Anterior Directing Attention Negativity Research Articles

Acute hypoxia alters visuospatial attention orienting: an electrical neuroimaging study

Our study investigated the effects of hypoxia on visuospatial attention processing during preparation for a single/double-choice motor response. ERPs were recorded in two sessions in which participants breathed either ambient-air or oxygen-impoverished air. During each session, participants performed four cue-target attention orienting and/or alerting tasks. Replicating the classic findings of valid visuospatial attentional orienting modulation, ERPs to pre-target cues elicited both an Anterior directing attention negativity (ADAN)/CNV and a posterior Late directing attention positivity (LDAP)/TP, which in ambient air were larger for attention orienting than for alerting. Hypoxia increased the amplitude of both these potentials in the spatial orienting conditions for the upper visual hemifield, while, for the lower hemifield, it increased ADAN/CNV, but decreased LDAP/TP for the same attention conditions. To these ERP changes corresponded compensatory enhanced activation of right anterior cingulate cortex, left superior parietal lobule and frontal gyrus, as well as detrimental effects of hypoxia on behavioral overt performance. Together, these findings reveal for the first time, to our knowledge, that (1) these reversed alterations of the activation patterns during the time between cue and target occur at a larger extent in hypoxia than in air, and (2) acute normobaric hypoxia alters visuospatial attention orienting shifting in space.

The Electrophysiological Markers of Statistically Learned Attentional Enhancement: Evidence for a Saliency-based Mechanism.

It is well established that attention can be sharpened through the process of statistical learning (e.g., visual search becomes faster when targets appear at high-relative-to-low probability locations). Although this process of statistically learned attentional enhancement differs behaviorally from the well-studied top-down and bottom-up forms of attention, relatively little work has been done to characterize the electrophysiological correlates of statistically learned attentional enhancement. It thus remains unclear whether statistically learned enhancement recruits any of the same cognitive mechanisms as top-down or bottom-up attention. In the current study, EEG data were collected from while participants searched for an ambiguous unique shape in a visual array (the additional singleton task). Unbeknown to the participants, targets appeared more frequently in one location in space (probability cuing). Encephalographic data were then analyzed in two phases: an anticipatory phase and a reactive phase. In the anticipatory phase preceding search stimuli onset, alpha lateralization as well as the Anterior Directing Attention Negativity and Late Directing Attention Positivity components-signs of preparatory attention known to characterize top-down enhancement-were tested. In the reactive phase, the N2pc component-a well-studied marker of target processing-was examined following stimuli onset. Our results showed that statistically learned attentional enhancement is not characterized by any of the well-known anticipatory markers of top-down attention; yet targets at high probability locations did reliably evoke larger N2pc amplitudes, a finding that is associated with bottom-up attention and saliency. Overall, our findings are consistent with the notion that statistically learned attentional enhancement increases the perceptual salience of items appearing at high-probability locations relative to low-probability locations.

Spatial Attention in Serial Order Working Memory: An EEG Study.

Theoretical models explaining serial order processing link order information to specified position markers. However, the precise characteristics of position marking have remained largely elusive. Recent studies have shown that space is involved in marking serial position of items in verbal working memory (WM). Furthermore, it has been suggested, but not proven, that accessing these items involves horizontal shifts of spatial attention. We used continuous electroencephalography recordings to show that memory search in serial order verbal WM involves spatial attention processes that share the same electrophysiological signatures as those operating on the visuospatial WM and external space. Accessing an item from a sequence in verbal WM induced posterior "early directing attention negativity" and "anterior directing attention negativity" contralateral to the position of the item in mental space (i.e., begin items on the left; end items on the right). In the frequency domain, we observed posterior alpha suppression contralateral to the position of the item. Our results provide clear evidence for the involvement of spatial attention in retrieving serial information from verbal WM. Implications for WM models are discussed.

Retroactive Attentional Shifts Predict Performance in a Working Memory Task: Evidence by Lateralized EEG Patterns.

