Management of attentional resources in within-modal and cross-modal divided attention tasks: an fMRI study

In the present study, we were interested in distinguishing the cortical representations of within-modal and cross-modal divided attention tasks by using functional magnetic resonance imaging. Sixteen healthy male subjects aged between 21 and 30 years underwent two within-modal (auditory/auditory, visual/visual) and one cross-modal (auditory/visual) divided attention task, as well as related selective attention control conditions. After subtraction of the corresponding control task the three divided attention tasks, irrespective of sensory modality, revealed significant activation in a predominantly right hemisphere network involving the prefrontal cortex, the inferior parietal cortex, and the claustrum. Under the cross-modal condition, however, the frontal and parietal activation was more extended and more bilateral and there also was stronger right hemisphere activation of the anterior cingulate cortex and the thalamus. In comparison to the within-modal conditions additional bilateral frontal and left inferior parietal activation was found for the cross-modal condition. The supplementary fronto-parietal, anterior cingulate cortex, and thalamus activation in the auditory/visual condition could be argued to reflect an additional demand for coordination of two ongoing cross-modal cognitive processes.     

Attending to and remembering tactile stimuli: a review of brain imaging data and single-neuron responses.

Clinical and neuroimaging observations of the cortical network implicated in tactile attention have identified foci in parietal somatosensory, posterior parietal, and superior frontal locations. Tasks involving intentional hand-arm movements activate similar or nearby parietal and frontal foci. Visual spatial attention tasks and deliberate visuomotor behavior also activate overlapping posterior parietal and frontal foci. Studies in the visual and somatosensory systems thus support a proposal that attention to the spatial location of an object engages cortical regions responsible for the same coordinate referents used for guiding purposeful motor behavior. Tactile attention also biases processing in the somatosensory cortex through amplification of responses to relevant features of selected stimuli. Psychophysical studies demonstrate retention gradients for tactile stimuli like those reported for visual and auditory stimuli, and suggest analogous neural mechanisms for working memory across modalities. Neuroimaging studies in humans using memory tasks, and anatomic studies in monkeys support the idea that tactile information relayed from the somatosensory cortex is directed ventrally through the insula to the frontal cortex for short-term retention and to structures of the medial temporal lobe for long-term encoding. At the level of single neurons, tactile (such as visual and auditory) short-term memory appears as a persistent response during delay intervals between sampled stimuli.     

Sparse imaging of the auditory oddball task with functional MRI

Cortical reactions to rare task-related stimuli have been studied with electrophysiological methods in the assessment of the P300 component as well as in functional imaging studies with regard to oddball tasks. While functional magnetic resonance imaging studies using auditory stimuli have to deal with interference between auditory stimuli and scanner noise, the aim of our study was to assess auditory target processing with the sparse imaging method. Single volumes of echo-planar imaging were acquired 4 s following the onset of the stimuli of interest. In keeping with previous studies, target stimuli activated the insula, superior temporal gyrus, cingulate gyrus, middle frontal gyrus, parietal cortex and the cerebellum. Our results encourage the application of the sparse imaging method in experiments on cognitive processing elicited by auditory stimulation in a silent environment.     

A visual or tactile signal makes auditory speech detection more efficient by reducing uncertainty

Acoustic speech is easier to detect in noise when the talker can be seen. This finding could be explained by integration of multisensory inputs or refinement of auditory processing from visual guidance. In two experiments, we studied two‐interval forced‐choice detection of an auditory ‘ba’ in acoustic noise, paired with various visual and tactile stimuli that were identically presented in the two observation intervals. Detection thresholds were reduced under the multisensory conditions vs. the auditory‐only condition, even though the visual and/or tactile stimuli alone could not inform the correct response. Results were analysed relative to an ideal observer for which intrinsic (internal) noise and efficiency were independent contributors to detection sensitivity. Across experiments, intrinsic noise was unaffected by the multisensory stimuli, arguing against the merging (integrating) of multisensory inputs into a unitary speech signal, but sampling efficiency was increased to varying degrees, supporting refinement of knowledge about the auditory stimulus. The steepness of the psychometric functions decreased with increasing sampling efficiency, suggesting that the ‘task‐irrelevant’ visual and tactile stimuli reduced uncertainty about the acoustic signal. Visible speech was not superior for enhancing auditory speech detection. Our results reject multisensory neuronal integration and speech‐specific neural processing as explanations for the enhanced auditory speech detection under noisy conditions. Instead, they support a more rudimentary form of multisensory interaction: the otherwise task‐irrelevant sensory systems inform the auditory system about when to listen.     

