Attending to quadrants and ring-shaped regions: ERP effects of visual attention in different spatial selection tasks.

Attentional modulations of event-related brain potentials (ERPs) were measured when subjects were cued to attend to a visual quadrant or to a ring-shaped region of visual space to detect infrequently presented targets within the attended region. Spatial attention directed to quadrants was reflected in modulations of sensory-evoked P1 and N1 components at lateral posterior sites and enhanced negativities (Nds) at midline electrodes that started around 150 ms poststimulus. When attention was directed to ring-shaped regions, no modulations of P1 and N1 amplitudes were found, and Nd effects observed at midline electrodes were delayed by about 50 ms. These findings indicate that behavioral effects observed both when attention is directed to contiguous regions and to general areas of visual space may be caused by different underlying processes. Intraperceptual "sensory gating" mechanisms operating in a way suggested by the notion of an attentional "zoom-lens" may be responsible for the selection of single regions, quadrants, or hemifields. When relevant regions are more complex, spatial selectivity will affect primarily postperceptual processes.     

Reflexive attention in touch: an investigation of event related potentials and behavioural responses

Exogenous attention has been extensively studied in vision but little is known about its behavioural and neural correlates in touch. To investigate this, non-informative tactile cues were followed after 800 ms by tactile targets and participants either detected targets or discriminated their location. Responses were slowed for targets at cued compared to uncued locations (i.e. inhibition of return (IOR)) only in the detection task. Concurrently recorded ERPs showed enhanced negativity for targets at uncued compared to cued locations at the N80 component and this modulation overlapped with the P100 component but only for the detection task indicating IOR may, if anything, be linked to attentional modulations at the P100. Further, cue-target interval analysis showed an enhanced anterior negativity contralateral to the cue side in both tasks, analogous to the anterior directed attention negativity (ADAN) previously only reported during endogenous orienting.     

Selective visual attention to emotional pictures: Interactions of task-relevance and emotion are restricted to the late positive potential

The temporal dynamics of top-down voluntary and bottom-up driven emotional attention are a matter of debate. Both emotion and task-relevance have been shown to affect the early posterior negativity (EPN) and the late positive potential (LPP) during the processing of emotional pictures, whereas there are mixed findings related to interactions of emotion and task-relevance. In this preregistered study, we used Bayesian models to test this interplay between emotion and task-relevance. Participants (N = 104) were presented negative, neutral, and positive International Affective Picture System (IAPS) pictures, and block-wise were asked to respond either to negative, neutral, or positive pictures, rendering one stimulus category task-relevant and the remaining stimuli task-irrelevant. Bayesian models showed evidence for the absence of interactions between task-relevance and emotion. Furthermore, models showed parallel emotion and task effects for the EPN and late stages of the LPP. Additional interactive effects were found during an early LPP interval (400 to 600 ms), with increased LPP amplitudes when emotion was rendered task-relevant. Taken together, the results revealed distinct but parallel temporal onsets of task-relevance and emotion effects, followed by task and emotion interactions in the early LPP. Thus, based on this high-powered study employing Bayesian analysis, evidence is provided that interactions between task-relevance and emotion do not emerge before the LPP time window. These results further inform theories proposing early parallel and late interactive processes of the emotional salience of stimuli and top-down attention.     

Intermodal spatial attention differs between vision and audition: an event-related potential analysis

Subjects were required to attend to a combination of stimulus modality (vision or audition) and location (left or right). Intermodal attention was measured by comparing event-related potentials (ERPs) to visual and auditory stimuli when the modality was relevant or irrelevant, while intramodal (spatial) attention was measured by comparing ERPs to visual and auditory stimuli presented at relevant and irrelevant spatial locations. Intramodal spatial attention was expressed differently in visual and auditory ERPs. When vision was relevant, spatial attention showed a contralateral enhancement of posterior N1 and P2 components and enhancement of parietal P3. When audition was relevant, spatial attention showed a biphasic fronto-central negativity, starting after around 100 ms. The same effects were also present in ERPs to stimuli that were presented in the irrelevant modality. Thus, spatial attention was not completely modality specific. Intermodal attention effects were also expressed differently in vision and audition. Taken together, the obtained ERP patterns of the present study show that stimulus attributes such as modality and location are processed differently in vision and audition.     