Shifts of attention within working memory based on retroactive (retro-) cues were shown to facilitate performance in working memory tasks. Although posterior asymmetries in the EEG, such as the contralateral delay activity (CDA), have been used to study the active storage of lateralized working memory representations, results on the relation of such asymmetric effects to retro-cue benefits remain inconclusive. We recorded EEG in a retro-cue working memory task with lateralized items and a continuous performance response. Following either a selective or neutral retro-cue, participants adjusted the orientation of a central memory probe to the cued item. Selective retro-cues elicited an early posterior contralateral negativity (PCN), anterior directing attention negativity (ADAN) and a later modulation of CDA indicating that active storage was concentrated on the cued information. By dividing all trials into three within-condition performance quantiles, we could further show that high working memory accuracy was associated with a sustained increase of the CDA effect following the retro-cue. These results suggest that focusing resources on the active storage of relevant representations is an important factor regarding retro-cue benefits in working memory tasks.

EEG Correlates of Preparatory Orienting, Contextual Updating, and Inhibition of Sensory Processing in Left Spatial Neglect.

Studies with event-related potentials have highlighted deficits in the early phases of orienting to left visual targets in right-brain-damaged patients with left spatial neglect (N+). However, brain responses associated with preparatory orienting of attention, with target novelty and with the detection of a match/mismatch between expected and actual targets (contextual updating), have not been explored in N+. Here in a study in healthy humans and brain-damaged patients of both sexes we demonstrate that frontal activity that reflects supramodal mechanisms of attentional orienting (Anterior Directing Attention Negativity, ADAN) is entirely spared in N+. In contrast, posterior responses that mark the early phases of cued orienting (Early Directing Attention Negativity, EDAN) and the setting up of sensory facilitation over the visual cortex (Late Directing Attention Positivity, LDAP) are suppressed in N+. This uncoupling is associated with damage of parietal-frontal white matter. N+ also exhibit exaggerated novelty reaction to targets in the right side of space and reduced novelty reaction for those in the left side (P3a) together with impaired contextual updating (P3b) in the left space. Finally, we highlight a drop in the amplitude and latency of the P1 that over the left hemisphere signals the early blocking of sensory processing in the right space when targets occur in the left one: this identifies a new electrophysiological marker of the rightward attentional bias in N+. The heterogeneous effects and spatial biases produced by localized brain damage on the different phases of attentional processing indicate relevant functional independence among their underlying neural mechanisms and improve the understanding of the spatial neglect syndrome.SIGNIFICANCE STATEMENT Our investigation answers important questions: are the different components of preparatory orienting (EDAN, ADAN, LDAP) functionally independent in the healthy brain? Is preparatory orienting of attention spared in left spatial neglect? Does the sparing of preparatory orienting have an impact on deficits in reflexive orienting and in the assignment of behavioral relevance to the left space? We show that supramodal preparatory orienting in frontal areas is entirely spared in neglect patients though this does not counterbalance deficits in preparatory parietal-occipital activity, reflexive orienting, and contextual updating. This points at relevant functional dissociations among different components of attention and suggests that improving voluntary attention in N+ might be behaviorally ineffective unless associated with stimulations boosting the response of posterior parietal-occipital areas.

How microsaccades relate to lateralized ERP components of spatial attention: A co-registration study

Covert shifts of attention that follow the presentation of a cue are associated with lateralized components in the event-related potential (ERP): the “early directing attention negativity” (EDAN) and the “anterior directing attention negativity” (ADAN). Traditionally, these shifts are thought to take place while gaze is fixated and, thus, in the absence of saccades. However, microsaccades of small amplitude (<1°) occur frequently and involuntarily also during fixation and are closely correlated with spatial attention. To investigate potential links between microsaccades and lateralized ERP components, we simultaneously recorded eye movements and ERPs in a spatial cueing task. As a first major result, we show that both the posterior EDAN and the orientation of microsaccades align more strongly with the location of the task-relevant part of the cue stimulus than with the direction of the attention shift indicated by that cue. A coupling between microsaccades and EDAN was also present on the single-trial level: The EDAN was largest when microsaccades were oriented toward the relevant cue, but absent when microsaccades were oriented away from it, suggesting that EDAN and microsaccades are generated by the same neural network, which selects relevant stimuli and orients behavior toward them. As a second major result, we show that small corneoretinal artifacts from microsaccades, which fall below conventional EOG rejection thresholds, contaminate the measurement of the ADAN. After correcting the EEG for microsaccade-related artifacts with an optimized variant of independent component analysis, ADAN was abolished at frontal sites, but a genuine ADAN was still present at central sites. Thus, the combined measurement of microsaccades and lateralized ERPs sheds new light onto cue-elicited shifts of covert attention.