Facilitative effect of repetitive presentation of one stimulus on cortical responses to other stimuli in macaque monkeys – a possible neural mechanism for mismatch negativity

The event‐related potential ‘mismatch negativity’ (MMN) is an indicator of a perceiver's ability to detect deviations in sensory signal streams. MMN and its homologue in animals, mismatch activity (MMA), are differential neural responses to a repeatedly presented stimulus and a subsequent deviant stimulus (oddball). Because neural mechanisms underlying MMN and MMA remain unclear, there is a controversy as to whether MMN and MMA arise solely from stimulus‐specific adaptation (SSA), in which the response to a stimulus cumulatively attenuates with its repetitive presentation. To address this issue, we used electrocorticography and the auditory roving‐oddball paradigm in two awake macaque monkeys. We examined the effect of stimulus repetition number on MMA and on responses to repeated stimuli and oddballs across the cerebral cortex in the time–frequency domain. As the repetition number increased, MMA spread across the temporal, frontal and parietal cortices, and each electrode yielded a larger MMA. Surprisingly, this increment in MMA largely depended on response augmentation to the oddball rather than on SSA to the repeated stimulus. Following sufficient repetition, the oddball evoked a spectral power increment in some electrodes on the frontal cortex that had shown no power increase to the stimuli with less or no preceding repetition. We thereby revealed that repetitive presentation of one stimulus not only leads to SSA but also facilitates the cortical response to oddballs involving a wide range of cortical regions. This facilitative effect might underlie the generation of MMN‐like scalp potentials in macaques that potentially shares similar neural mechanisms with MMN in humans.     

Behavioural benefits of multisensory processing in ferrets

Enhanced detection and discrimination, along with faster reaction times, are the most typical behavioural manifestations of the brain's capacity to integrate multisensory signals arising from the same object. In this study, we examined whether multisensory behavioural gains are observable across different components of the localization response that are potentially under the command of distinct brain regions. We measured the ability of ferrets to localize unisensory (auditory or visual) and spatiotemporally coincident auditory–visual stimuli of different durations that were presented from one of seven locations spanning the frontal hemifield. During the localization task, we recorded the head movements made following stimulus presentation, as a metric for assessing the initial orienting response of the ferrets, as well as the subsequent choice of which target location to approach to receive a reward. Head‐orienting responses to auditory–visual stimuli were more accurate and faster than those made to visual but not auditory targets, suggesting that these movements were guided principally by sound alone. In contrast, approach‐to‐target localization responses were more accurate and faster to spatially congruent auditory–visual stimuli throughout the frontal hemifield than to either visual or auditory stimuli alone. Race model inequality analysis of head‐orienting reaction times and approach‐to‐target response times indicates that different processes, probability summation and neural integration, respectively, are likely to be responsible for the effects of multisensory stimulation on these two measures of localization behaviour.     