Varieties of attention in neutral trials: linking RT to ERPs and EEG frequencies

It is often assumed that when a neutral cue is presented in a spatial cueing task, attention remains at fixation until target onset. We hypothesized that variance in nonspatial attention and switches of attention toward target locations can account for variance in reaction times of neutral trials. Lateralized event-related potentials (ERPs) and changes in electroencephalogram (EEG) frequency bands served as predictor variables in a single-trial logistic regression analysis to predict the direction of spatial attention in cued and neutral trials. The contingent negative variation (CNV) and non-lateralized changes in the alpha band served as markers of nonspatial attention. The direction of attention in cued trials was reliably predicted from single-trial lateralized ERP components. In neutral trials, only evidence for nonspatial attention was found, indicated by increases in the CNV and decreases in alpha preceding targets to which responses were relatively fast.     

Age-related frontoparietal changes during the control of bottom-up and top-down attention: an ERP study

We investigated age-related changes in frontal and parietal scalp event-related potential (ERP) activity during bottom-up and top-down attention. Younger and older participants were presented with arrays constructed to induce either automatic “pop-out” (bottom-up) or effortful “search” (top-down) behavior. Reaction times (RTs) increased and accuracy decreased with age, with a greater age-related decline in accuracy for the search than for the pop-out condition. The latency of the P300 elicited by the visual search array was shorter in both conditions in the younger than in the older adults. Pop-out target detection was associated with greater activity at parietal than at prefrontal locations in younger participants and with a more equipotential prefrontal-parietal distribution in older adults. Search target detection was associated with greater activity at prefrontal than at parietal locations in older relative to younger participants. Thus, aging was associated with a more prefrontal P300 scalp distribution during the control of bottom-up and top-down attention. Early latency extrastriate potentials were enhanced and N2-posterior-contralateral (N2pc) was reduced in the older group, supporting the idea that the frontal enhancements may be due to a compensation for disinhibition and distraction in the older adults. Taken together these findings provide evidence that younger and older adults recruit different frontal-parietal networks during top-down and bottom-up attention, with older adults increasing their recruitment of a more frontally distributed network in both of these types of attention. This work is in accord with previous neuroimaging findings suggesting that older adults recruit more frontal activity in the service of a variety of tasks than younger adults.     

Reducing attentional capture of emotion by broadening attention: increased global attention reduces early electrophysiological responses to negative stimuli

Decades of research has shown the influence of emotion on attentional capture, and more recently, the influence of emotion on neurophysiological processes related to attentional capture. The current research tested whether some of the earliest neurophysiological underpinnings of emotive attentional processes can be influenced by attentional manipulations of broadening versus narrowing. Previous research has shown that negative affects high in motivational intensity (e.g., disgust, fear) cause a relative narrowing of attentional scope (Gable and Harmon-Jones, 2010a; Easterbrook, 1959). Because of the strong link between motivation and attention, attentional scope should also influence the attentional capture of negative stimuli. The current study manipulated a local attentional scope or global attentional scope, then measured attentional capture towards disgust and neutral pictures using the N1 event-related potential component. Results revealed that a manipulated global attentional scope reduced N1 amplitude towards disgust pictures compared to a manipulated local attentional scope.     

Attending to the heart is associated with posterior alpha band increase and a reduction in sensitivity to concurrent visual stimuli

Attentional mechanisms have been studied mostly in specific sensory domains, such as auditory, visuospatial, or tactile modalities. In contrast, attention to internal interoceptive visceral targets has only recently begun to be studied, despite its potential importance in emotion, empathy, and self‐awareness. Here, we studied the effects of shifting attention to the heart using a cue‐target detection paradigm during continuous EEG recordings. Subjects were instructed to count either a series of visual stimuli (visual condition) or their own heartbeats (heart condition). Visual checkerboard stimuli were used as attentional probes throughout the task. Consistent with previous findings, attention modulated the amplitude of the heartbeat‐evoked potentials. Directing attention to the heart significantly reduced the visual P1/N1 amplitude evoked by the attentional probe. ERPs locked to the attention‐directing cue revealed a novel frontal positivity around 300 ms postcue. Finally, spectral power in the alpha band over parieto‐occipital regions was higher while attending to the heart—when compared to the visual task—and correlated with subject's performance in the interoceptive task. These results are consistent with a shared, resource‐based attentional mechanism whereby allocating attention to bodily signals can affect early responses to visual stimuli.     

