Do ERP components triggered during attentional orienting represent supramodal attentional control?

Lateralized ERP components triggered during cued shifts of spatial attention (anterior directing attention negativity [ADAN], late directing attention positivity [LDAP]) have been observed during visual, auditory, and tactile attention tasks, suggesting that these components reflect supramodal attentional control processes. This interpretation has recently been called into question by the finding that the ADAN is absent in response to auditory attention cues. Here we demonstrate that ADAN and LDAP components are reliably elicited in a purely unimodal auditory attention task where auditory cues are followed by auditory imperative stimuli. The fact that the ADAN is not restricted to task contexts where visual or tactile stimuli are relevant is consistent with the hypothesis that this component is linked to supramodal attentional control.     

Control mechanisms mediating shifts of attention in auditory and visual space: a spatio-temporal ERP analysis

The neural systems that mediate voluntary shifts of attention to visual and auditory stimuli were investigated by examining the patterns of human brain electricity elicited by attention-directing cues in auditory and visual tasks. Several lateralized event-related potential (ERP) components were observed when participants shifted attention in expectation of visual targets (experiment 1). One component was focused over frontal cortex and a second was focused primarily over the occipital-temporal cortex but also spread to parietal regions of the scalp. Previous work has indicated that the frontal component reflects supramodal processes involved in the executive control of attention and that the posterior component reflects either spatial attentional control processes in the posterior parietal lobe or modulation of processes in visual cortex. Here, the posterior component was observed when participants shifted attention in expectation of auditory targets (experiments 2-4), but the frontal component was found only in the visual task. The posterior component seemed to be generated in parietal and occipital areas even when there was no visual information about the to-be-attended locations. These results are consistent with the view that voluntary shifts of attention are mediated by supramodal processes in the parietal lobe and that the spatial coordinates of the to-be-attended location are based on visual representations of space.     

Spatial attention triggered by unimodal, crossmodal, and bimodal exogenous cues: a comparison of reflexive orienting mechanisms

The aim of this study was to establish whether spatial attention triggered by bimodal exogenous cues acts differently as compared to unimodal and crossmodal exogenous cues due to crossmodal integration. In order to investigate this issue, we examined cuing effects in discrimination tasks and compared these effects in a condition wherein a visual target was preceded by both visual and auditory exogenous cues delivered simultaneously at the same side (bimodal cue), with conditions wherein the visual target was preceded by either a visual (unimodal cue) or an auditory cue (crossmodal cue). The results of two experiments revealed that cuing effects on RTs in these three conditions with an SOA of 200 ms had comparable magnitudes. Differences at a longer SOA of 600 ms (inhibition of return for bimodal cues, Experiment 1) disappeared when catch trials were included (in Experiment 2). The current data do not support an additional influence of crossmodal integration on exogenous orienting, but are well in agreement with the existence of a supramodal spatial attention module that allocates attentional resources towards stimulated locations for different sensory modalities.     

The N2pc component and its links to attention shifts and spatially selective visual processing

The N2pc component has recently become a popular tool in attention research. To investigate whether this component exclusively reflects attentional target selection or also prior stages in attentional processing (covert orienting, target-unspecific spatial attention), a spatial cuing procedure was combined with a visual search task. In some blocks, informative cues indicated the side of upcoming singleton targets that were present on most trials among uniform distractors. In other blocks, cues were spatially uninformative, and no preparatory shifts of attention were possible. The N2pc in response to targets was unaffected by this manipulation, showing that this component is not associated with attention shifts. Following informative cues, an attenuated N2pc was elicited by uniform nontarget arrays, suggesting that the N2pc may also reflect spatially specific processing of stimulus features at task-relevant locations prior to target selection.     