Interplay between supramodal attentional control and capacity limits in the low-level visual processors modulate the tendency to inattention.

When engaged in a demanding task, individuals may neglect unexpected visual stimuli presented concomitantly. Here we use a change detection task to show that propensity to inattention is associated with a flexible allocation of attentional resources to filter and represent visual information. This was reflected by N2 posterior contralateral (N2pc) and contralateral delay activity (CDA) respectively, but also during high-order reorienting of attentional resources (known as anterior directing attention negativity, ADAN). Results show that differences in noticing and failing to notice unexpected stimuli/changes are associated with different patterns of brain activity. When processing (N2) and working memory (CDA) capacities are low, resources are mostly allocated to small set-sizes and associated with a tendency to filter information during early low-level processing (N2). When resources are high, saturation is obtained with larger set-sizes. This is also associated to a tendency to select (N2) and reorient resources (ADAN) to maintain extra information (CDA).

Neural activity associated with attention orienting triggered by implied action cues

Spatial attention can be directed by the actions of others. We used ERPs method to investigate the neural underpins associated with attention orienting which is induced by implied body action. Participants performed a standard non-predictive cuing task, in which a directional implied action (throwing and running) or non-action (standing) cue was randomly presented and then followed by a target to the left or right of the central cue, despite cue direction. The cue-triggered ERPs results demonstrated that implied action cues, rather than the non-action cue, could shift the observers' spatial attention as demonstrated by the robust anterior directing attention negativity (ADAN) effects in throwing and running cues. Further, earlier N1 (100–170ms) and P2 (170–260ms) waveform differences occurred between implied action and non-action cues over posterior electrodes. The P2 component might reflect implied motion signal perception of implied action cues, and this implied motion perception might play an important role in facilitating the attentional shifts induced by implied action cues. Target-triggered ERPs data (mainly P3a component) indicated that implied action cues (throwing and running) speeded and enhanced the responses to valid targets compared to invalid targets. Furthermore, P3a might imply that implied action orienting may share similar mechanisms of action with voluntary attention, especially at the novel stimuli processing decision-level. These results further support previous behavioral findings that implied body actions direct spatial attention and extend our understanding about the nature of the attentional shifts that are elicited by implied action cues.

Cue-locked lateralized components in a tactile spatial attention task: Evidence for a functional dissociation between ADAN and LSN.

ERP studies investigating the control processes responsible for spatial orienting in touch have consistently observed that the anterior directing attention negativity (ADAN) elicited by an attention-directing cue is followed by a sustained negativity contralateral to the cued hand. Recent evidence suggested that the later negativity, labeled late somatotopic negativity (LSN), might reflect distinct neurocognitive processes from those associated with the ADAN. To investigate the functional meaning of the ADAN and LSN components, we measured ERPs elicited by bilateral tactile cues indicating to covertly shift tactile attention to the left or right hand. Participants performed two spatial attention tasks that differed only for the difficulty of the target/nontarget discrimination at attended locations. The LSN but not the ADAN was sensitive to our experimental manipulation of task difficulty, suggesting that this component might reflect sensory-specific preparatory processes prior to a forthcoming tactile stimulus.