Allocation of attention to self-name and self-face: An ERP study

Self-related information, due to its high social/adaptive value, seems to have a preferential access to our attentional resources (cf. the cocktail party effect). However, it remains uncertain whether this attention preference is the same for different kinds of self-related cues. In this ERP study we showed that self-name and self-face when compared with other names and faces, produced very similar patterns of behavioral and neural responses, i.e., shorter reaction times (RTs) and enhanced P300. The processing of the two self-related cues did not differ between each other, neither in RTs nor in P300 responses. In fact, the amplitudes of P300 to self-name and self-face were correlated. These results suggest that the adaptive value of different kinds of self-related cues tends to be equal and they engage attention resources to a similar extent.     

Filtering performance in visual working memory is improved by reducing early spatial attention to the distractors

We investigated the underlying processes that enable improving filtering irrelevant items from entering visual working memory (WM). To this end, participants performed a bilateral change‐detection task in which either targets or targets along with distractors (i.e., the filtering condition) appeared in the memory array while ERPs were recorded. In the cue‐present condition, we provided a spatial cue coupled with a temporal cue regarding where and when the distractors would appear. On some of the filtering trials, after the offset of the memory array, task‐irrelevant probes were briefly flashed either at the locations of the targets or at the locations of the distractors. This enabled measuring whether reactivating the filtering settings resulted in reducing spatial attentional resources to the distractors, allocating additional spatial attentional resources to the targets, or both, as was measured by the P1/N1 amplitude. Results revealed that, relative to the cue‐absent condition, in the cue‐present condition the P1/N1 amplitude was reduced for probes at the distractors and was similar for probes at the targets. In addition, the reduction in the P1/N1 amplitude was accompanied by a reduced filtering cost in accuracy performance in the cue‐present condition relative to the cue‐absent condition. These findings suggest that reactivating the distractor filtering settings improved filtering performance in visual WM by reducing the allocation of spatial attention to the distractors already at early processing stages, and not by allocating additional spatial attentional resources to the targets.     

The effect of category learning on visual attention and visual representation

Subordinate‐level category learning recruits neural resources associated with perceptual expertise, including the N250 component of the ERP, a posterolateral negative wave maximal between 230 and 330 ms. The N250 is a relatively late visual ERP and could plausibly be driven by attention to the features of categorized objects. Indeed, it has a latency and scalp distribution similar to the selection negativity (SN), an ERP component long known to be sensitive to attentional selection of target features. To clarify sensitivity of the N250 to attention and to more generally investigate the effect of category learning on attentional modulation of learned features, we independently manipulated subordinate‐level category learning and target detection in a speeded paradigm designed to optimally elicit the SN and accompanying frontal selection positivity (FSP). Participants first practiced categorizing a set of artificial animal stimuli and then performed a speeded target detection task on trained and untrained stimuli while ERPs were recorded. SN and FSP were roughly linearly related to the number of target features in the stimulus. Trained stimuli elicited a significantly larger N250 than untrained stimuli. The SN and N250 effects were additive, with all levels of target similarity equally affected by training, and had different time courses. Training had little effect on the FSP. The results suggest that (a) the N250 and SN have different sources, and (b) at the very least, the learning‐induced N250 indexes a different attentional subprocess from the target‐induced SN and could be driven by a different cognitive process altogether.     

Early interaction between perceptual load and involuntary attention: An event-related potential study

Whether selective attention affects C1, the first (earliest) visual cortical component of the event-related potential (ERP), remains controversial. We used a cued, involuntary attention task requiring discrimination of targets under low and high levels of perceptual load to examine early attentional modulation in visual cortex. Potential confounds due to physical stimulus differences between load conditions and cue–target sensory interaction were minimized. An interaction between perceptual load and involuntary attention was observed for the P1m component (peak latency between 100 and 140 ms). Furthermore, the parieto-central C1 component (peak latency 80 ms) was modulated by attention, but only under the high-load condition. Thus, whereas attention typically modulates the later P1 component, attentional modulation of C1 is possible under optimal conditions. Specifically, a high perceptual load is necessary for eliciting this earliest attentional effect on cortical processing.     