Top‐down control of audiovisual search by bimodal search templates

To test whether the attentional selection of targets defined by a combination of visual and auditory features is guided in a modality‐specific fashion or by control processes that are integrated across modalities, we measured attentional capture by visual stimuli during unimodal visual and audiovisual search. Search arrays were preceded by spatially uninformative visual singleton cues that matched the current target‐defining visual feature. Participants searched for targets defined by a visual feature, or by a combination of visual and auditory features (e.g., red targets accompanied by high‐pitch tones). Spatial cueing effects indicative of attentional capture were reduced during audiovisual search, and cue‐triggered N2pc components were attenuated and delayed. This reduction of cue‐induced attentional capture effects during audiovisual search provides new evidence for the multimodal control of selective attention.     

Target processing is facilitated by motivationally relevant cues

The effect that motivationally relevant stimuli have on processes of attentional engagement and disengagement was investigated during two modified peripheral cueing paradigms. Sexual, mutilation, threatening, and neutral stimuli served as peripheral cues in both experiments. Responses were made to target location in Experiment 1 (N=19 female) and target identity in Experiment 2 (N=18 female). As indexed by enhanced target-evoked P1 and P3b component amplitudes, target processing was facilitated by the presentation of sexual and mutilation stimuli in both experiments. This facilitation in response to targets cued by sexual and mutilation stimuli occurred regardless of whether cueing was valid or invalid as demonstrated by the non-significant cue validity x picture-type interaction. As such, the processes of attentional engagement (as inferred by responses to validly cued targets) and attentional disengagement (as inferred by responses to invalidly cued targets) were not differentially affected by the motivational relevance of the preceding cue. These results indicate that in a non-clinical sample, participants can shift attention rapidly to process information following the onset of motivationally relevant stimuli at attended (valid) and unattended (invalid) locations and that target processing is facilitated by the presence of appetitive and aversive cues.     

ERP correlates of tactile spatial attention differ under intra- and intermodal conditions

To investigate whether the mechanisms underlying endogenous tactile spatial attention differ under pure tactile compared to mixed modality conditions event-related brain potentials (ERPs) were recorded to bilateral tactile and visual cues and tactile imperative stimuli. In the cue-stimulus interval the anterior directing attention negativity (ADAN) was present contralateral to the side of the attentional shift. Importantly, under pure tactile conditions this component persisted until imperative stimulus onset, while it diminished under intermodal conditions. Furthermore, post-tactile stimulus onset attentional modulations were present for the P100 component and later latencies under intermodal conditions. In contrast, under pure tactile conditions attentional modulations only emerged for the N140 component and later latencies. It is suggested that mechanisms underlying attentional orienting and selection are not entirely supramodal but depend in part on the modalities involved.     

ERP correlates of tactile spatial attention differ under intra- and intermodal conditions

To investigate whether the mechanisms underlying endogenous tactile spatial attention differ under pure tactile compared to mixed modality conditions event-related brain potentials (ERPs) were recorded to bilateral tactile and visual cues and tactile imperative stimuli. In the cue-stimulus interval the anterior directing attention negativity (ADAN) was present contralateral to the side of the attentional shift. Importantly, under pure tactile conditions this component persisted until imperative stimulus onset, while it diminished under intermodal conditions. Furthermore, post-tactile stimulus onset attentional modulations were present for the P100 component and later latencies under intermodal conditions. In contrast, under pure tactile conditions attentional modulations only emerged for the N140 component and later latencies. It is suggested that mechanisms underlying attentional orienting and selection are not entirely supramodal but depend in part on the modalities involved.     

Attention to endogenous and exogenous cues affects auditory localization

Three experiments examine attentional differences in auditory localization using either endogenous or exogenous visual cues. Participants were presented with visual cues on a computer screen and asked to localize auditory targets presented through headphones. In conditions in which the auditory and visual stimuli traveled in the same direction, participants showed illusory directional hearing (Hari in Neurosci Lett 189:29–30, 1995) in that the targets were perceived to travel through the head. In conditions in which the directions of the auditory and visual stimuli were conflicting, participants localized the auditory targets as traveling in the direction of the visual cues. These data suggest that visual capture plays a predominate role in the processes of auditory localization that occurs within the head. Additionally, endogenous response times were significantly greater than exogenous response times. We propose this is the result of additional time required to shift one’s attentional window in the endogenous condition.     