Contextual factors multiplex to control multisensory processes

This study analyzed high‐density event‐related potentials (ERPs) within an electrical neuroimaging framework to provide insights regarding the interaction between multisensory processes and stimulus probabilities. Specifically, we identified the spatiotemporal brain mechanisms by which the proportion of temporally congruent and task‐irrelevant auditory information influences stimulus processing during a visual duration discrimination task. The spatial position (top/bottom) of the visual stimulus was indicative of how frequently the visual and auditory stimuli would be congruent in their duration (i.e., context of congruence). Stronger influences of irrelevant sound were observed when contexts associated with a high proportion of auditory‐visual congruence repeated and also when contexts associated with a low proportion of congruence switched. Context of congruence and context transition resulted in weaker brain responses at 228 to 257 ms poststimulus to conditions giving rise to larger behavioral cross‐modal interactions. Importantly, a control oddball task revealed that both congruent and incongruent audiovisual stimuli triggered equivalent non‐linear multisensory interactions when congruence was not a relevant dimension. Collectively, these results are well explained by statistical learning, which links a particular context (here: a spatial location) with a certain level of top‐down attentional control that further modulates cross‐modal interactions based on whether a particular context repeated or changed. The current findings shed new light on the importance of context‐based control over multisensory processing, whose influences multiplex across finer and broader time scales. Hum Brain Mapp 37:273–288, 2016. © 2015 Wiley Periodicals, Inc.     

A biologically inspired neurocomputational model for audiovisual integration and causal inference

Recently, experimental and theoretical research has focused on the brain's abilities to extract information from a noisy sensory environment and how cross‐modal inputs are processed to solve the causal inference problem to provide the best estimate of external events. Despite the empirical evidence suggesting that the nervous system uses a statistically optimal and probabilistic approach in addressing these problems, little is known about the brain's architecture needed to implement these computations. The aim of this work was to realize a mathematical model, based on physiologically plausible hypotheses, to analyze the neural mechanisms underlying multisensory perception and causal inference. The model consists of three layers topologically organized: two encode auditory and visual stimuli, separately, and are reciprocally connected via excitatory synapses and send excitatory connections to the third downstream layer. This synaptic organization realizes two mechanisms of cross‐modal interactions: the first is responsible for the sensory representation of the external stimuli, while the second solves the causal inference problem. We tested the network by comparing its results to behavioral data reported in the literature. Among others, the network can account for the ventriloquism illusion, the pattern of sensory bias and the percept of unity as a function of the spatial auditory–visual distance, and the dependence of the auditory error on the causal inference. Finally, simulations results are consistent with probability matching as the perceptual strategy used in auditory–visual spatial localization tasks, agreeing with the behavioral data. The model makes untested predictions that can be investigated in future behavioral experiments.     

Involvement of the dorsal and ventral attention networks in oddball stimulus processing: A meta‐analysis

The aim of this study was to provide the first, comprehensive meta‐analysis of the neuroimaging literature regarding greater neural responses to a deviant stimulus in a stream of repeated, standard stimuli, termed here oddball effects. The meta‐analysis of 75 independent studies included a comparison of auditory and visual oddball effects and task‐relevant and task‐irrelevant oddball effects. The results were interpreted with reference to the model in which a large‐scale dorsal frontoparietal network embodies a mechanism for orienting attention to the environment, whereas a large‐scale ventral frontoparietal network supports the detection of salient, environmental changes. The meta‐analysis yielded three main sets of findings. First, ventral network regions were strongly associated with oddball effects and largely common to auditory and visual modalities, indicating a supramodal “alerting” system. Most ventral network components were more strongly associated with task‐relevant than task‐irrelevant oddball effects, indicating a dynamic interplay of stimulus saliency and internal goals in stimulus‐driven engagement of the network. Second, the bilateral inferior frontal junction, an anterior core of the dorsal network, was strongly associated with oddball effects, suggesting a central role in top‐down attentional control. However, other dorsal network regions showed no or only modest association with oddball effects, likely reflecting active engagement during both oddball and standard stimulus processing. Finally, prominent oddball effects outside the two networks included the sensory cortex regions, likely reflecting attentive and preattentive modulation of early sensory activity, and subcortical regions involving the putamen, thalamus, and other areas, likely reflecting subcortical involvement in alerting responses. Hum Brain Mapp 35:2265–2284, 2014. © 2013 Wiley Periodicals, Inc .     