Normal aging selectively diminishes alpha lateralization in visual spatial attention

EEG studies of cue-induced visual alpha power (8–13Hz) lateralization have been conducted on young adults without examining differences that may develop as a consequence of normal aging. Here, we examined age-related differences in spatial attention by comparing healthy older and younger adults. Our key finding is that cue-induced alpha power lateralization was observed in younger, but not older adults, even though both groups exhibited classic event-related potential signatures of spatial orienting. Specifically, both younger and older adults showed significant early directing-attention negativity (EDAN), anterior directing-attention negativity (ADAN), late directing-attention positivity (LDAP) and contingent negative variation (CNV). Furthermore, target-evoked sensory components were enhanced for attended relative to unattended targets in both younger and older groups. This pattern of results suggests that although older adults can successfully allocate spatial attention, they do so without the lateralization of alpha power that is commonly observed in younger adults. Taken together, our findings demonstrate that younger and older adults might engage different neural mechanisms for attentional orienting, and that alpha power lateralization during visual spatial attention is a phenomenon that diminishes during normal aging.

Attention Allocation and Task Representation during Joint Action Planning

We investigated whether people take into account an interaction partner's attentional focus and whether they represent in advance their partner's part of the task when planning to engage in a synchronous joint action. The experiment involved two participants planning and performing joint actions (i.e., synchronously lifting and clinking glasses), unimanual individual actions (i.e., lifting and moving a glass as if clinking with another person), and bimanual individual actions. EEG was recorded from one of the participants. We employed a choice reaction paradigm where a visual cue indicated the type of action to be planned, followed 1.5 sec later by a visual go stimulus, prompting the participants to act. We studied attention allocation processes by examining two lateralized EEG components, namely the anterior directing attention negativity and the late directing attention positivity. Action planning processes were examined using the late contingent negative variation and the movement-related potential. The results show that early stages of joint action planning involve dividing attention between locations in space relevant for one's own part of the joint action and locations relevant for one's partner's part of the joint action. At later stages of joint action planning, participants represented in advance their partner's upcoming action in addition to their own action, although not at an effector-specific level. Our study provides electrophysiological evidence supporting the operation of attention sharing processes and predictive self/other action representation during the planning phase of a synchronous joint task.

Eye gaze triggers reflexive attention shifts: Evidence from lateralised ERPs

Social cues, such as another individual׳s eye gaze, provide valuable information regarding the actions and intentions of others. Previous studies have suggested that seeing another׳s gaze automatically orients one׳s attention in the gaze direction. In this event-related potential (ERP) study, a spatial cuing paradigm was combined with a visual search task in which targets were defined by feature conjunctions in order to eliminate effects of target/distractor salience. Participants viewed centrally presented faces with neutral expressions in which eyes looked to the left or right. The participants׳ task was to identify a target object (with or without gap) defined by a combination of shape and orientation, which appeared in either the same (cued) or the opposite (uncued) location as the direction of the eye gaze. There was behavioural evidence of a gaze congruency effect, as reaction times (RTs) were faster when the eyes looked towards the target rather than away from the location of the target. The ERP data indicated the presence of significant gaze-congruent early directing attention negativity (EDAN) and anterior directing attention negativity (ADAN), reflecting attention shifts to the cued location in advance of the target presentation. ERP data did not show evidence of later orienting of attention triggered by gaze cues in the late attention-directing attention positivity (LDAP) at posterior sites. The results disclosed the neural response during reflexive attention shifting triggered by gazes and ascertained the relationship among EDAN, ADAN, LDAP and gaze-elicited attention shifts. After the presentation of the target array without salient stimuli, the presence of the N2-posterior-contralateral (N2pc) in the cued trials and the absence in the uncued trials further supported that attention had been directed to the possible target location prior to the target onset. The ERPs in response to the target array also extend our understanding of the neural response that orients spatial attention by providing valuable information about the temporal dynamics without the influence of salience.

Autistic traits influence gaze-oriented attention to happy but not fearful faces

The relationship between autistic traits and gaze-oriented attention to fearful and happy faces was investigated at the behavioral and neuronal levels. Upright and inverted dynamic face stimuli were used in a gaze-cueing paradigm while event related potentials (ERPs) were recorded. Participants responded faster to gazed-at than to non-gazed-at targets, and this gaze orienting effect (GOE) diminished with inversion, suggesting it relies on facial configuration. It was also larger for fearful than happy faces but only in participants with high autism-spectrum quotient (AQ) scores. While the GOE to fearful faces was of similar magnitude regardless of AQ scores, a diminished GOE to happy faces was found in participants with high AQ scores.At the ERP level, a congruency effect on target-elicited P1 component reflected enhanced visual processing of gazed-at targets. In addition, cue-triggered early directing attention negativity and anterior directing attention negativity reflected, respectively, attention orienting and attention holding at gazed-at locations. These neural markers of spatial attention orienting were not modulated by emotion and were not found in participants with high AQ scores. Together, these findings suggest that autistic traits influence attention orienting to gaze and its modulation by social emotions such as happiness.