The effects of auditory and visual linguistic distractors on target localization

In this experiment, a Stroop-like paradigm was used to investigate the ability to attend to visuospatial cues while ignoring distracting stimuli in the auditory or visual modality. In Part 1, the authors investigated whether linguistic cue words (i.e., RIGHT, LEFT, DOWN, and UP) would induce endogenous shifts of attention to visual targets. In Part 2, a relevant distractor stimulus was introduced in a different modality from the endogenous cues to investigate effects of interference. Twenty-five right-handed students served as participants. Auditory and visual linguistic cues were effective in inducing shifts of visual attention when cues were presented alone. Furthermore, introducing a distractor stimulus decreased the efficacy of these cues differently depending on modality, suggesting that language processing and visuospatial attention may share neuronal resources. Implications for unimodal and supramodal mechanisms of selective attention and relevant neuronal networks are discussed.     

Attentional capture by a color singleton is stronger at spatially relevant than irrelevant locations: Evidence from an ERP study

Top-down spatial attention can modulate contingent attentional capture, but the underlying mechanism is still not clear. Using variants of spatial cueing paradigms, our previous event-related potential study showed that peripheral color singleton cues with task-relevant features captured attention (indexed by cue-elicited N2pc) even when the targets appeared at central locations, but the magnitude of attentional capture was smaller than when the targets appeared at same peripheral locations. One reasonable explanation is that the modulation effect is due to spatial relevance of cues. However, several other confounding factors might also explain the modulation effect, such as task difficulty, spread of attentional window, and inside/outside relation between cue and attentional window. In the present study, we rearranged the relative locations between cues and targets to control these factors and to further examine whether inequivalence of attentional capture across attended and unattended locations was a common phenomenon. In two experiments, color singleton cues elicited apparent N2pc components when participants were searching for targets possessing the same color, which replicated typical findings of contingent attentional capture. More importantly, the modulation effect of top-down spatial attention on the cue-elicited N2pc still existed even when the factors of task difficulty (Experiment 1), spread of attentional window (Experiment 1), or inside/outside relation between cue and attentional window (Experiment 2) were controlled. These results consistently demonstrate that attentional capture by a color singleton is stronger at spatially relevant locations than at spatially irrelevant locations, suggesting an important role of spatial relevance on contingent attentional capture.     

Inhibition of return following an auditory cue The role of central reorienting events

 It is currently controversial whether auditory events produce inhibition of return (IOR). Although some authors argue that this can arise, others propose that peripheral auditory cues do not produce the characteristic IOR pattern of delayed detection latencies for ipsilaterally presented auditory or visual targets, unless a saccade is made to the cued location. We suggest that these previous discrepancies may depend on whether attention is reoriented centrally following the peripheral sound. We presented spatially uninformative peripheral auditory cues prior to visual targets requiring speeded detection responses. IOR was found in the absence of eye movements, provided an auditory reorienting event was presented at central fixation between onset of the peripheral cue and the subsequent target, but not when the central reorienting event was visual. A subsequent experiment demonstrated that auditory IOR between successive targets is similarly significantly reduced in the absence of an appropriate central reorienting event. These results imply that auditory stimuli can induce IOR directly. Previous failures to demonstrate IOR following auditory cues may have been due to an opposing influence of long-lasting attentional facilitation at the cued location, rather than to the putative inability of auditory stimuli to engage oculomotor processes generating IOR. Received: 17 March 1997 / Accepted: 8 August 1997     

Preservation of crossmodal selective attention in healthy aging

The goal of the present study was to determine if older adults benefited from attention to a specific sensory modality in a voluntary attention task and evidenced changes in voluntary or involuntary attention when compared to younger adults. Suppressing and enhancing effects of voluntary attention were assessed using two cued forced-choice tasks, one that asked participants to localize and one that asked them to categorize visual and auditory targets. Involuntary attention was assessed using the same tasks, but with no attentional cues. The effects of attention were evaluated using traditional comparisons of means and Cox proportional hazards models. All analyses showed that older adults benefited behaviorally from selective attention in both visual and auditory conditions, including robust suppressive effects of attention. Of note, the performance of the older adults was commensurate with that of younger adults in almost all analyses, suggesting that older adults can successfully engage crossmodal attention processes. Thus, age-related increases in distractibility across sensory modalities are likely due to mechanisms other than deficits in attentional processing.     