An event-related functional magnetic resonance imaging study of an auditory oddball task in schizophrenia

Schizophrenia is a diffuse brain disease that affects many facets of cognitive function. One of the most replicated findings in the neurobiology of schizophrenia is that the event-related potentials to auditory oddball stimuli are abnormal, effects believed to be related to abnormalities in attentional and memory processes. Although event-related potentials provide excellent resolution regarding the time course of information processing, such studies are poor at characterizing the spatial location of these abnormalities. To address this issue, we used event-related functional magnetic resonance imaging techniques to elucidate the neural areas underlying target detection in schizophrenia. Consistent with recent event-related functional magnetic resonance imaging results, target processing by control participants was associated with bilateral activation in the anterior superior temporal gyri, inferior and superior parietal lobules, and activation in anterior and posterior cingulate, thalamus, and right lateral frontal cortex. For the schizophrenic patients, selective deficits were observed in both the extent and strength of activation associated with target processing in the right lateral frontal cortex, thalamus, bilateral anterior superior temporal gyrus, anterior and posterior cingulate, and right inferior and superior parietal lobules. These findings are consistent with the evidence for abnormal processing of oddball stimuli suggested by event-related potential studies in schizophrenic patients, but provide much more detailed evidence regarding the anatomical sites implicated. These data are consistent with the hypothesis that schizophrenia is characterized by a widespread pathological process affecting many cerebral areas, including association cortex and thalamus.     

Modulating auditory selective attention by non‐invasive brain stimulation: Differential effects of transcutaneous vagal nerve stimulation and transcranial random noise stimulation

Selective attention is a basic process required to maintain goal‐directed behavior by appropriately responding to target stimuli and suppressing reactions to non‐target stimuli. It has been proposed that auditory selective attention is linked to the activity of the locus coeruleus‐norepinergic (LC‐NE) system and a large‐scale fronto‐parietal cortical network, but there is still sparse causal evidence for these assumptions. By applying transcutaneous vagal nerve stimulation (tVNS) and transcranial random noise stimulation (tRNS) over the frontal cortex, we systematically assessed the involvement of these subcortical and cortical components in the regulation of auditory selective attention. Using a single‐blinded, sham‐controlled, within‐subject design we analyzed online effects of tVNS and tRNS in 20 healthy participants during an auditory oddball paradigm. We show significant stimulation‐dependent modulations of auditory selective attention on the behavioral and electrophysiological level. Compared to sham, tVNS increased the P3 amplitude, while tRNS reduced the reaction time to target stimuli. Moreover, both techniques reduced the P3 latency. Our data provide evidence for the functional relevance of the subcortical NE system in the regulation of neural resources that allows a phasic response to incoming target stimuli. They indicate that frontal cortex structures are crucially involved in the successful evaluation of the respective information. Moreover, our results are in favor of the LC‐P3 hypothesis claiming the vital role of the NE system in auditory selective attention and in the generation of the P3. Of note, the effects of tVNS on auditory selective attention are comparable with those evoked by pharmacological interventions and invasive vagal nerve stimulation.     

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.     

Noise‐rearing disrupts the maturation of multisensory integration

It is commonly believed that the ability to integrate information from different senses develops according to associative learning principles as neurons acquire experience with co‐active cross‐modal inputs. However, previous studies have not distinguished between requirements for co‐activation versus co‐variation. To determine whether cross‐modal co‐activation is sufficient for this purpose in visual–auditory superior colliculus (SC) neurons, animals were reared in constant omnidirectional noise. By masking most spatiotemporally discrete auditory experiences, the noise created a sensory landscape that decoupled stimulus co‐activation and co‐variance. Although a near‐normal complement of visual–auditory SC neurons developed, the vast majority could not engage in multisensory integration, revealing that visual–auditory co‐activation was insufficient for this purpose. That experience with co‐varying stimuli is required for multisensory maturation is consistent with the role of the SC in detecting and locating biologically significant events, but it also seems likely that this is a general requirement for multisensory maturation throughout the brain.     