Age Group and Individual Differences in Attentional Orienting Dissociate Neural Mechanisms of Encoding and Maintenance in Visual STM

Selective attention biases the encoding and maintenance of representations in visual STM (VSTM). However, precise attentional mechanisms gating encoding and maintenance in VSTM and across development remain less well understood. We recorded EEG while adults and 10-year-olds used cues to guide attention before encoding or while maintaining items in VSTM. Known neural markers of spatial orienting to incoming percepts, that is, Early Directing Attention Negativity, Anterior Directing Attention Negativity, and Late Directing Attention Positivity, were examined in the context of orienting within VSTM. Adults elicited a set of neural markers that were broadly similar in preparation for encoding and during maintenance. In contrast, in children these processes dissociated. Furthermore, in children, individual differences in the amplitude of neural markers of prospective orienting related to individual differences in VSTM capacity, suggesting that children with high capacity are more efficient at selecting information for encoding into VSTM. Finally, retrospective, but not prospective, orienting in both age groups elicited the well-known marker of visual search (N2pc), indicating the recruitment of additional neural circuits when orienting during maintenance. Developmental and individual differences differentiate seemingly similar processes of orienting to perceptually available representations and to representations held in VSTM.

Fearful, surprised, happy, and angry facial expressions modulate gaze-oriented attention: Behavioral and ERP evidence

The impact of emotions on gaze-oriented attention was investigated in non-anxious participants. A neutral face cue with straight gaze was presented, which then averted its gaze to the side while remaining neutral or expressing an emotion (fear/surprise in Exp.1 and anger/happiness in Exp.2). Localization of a subsequent target was faster at the gazed-at location (congruent condition) than at the non-gazed-at location (incongruent condition). This Gaze-Orienting Effect (GOE) was enhanced for fear, surprise, and anger, compared to neutral expressions which did not differ from happy expressions. In addition, Event Related Potentials (ERPs) to the target showed a congruency effect on P1 for fear and surprise and a left lateralized congruency effect on P1 for happy faces, suggesting that target visual processing was also influenced by attention to gaze and emotions. Finally, at cue presentation, early postero-lateral (Early Directing Attention Negativity (EDAN)) and later antero-lateral (Anterior Directing Attention Negativity (ADAN)) attention-related ERP components were observed, reflecting, respectively, the shift of attention and its holding at gazed-at locations. These two components were not modulated by emotions. Together, these findings show that the processing of social signals such as gaze and facial expression interact rather late and in a complex manner to modulate spatial attention.

ERP evidence for selective drop in attentional costs in uncertain environments: Challenging a purely premotor account of covert orienting of attention

Several studies have proved that the reliability of endogenous spatial cues linearly modulates the reaction time advantage in the processing of targets at validly cued vs. invalidly cued locations, i.e. the “validity effect”. This would imply that with non-predictive cues, no “validity effect” should be observed. However, contrary to this prediction, one could hypothesize that attentional benefits by valid cuing (i.e. the RT advantage for validly vs. neutrally cued targets) can still be maintained with non-predictive cues, if the brain were endowed with mechanisms allowing the selective reduction in costs of reorienting from invalidly cued locations (i.e. the reduction of the RT disadvantage for invalidly vs. neutrally cued targets). This separated modulation of attentional benefits and costs would be adaptive in uncertain contexts where cues predict at chance level the location of targets. Through the joint recording of manual reaction times and event-related cerebral potentials (ERPs), we have found that this is the case and that relying on non-predictive endogenous cues results in abatement of attentional costs and the difference in the amplitude of the P1 brain responses evoked by invalidly vs. neutrally cued targets. In contrast, the use of non-predictive cues leaves unaffected attentional benefits and the difference in the amplitude of the N1 responses evoked by validly vs. neutrally cued targets. At the individual level, the drop in costs with non-predictive cues was matched with equivalent lateral biases in RTs to neutrally and invalidly cued targets presented in the left and right visual field. During the cue period, the drop in costs with non-predictive cues was preceded by reduction of the Early Directing Attention Negativity (EDAN) on posterior occipital sites and by enhancement of the frontal Anterior Directing Attention Negativity (ADAN) correlated to preparatory voluntary orienting. These findings demonstrate, for the first time, that the segregation of mechanisms regulating attentional benefits and costs helps efficiency of orienting in “uncertain” visual spatial contexts characterized by poor probabilistic association between cues and targets.