Comparing intramodal and crossmodal cuing in the endogenous orienting of spatial attention

The endogenous orienting of spatial attention has been studied with both informative central cues and informative peripheral cues. Central cues studies are difficult to compare with studies that have used uninformative peripheral cues due to the differences in stimulus presentation. Moreover, informative peripheral cues attract both endogenous and exogenous attention, thus making it difficult to disentangle the contribution of each process to any behavioural results observed. In the present study, we used an informative peripheral cue (either tactile or visual) that predicted that the target would appear (in different blocks of trials) on either the same or opposite side as the cue. By using this manipulation, both expected and unexpected trials could either be exogenously cued or uncued, thus making it possible to isolate expectancy effects from cuing effects. Our aim was to compare the endogenous orienting of spatial attention to tactile (Experiment 1) and to visual targets (Experiment 2) under conditions of intramodal and crossmodal spatial cuing. The results suggested that the endogenous orienting of spatial attention should not be considered as being a purely supramodal phenomenon, given that significantly larger expectancy effects were observed in the intramodal cuing conditions than in the crossmodal cuing conditions in both experiments.     

Spatial and semantic inhibitory processing in schizophrenia.

Two experiments assessed inhibitory mechanisms associated with the posterior and anterior attention networks in schizophrenia. Experiment 1 assessed the inhibition of return effect of the posterior network. Both healthy adults and schizophrenic adults showed inhibition of return, suggesting that this inhibitory mechanism of visual orienting is preserved in schizophrenia. Experiment 2 assessed semantic inhibition, which supposedly taps the anterior network, in a lexical-decision task. Healthy adults showed semantic inhibition effects in both visual fields. Schizophrenic adults showed semantic inhibition effects when targets were presented to the left visual field, involving the right hemisphere. However, semantic facilitation rather than inhibition was observed when targets were presented to the right visual field, involving the left hemisphere. These results reflect left hemisphere dysfunction associated with deficits in attentional control in schizophrenia.     

Lateralized irrelevant speech alters visuospatial selective attention mechanisms

Recent studies indicate that the coordination of spatial attention across modalities may in part be mediated by a supramodal attentional system. We try to extend the concept of a supramodal system and hypothesized that involuntary modulations of auditory attentional processes by irrelevant speech signals influence visuospatial attention, suggesting crossmodal links between vision and speech. In order to test this we recorded event-related brain potentials (ERPs) of 12 healthy subjects in a visuospatial selective attention task. The task to identify target stimuli appearing at lateral visual field locations caused the expected enhancements of the early P1 and N1 ERP components to attended visual stimuli. Understandable and ununderstandable task irrelevant speech was presented either at the visually attended position or in the opposite visual field location. Speech contralateral to unattended visual stimuli led to a decreased N1 amplitude. This effect was stronger for understandable speech. Thus, speech influences the allocation of visual spatial attention if it is presented in the unattended location. The results suggest crossmodal links of speech and visuospatial attention mechanisms at a very early stage of human perception.     

Lateralized ERP components related to spatial orienting: discriminating the direction of attention from processing sensory aspects of the cue

Two spatial cueing experiments were conducted to examine the functional significance of lateralized ERP components after cue-onset and to discriminate components related to sensory cue aspects and components related to the direction of attention. In Experiment 1, a simple detection task was presented. In Experiment 2, attentional selection was augmented. Two unimodal visual cueing tasks were presented using nonlateralized line cues and lateralized arrow cues. Lateralized cue effects and modulation after stimulus onset were stronger in Experiment 2. An early posterior component was related to the physical shape of arrows. A posterior negativity (EDAN) may be related to the encoding of direction from arrow cues. An anterior negativity (ADAN) and a posterior positivity (LDAP) were related to the direction of attention. The ADAN was delayed when it was more difficult to derive cue meaning. Finally, the data suggested an overlap of the LDAP and the EDAN.     