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 neural circuitry underlying the executive control of auditory spatial attention

Although a fronto-parietal network has consistently been implicated in the control of visual spatial attention, the network that guides spatial attention in the auditory domain is not yet clearly understood. To investigate this issue, we measured brain activity using functional magnetic resonance imaging while participants performed a cued auditory spatial attention task. We found that cued orienting of auditory spatial attention activated a medial-superior distributed fronto-parietal network. In addition, we found cue-triggered increases of activity in the auditory sensory cortex prior to the occurrence of an auditory target, suggesting that auditory attentional control operates in part by biasing processing in sensory cortex in favor of expected target stimuli. Finally, an exploratory cross-study comparison further indicated several common frontal and parietal regions as being involved in the control of both visual and auditory spatial attention. Thus, the present findings not only reveal the network of brain areas underlying endogenous spatial orienting in the auditory modality, but also suggest that the control of spatial attention in different sensory modalities is enabled in part by some common, supramodal neural mechanisms.     

Prestimulus oscillatory alpha power and connectivity patterns predispose perceptual integration of an audio and a tactile stimulus

To efficiently perceive and respond to the external environment, our brain has to perceptually integrate or segregate stimuli of different modalities. The temporal relationship between the different sensory modalities is therefore essential for the formation of different multisensory percepts. In this magnetoencephalography study, we created a paradigm where an audio and a tactile stimulus were presented by an ambiguous temporal relationship so that perception of physically identical audiotactile stimuli could vary between integrated (emanating from the same source) and segregated. This bistable paradigm allowed us to compare identical bimodal stimuli that elicited different percepts, providing a possibility to directly infer multisensory interaction effects. Local differences in alpha power over bilateral inferior parietal lobules (IPLs) and superior parietal lobules (SPLs) preceded integrated versus segregated percepts of the two stimuli (audio and tactile). Furthermore, differences in long‐range cortical functional connectivity seeded in rIPL (region of maximum difference) revealed differential patterns that predisposed integrated or segregated percepts encompassing secondary areas of all different modalities and prefrontal cortex. We showed that the prestimulus brain states predispose the perception of the audiotactile stimulus both in a global and a local manner. Our findings are in line with a recent consistent body of findings on the importance of prestimulus brain states for perception of an upcoming stimulus. This new perspective on how stimuli originating from different modalities are integrated suggests a non‐modality specific network predisposing multisensory perception. Hum Brain Mapp 36:3486–3498, 2015. © 2015 Wiley Periodicals, Inc .     

Visual attention circuitry in schizophrenia investigated with oddball event-related functional magnetic resonance imaging

OBJECTIVE: Schizophrenia patients have problems directing attention. Sustained attention requires ensuring that brain resources are focused on a selected target (top-down task) while ignoring irrelevant distractors (bottom-up interference). Whether patients have too little ability to focus or too much interference from distraction has not been clarified. The oddball paradigm embeds infrequent targets and distractors into the stimulus train, and schizophrenia deficits have been linked to diminished responses to both. Cerebral activity underlying abnormal attention can be examined with event-related functional magnetic resonance imaging. METHOD: A visual oddball task was presented to 22 patients with schizophrenia and 28 comparison subjects. Statistical probability maps reflecting blood-oxygenation-level-dependent changes were generated for infrequent targets and novel distractors relative to frequent standard stimuli. Activation was related to performance and symptoms. RESULTS: Activation specific to targets and distractors was associated with faster performance. For targets, patients had diminished activation in superior temporal and frontal gyri, cingulate, thalamus, and basal ganglia. They had increased activation in right insula, mid-frontal gyrus, posterior cingulate, and left inferior parietal lobule. For distractors, patients showed less activation in occipital regions and left inferior parietal lobule but increased activation in parietal-occipital, right mid-frontal, and left inferior frontal gyri. Abnormal activation correlated with positive and negative symptoms. CONCLUSIONS: Abnormal activation in schizophrenia in response to attentional demands reflects both insufficient recruitment of brain systems required for target detection and overcommitment of resources for processing irrelevant distractors. Schizophrenia patients appear to have an inability both to focus on targets and ignore distraction.     

