group differences in this verbal ability might account for neurocognitive differences elicited between men and women. The aim of this study was to test sex and performance level effects and the combination of these on cerebral activation. Four samples of 11 healthy students (N = 44) selected on the basis of sex and contrasted VF scores, high fluency (HF) versus low fluency (LF), performed a covert phonological VF task during scans. Within- and between-group analyses were conducted. Consistent with previous studies, for each sample, the whole-group analysis reported activation in the inferior frontal gyrus (IFG), insula, anterior cingulate cortex (ACC), medial frontal gyrus (mFG), superior (SPL) and inferior parietal lobules (IPL), inferior visual areas, cerebellum, thalamus and basal ganglia. Between-group analyses showed an interaction between sexes and performances in the right precuneus, left ACC, right IFG and left dorsolateral prefrontal cortex (dlPFC). HF men showed more activation than LF ones in the right precuneus and left dlPFC. LF men showed more activation in the right IFG than HF ones and LF women elicited more activation in the left ACC than HF ones. A sex main effect was found regardless of performance in the left inferior temporal gyrus (ITG), cerebellum, anterior and posterior cingulate cortexes and in the right superior frontal gyrus (SFG) and dlPFC, lingual gyrus and ACC, with men eliciting significantly greater activations than women. A performance main effect was found for the left ACC and the left cerebellum regardless of sex. LF subjects had stronger activations than HF ones in the ACC whereas HF subjects showed stronger activations in the cerebellum. Activity in three discrete subregions of the ACC is related to sex, performance and their interaction, respectively. Our findings emphasize the need to consider sex and performance level in functional imaging studies of VF.     

Involvement of the human frontal eye field and multiple parietal areas in covert visual selection during conjunction search

Searching for a target object in a cluttered visual scene requires active visual attention if the target differs from distractors not by elementary visual features but rather by a feature conjunction. We used functional magnetic resonance imaging (fMRI) in human subjects to investigate the functional neuroanatomy of attentional mechanisms employed during conjunction search. In the experimental condition, subjects searched for a target defined by a conjunction of colour and orientation. In the baseline condition, subjects searched for a uniquely coloured target, regardless of its orientation. Eye movement recordings outside the scanner verified subjects' ability to maintain fixation during search. Reaction times indicated that the experimental condition was attentionally more demanding than the baseline condition. Differential activations between conditions were therefore ascribed to top-down modulation of neural activity. The frontal eye field, the ventral precentral sulcus and the following posterior parietal regions were consistently activated: (i) the postcentral sulcus; (ii) the posterior; and (iii) the anterior part of the intraparietal sulcus; and (iv) the junction of the intraparietal with the transverse occipital sulcus. Parietal regions were spatially distinct and displayed differential amplitudes of signal increase with a maximal amplitude in the posterior intraparietal sulcus. Less consistent activation was found in the lateral fusiform gyrus. These results suggest an involvement of the human frontal eye field in covert visual selection of potential targets during search. These results also provide evidence for a subdivision of posterior parietal cortex in multiple areas participating in covert visual selection, with a major contribution of the posterior intraparietal sulcus.     

Neural basis of endogenous and exogenous spatial orienting. A functional MRI study

Whole-brain functional magnetic resonance imaging (MRI) was used to examine the neural substrates of internally (endogenous) and externally (exogenous) induced covert shifts of attention. Thirteen normal subjects performed three orienting conditions: endogenous (location of peripheral target predicted by a central arrow 80% of the time), exogenous (peripheral target preceded by noninformative central cue). Behavioral results indicated faster reaction times (RTs) for valid than for invalid trials for the endogenous condition but slower RTs for valid than for invalid trials for the exogenous condition (inhibition of return). The spatial extent and intensity of activation was greatest for the endogenous condition, consistent with the hypothesis that endogenous orienting is more effortful (less automatic) than exogenous orienting. Overall, we did not observe distinctly separable neural systems associated with the endogenous and exogenous orienting conditions. Both exogenous and endogenous orienting, but not the control condition, activated bilateral parietal and dorsal premotor regions, including the frontal eye fields. These results suggest a specific role for these regions in preparatory responding to peripheral stimuli. The right dorsolateral prefrontal cortex (BA 46) was activated selectively by the endogenous condition. This finding suggests that voluntary, but not reflexive, shifts of attention engage working memory systems.