Interruption from irrelevant auditory and visual onsets even when attention is in a focused state

The common view on the interplay between exogenous and endogenous orienting holds that abrupt onsets are not capable of attracting attention when they occur outside the current focus of attention. Does this also apply to sudden irrelevant auditory onsets and when irrelevant visual onsets occur far in the periphery? In addition, does focused attention also reduce the alerting effect of auditory onsets, or vice versa, do highly alerting stimuli distort the attentional state? Crossmodal and unimodal variants of the Posner paradigm were examined in two experiments with targets and irrelevant onsets occurring at 28.3 and 19.3° from fixation. Either centrally presented arrows indicated the forthcoming position of visual targets to be discriminated, or warning cues signaled the likely moment of target occurrence. The targets could be preceded by peripheral auditory or visual onsets that were to be ignored. Crossmodal and unimodal exogenous orienting effects of these irrelevant onsets were observed while participants focused at the relevant side. In addition, no evidence was found that the alerting effect of auditory onsets was dependent on focused attention. Our findings indicate that, at least under the current conditions, neither crossmodal nor unimodal orienting effects of peripheral events dissipate when attention is in a focused state.     

Temporal dynamics of lateralized ERP components elicited during endogenous attentional shifts to relevant tactile events

To investigate the temporal dynamics of lateralized event-related brain potential (ERP) components elicited during covert shifts of spatial attention, ERPs were recorded in a task where central visual symbolic cues instructed participants to direct attention to their left or right hand in order to detect infrequent tactile targets presented to that hand, and to ignore tactile stimuli presented to the other hand, as well as all randomly intermingled peripheral visual stimuli. In different blocks, the stimulus onset asynchrony (SOA) between cue and target was 300 ms, 700 ms, or 1,100 ms. Anterior and posterior ERP modulations sensitive to the direction of an attentional shift were time-locked to the attentional cue, rather than to the anticipated arrival of a task-relevant stimulus. These components thus appear to reflect central attentional control rather than the anticipatory preparation of sensory areas. In addition, attentional modulations of ERPs to task-irrelevant visual stimuli were found, providing further evidence for crossmodal links in spatial attention between touch and vision.     

Visual and auditory alertness: modality-specific and supramodal neural mechanisms and their modulation by nicotine

Alertness is a nonselective attention component that refers to a state of general readiness that improves stimulus processing and response initiation. We used functional magnetic resonance imaging (fMRI) to identify neural correlates of visual and auditory alertness. A further aim was to investigate the modulatory effects of the cholinergic agonist nicotine. Nonsmoking participants were given either placebo or nicotine (NICORETTE gum, 2 mg) and performed a target-detection task with warned and unwarned trials in the visual and auditory modality. Our results provide evidence for modality-specific correlates of visual and auditory alertness in respective higher-level sensory cortices and in posterior parietal and frontal brain areas. The only region commonly involved in visual and auditory alertness was the right superior temporal gyrus. A connectivity analysis showed that this supramodal region exhibited modality-dependent coupling with respective higher sensory cortices. Nicotine was found to mainly decrease visual and auditory alertness-related activity in several brain regions, which was evident as a significant interaction of nicotine-induced decreases in BOLD signal in warned trials and increases in unwarned trials. The cholinergic drug also affected alerting-dependent activity in the supramodal right superior temporal gyrus; here the effect was the result of a significant increase of neural activity in unwarned trials. We conclude that the role of the right superior temporal gyrus is to induce an "alert" state in response to warning cues and thereby optimize stimulus processing and responding. We speculate that nicotine increases brain mechanisms of alertness specifically in conditions where no extrinsic warning is provided.