Are you listening? Brain activation associated with sustained nonspatial auditory attention in the presence and absence of stimulation

Neuroimaging studies investigating the voluntary (top‐down) control of attention largely agree that this process recruits several frontal and parietal brain regions. Since most studies used attention tasks requiring several higher‐order cognitive functions (e.g. working memory, semantic processing, temporal integration, spatial orienting) as well as different attentional mechanisms (attention shifting, distractor filtering), it is unclear what exactly the observed frontoparietal activations reflect. The present functional magnetic resonance imaging study investigated, within the same participants, signal changes in (1) a “Simple Attention” task in which participants attended to a single melody, (2) a “Selective Attention” task in which they simultaneously ignored another melody, and (3) a “Beep Monitoring” task in which participants listened in silence for a faint beep. Compared to resting conditions with identical stimulation, all tasks produced robust activation increases in auditory cortex, cross‐modal inhibition in visual and somatosensory cortex, and decreases in the default mode network, indicating that participants were indeed focusing their attention on the auditory domain. However, signal increases in frontal and parietal brain areas were only observed for tasks 1 and 2, but completely absent for task 3. These results lead to the following conclusions: under most conditions, frontoparietal activations are crucial for attention since they subserve higher‐order cognitive functions inherently related to attention. However, under circumstances that minimize other demands, nonspatial auditory attention in the absence of stimulation can be maintained without concurrent frontal or parietal activations. Hum Brain Mapp 35:2233–2252, 2014. © 2013 Wiley Periodicals, Inc.     

Response amplification in sensory-specific cortices during crossmodal binding

Integrating information across the senses can enhance our ability to detect and classify stimuli in the environment. For example, auditory speech perception is substantially improved when the speaker's face is visible. In an fMRI study designed to investigate the neural mechanisms underlying these crossmodal behavioural gains, bimodal (audio-visual) speech was contrasted against both unimodal (auditory and visual) components. Significant response enhancements in auditory (BA 41/42) and visual (V5) cortices were detected during bimodal stimulation. This effect was found to be specific to semantically congruent crossmodal inputs. These data suggest that the perceptual improvements effected by synthesizing matched multisensory inputs are realised by reciprocal amplification of the signal intensity in participating unimodal cortices.     

The cortical generators of P3a and P3b: a LORETA study

The P3 is probably the most well known component of the brain event-related potentials (ERPs). Using a three-tone oddball paradigm two different components can be identified: the P3b elicited by rare target stimuli and the P3a elicited by the presentation of rare non-target stimuli. Although the two components may partially overlap in time and space, they have a different scalp topography suggesting different neural generators. The present study is aimed at defining the scalp topography of the two P3 components by means of reference-independent methods and identifying their electrical cortical generators by using the low-resolution electromagnetic tomography (LORETA). ERPs were recorded during a three-tone oddball task in 32 healthy, right-handed university students. The scalp topography of the P3 components was assessed by means of the brain electrical microstates technique and their cortical sources were evaluated by LORETA. P3a and P3b showed different scalp topography and cortical sources. The P3a electrical field had a more anterior distribution as compared to the P3b and its generators were localized in cingulate, frontal and right parietal areas. P3b sources included bilateral frontal, parietal, limbic, cingulate and temporo-occipital regions. Differences in scalp topography and cortical sources suggest that the two components reflect different neural processes. Our findings on cortical generators are in line with the hypothesis that P3a reflects the automatic allocation of attention, while P3b is related to the effortful processing of task-relevant events.