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.     

Perceptual factors affecting age-related differences in focused attention: Performance and psychophysiological analyses

In this study, the impact of two perceptual factors, feature similarity and spacing, on age-related differences in performance and psychophysiological measures were investigated within a focused attention paradigm. Young and old subjects performed an Eriksen letter identification task, in which centrally presented targets were flanked by response-compatible or response-incompatible letters. In feature similarity conditions, targets and flankers had a low or high amount of feature overlap. In spacing conditions, flankers were presented at four different lateral positions from the target. In the condition with high feature overlap and shortest target-flanker distance, old subjects showed greater interference by incompatible flankers than young subjects. Feature similarity was of little influence on age-related differences. However, spacing turned out to be of critical importance. Age-related interference effects disappeared when the target-flanker distance increased. This appears to be due to a decrease in response competition.     

Testing the top-down contingent capture of attention for abrupt-onset cues: Evidence from cue-elicited N2pc

Many studies using N2pc as a marker of attentional capture have demonstrated top-down contingent capture for salient color singletons: Among all salient cues that are not predictive of the target location, only cues similar to searched-for target features, and thus, matching to the top-down attentional control settings capture attention. This is reflected in matching cue's elicitation of an N2pc and a cueing effect in behavior, and the absence of the corresponding effects for non-matching cues (with features dissimilar to that of the searched-for targets). Yet, with abrupt-onset cues, corresponding evidence is missing, inviting speculations about the potential of abrupt-onset cues to capture attention followed by quick suppression within the target displays. Here, we used two types of abrupt-onset cues to test if capture by such cues also adheres to the contingent-capture principle: matching abrupt-onset cues with a color similar to the top-down control settings and non-matching abrupt-onset cues with a color different from all searched-for targets. With the help of these cues, top-down contingent capture was supported. Only matching abrupt-onset cues elicited an N2pc and a behavioral cueing effect. Depending on the exact side conditions, non-matching cues either elicited no N2pc or a P_D (i.e., evidence of active suppression). Results are discussed against the background of competing theories on attention capture by abrupt-onset cues.     

Hemodynamic responses to fear and disgust-inducing pictures: an fMRI study

The majority of neuroimaging studies on affective processing have indicated that there are specific brain structures, which are selectively responsive to fear and disgust. Whereas the amygdala is assumed to be fear-related, the insular cortex is most likely involved in disgust processing. Since these findings are mainly a result of studies focusing exclusively either on fear, or on disgust, but rarely on both emotions together, the present experiment explored the neural effects of viewing disgusting and fear-inducing pictures in contrast to neutral pictures. This was done by means of functional magnetic resonance imaging (fMRI) with 19 subjects (nine males, ten females), who also gave affective ratings for the presented pictures. The fear and the disgust pictures were able to induce the target emotions and they received comparable valence and arousal ratings. The processing of both aversive picture types was associated with an increased brain activation in the occipital–temporal lobe, in the prefrontal cortex, and in the thalamus. The amygdala was significantly activated by disgusting, but not by fear-inducing, pictures. Thus, our data are in contrast with the idea of highly emotion-specific brain structures and rather suggest the existence of a common affective circuit.     

Auditory selective attention in young and elderly adults: the selection of single versus conjoint features

The effects of aging on the behavioral and neurophysiological correlates of auditory selective attention were investigated when selection was between either unidimensional or multidimensional stimuli. Attentional processes were studied by recording event-related brain potentials (ERPs) from 16 young (M = 22 years) and 16 elderly adults (M = 74 years) while they detected target tones based on a single location cue or a conjunction of location and pitch cues. Performance declined from the single- to the conjoint-cue task for both age groups but more so for the elderly. The ERP data showed that both age groups used a hierarchical processing strategy to perform the conjoint-cue task, but processing of the pitch dimension took longer for the elderly than for the young. The ERP data also showed that the scalp distribution of a late aspect of the waveform was more restricted in both anterior and posterior directions for the elderly. This suggests that frontal-lobe dependent attentional processes may be less efficacious with aging.     