Event-related potential evidence for the use of external coordinates in the preparation of tactile attention by the early blind

Recent studies have suggested that visual experience in childhood is crucial for the automatic activation of an external spatial reference frame in tactile perception. These findings are largely based on behavioural work, with limited exploration using event-related potentials (ERPs). The present study examined the role of external spatial frameworks on tactile perception by recording ERP correlates of both preparatory processes and somatosensory processing during a tactile attention task for a group of early blind participants and age-matched sighted controls who carried out the task in darkness. Participants had to shift attention to one hand or the other as indicated by an auditory cue presented at the start of each trial, in order to detect infrequent tactile targets delivered to the attended hand. Spatial information about the external environment was acquired in advance during tactile exploration of the testing booth. ERPs measured during the cue-target interval indicated a conflict between anatomical and external spatial reference frames for both early blind and sighted participants, as marked by the delayed onset of the anterior directing attention negativity, although the delay was more pronounced in the sighted. A delay was also observed, irrespective of visual experience, on the onset of attentional modulations of somatosensory ERPs elicited by tactile stimuli. Although these results confirm that neither concurrent nor developmental vision is necessary for the default use of an external spatial framework in tactile attention, they suggest that the relative impact of an external vs. an anatomical spatial coordinate system may be affected by visual experience.

Neural activity associated with attention orienting triggered by gaze cues: A study of lateralized ERPs

This study investigated the electrophysiological (event-related potential, ERP) and behavioral (reaction time, RT) correlates of gaze-cued shifts of visuospatial attention. Participants viewed centrally presented faces with neutral expressions in which eyes looked straight ahead and then shifted to the left or right. Upon movement of the eyes, the facial expression either stayed the same (neutral) or changed to a fearful or happy expression. Participants' task was to identify a probe letter (T or L), which appeared in either the same or the opposite location to the direction of the eye gaze. There was behavioral evidence of a gaze congruency effect, as RTs were faster when the eyes looked toward rather than away from the location of the target. The ERP data indicated the presence of a significant gaze-congruent anterior directing attention negativity (ADAN) at anterior locations (300–500 ms after the onset of the gaze cue). ERP data did not show evidence of initial orienting of attention triggered by gaze cues in the early directing attention negativity (EDAN) at posterior locations (200–300 ms post-cue onset). The gaze cueing effects in the RT and ERP data were not significantly influenced by the emotional expression of the faces. The presence of the ADAN may reflect neural mechanisms that underlie the holding of attention on gazed-at locations.

An early parietal ERP component of the frontoparietal system: EDAN ≠ N2pc

Directing spatial attention or manual response selection by means of arrow cues elicits a characteristic sequence of lateralized ERP components. Van Velzen and Eimer (2003) proposed that one of these components, the EDAN (early directing-attention negativity) is not related to the control of spatial attention but is instead an N2pc in disguise, related to the visual processing of arrow cues. While this proposal seems widely accepted, it has never been evaluated whether the hitherto not well-determined scalp distribution of the EDAN matches the well-established scalp distribution of the N2pc. This paper reviews evidence from earlier published work supporting a parietal distribution of the EDAN. This strongly argues against the proposed identification with the N2pc. The EDAN may instead reflect early parietal activity within the frontoparietal network, preceding the frontal activation represented in the anterior directing-attention negativity (ADAN).

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.