N1 and P2 to words and wordlike stimuli in late elementary school children and adults

In an investigation of the development of fine‐tuning for word processing across the late elementary school years as indexed by the posterior N1 and P2 components of the ERP waveform, third, fourth, and fifth graders and a comparison group of adults viewed words, pseudowords, nonpronounceable letter strings, and false font strings in a semantic categorization task. In adults, N1 was larger to and P2 was later to words as compared to pseudowords, a finely tuned effect of lexicality reflecting specialization for word processing. In contrast, in each group of children, N1 was larger to letter strings than false font strings and P2 was larger to false font strings than letter strings, reflecting coarse encoding for orthography. In regression analyses, scores on standardized behavioral test measures of orthographic knowledge, decoding skill, and fluency predicted N1 amplitude; these effects were not significant with age included as a separate predictor. None of the behavioral scores, in models including or not including age, predicted P2 amplitude. In direct comparisons between groups, there were multiple differences between the child and adult groups for both N1 and P2 amplitude effects, and only a single significant difference between two child groups. Overall, the findings suggest a lengthy developmental time course for the fine‐tuning of early word processing as indexed by N1 and P2.     

Distilling the distinct contralateral and ipsilateral attentional responses to lateral stimuli and the bilateral response to midline stimuli for upper and lower visual hemifield locations

A contralateral posterior negativity elicited by lateral oddballs (N2pc) and a bilateral posterior negativity elicited by vertical midline oddballs (bilateral N2) are ERP components reflecting attentional deployment that have been rarely compared. In different tasks, we explored to what extent they reflect similar underlying mechanisms of attention. We used a multiple-frame procedure to present pop-out color oddballs among distractors. A homogeneous condition contained only distractors (0 oddballs) and served as a control condition that was subtracted from oddball-present conditions to isolate attention effects. The number of oddballs and the vertical hemifield containing them (upper vs. lower) were two critical factors. For the lower hemifield, the signal amplitude increased with the number of oddballs, otherwise had similar effects and scalp distributions, suggesting the bilateral N2 acted as a bilateral N2pc and likely reflected similar underlying generators. For the upper hemifield, component amplitude also increased with the number of oddballs, but the scalp distributions were positive and more centered, suggesting inverted generators across the two vertical hemifields. An ipsilateral positivity occurred about 50 ms after a contralateral positivity, similar in magnitude, producing a biphasic contra-minus-ipsi difference wave. Previously reported smaller negative N2pc components for upper hemifield oddballs likely reflected a negative lobe artificially created by the subtraction of a lagged positive ipsilateral response. The results compel us to argue for a systematic separation of data for upper versus lower hemifields in studies of visuo-spatial attention, and the use of an experimental design permitting the separate estimation of contralateral and ipsilateral responses.     

Danger and disease: Electrocortical responses to threat- and disgust-eliciting images

Previous research suggests facilitated processing of evolutionarily significant stimuli (e.g., depictions of erotica, mutilation, threat), as reflected by augmented event-related potentials (ERPs), including the early posterior negativity (EPN) and late positive potential (LPP). Evolutionary models suggest that images that evoke disgust should be high in motivational salience, but evidence that the EPN and LPP are enhanced by disgusting images is lacking. Prior studies have employed only a small number of disgusting images that were limited in the types of content depicted. In the current study, participants viewed larger sets of disgusting, threatening, and neutral images with more varied content while electroencephalography (EEG) was recorded. Results showed that disgusting and threatening images elicited equivalent LPPs, which were both significantly increased relative to LPPs elicited by neutral images. EPN amplitudes were augmented for both disgusting and threatening relative to neutral images, though significantly more for disgust. These findings offer initial evidence that the EPN and the LPP are sensitive to disgust-eliciting pictures and that these pictures may receive processing that is at least on par with that of threatening images. Limitations of the current study and implications for future research are discussed.