On the Relevance of Posterior and Midfrontal Theta Activity for Visuospatial Attention.

The aim of this study was to examine whether oscillatory activity in the theta-band is relevant for selective visuospatial attention when there is a need for the suppression of interfering and distracting information. A variant of the Eriksen flanker task was employed with bilateral arrays: one array consisting of a target and congruent or incongruent flankers and the second array consisting of neutral distractors. The bilateral arrays were preceded either by a 100% valid spatial cue or by a neutral cue. In the cue-target interval, a major burst in medial frontal theta power was observed, which was largest in the spatial cue condition. In the latter condition, additionally a posterior theta increase was observed that was larger over sites ipsilateral to the forthcoming target array. Functional connectivity analyses revealed that this pretarget posterior theta was related to the midfrontal theta. No such effects were observed in the neutral cue condition. After onset of the bilateral arrays, a major burst in posterior theta activity was observed in both cue conditions, which again was larger above sites ipsilateral to the target array. Furthermore, this posterior theta was in all cases related to the midfrontal theta. Taken together, the findings suggest that a fronto-posterior theta network plays an important role in the suppression of irrelevant and conflicting visual information. The results also suggest that the reciprocal relation between visuospatial attention and executive response control may be closer than commonly thought.

Neural Underpinnings of Proactive and Preemptive Adjustments of Action Control.

This study aimed to trace the neural basis of proactive and preemptive adjustments of executive control and their effects on online processing of response conflict. In two EEG experiments, participants performed the flanker task with predictive cueing of conflict. The following questions were addressed: "Does conflict cueing improve performance?" We observed improved behavioral performance in the predictive condition, suggesting that participants proactively utilized the cues to prepare for the upcoming demands. "How is conflict processing affected by predictive cueing?" Conflict-related modulations of midfrontal N2 and theta power were smaller in the predictive than in the neutral condition. This suggests that proactive control suppressed the impact of incongruent flankers so that the conflict was reduced, and so was the involvement of online control. "Is proactive control implemented through preactivation of online control?" Conflict cueing increased midfrontal theta power also before target onset, suggesting preactivation of the control processes beforehand. "Do proactive and reactive control depend on common or unique processes?" Unlike the online control, the proactive control triggered a burst of theta power in the right hemisphere's dorsal and ventral lateral prefrontal cortices, connected with the midfrontal area via theta phase coherence. This indicates that the two control modes involve partially unique but coordinated neural processes. "Is preemptive control implemented through modulations of visual processing?" Predictive cueing modulated both the pretarget preparatory alpha desynchronization and the target selection-related posterior contralateral negativity (N2pc and sustained posterior contralateral negativity), in line with the hypothesis of preemptive tuning of sensory selection aimed at reducing the impact of conflicting stimuli.

Selection for Action: The Medial Frontal Cortex Is an Executive Hub for Stimulus and Response Selection.

This EEG study investigates the electrophysiological activity underlying processes of stimulus and response selection, and their executive orchestration via long-range functional connectivity under conflict condition, in order to shed more light on how these brain dynamics shape individual behavioral performance. Participants (n = 91) performed a modified flanker task, in which bilateral visual stimulation and a bimanual response pattern were employed to isolate the stimulus and response selection-related lateralized activity. First, we identified conflict-related markers of task-relevant processes; most importantly, the stimulus and response selection were evidenced by contra-ipsilateral differences in visual and motor activity, respectively, and executive control was evidenced by modulations of midfrontal activity. Second, we identified conflict-related functional connectivity between midfrontal and other task-relevant areas. The results showed that interregional phase synchronization in theta band was centered at the midfrontal site, interpreted here as a "hub" of executive communication. Importantly, the theta functional connectivity was more robust under the condition of increased demands for stimulus and response selection, including connectivity between the medial frontal cortex and the lateral frontal and motor areas, as well as cross-frequency theta-alpha coupling between the medial frontal cortex and contralateral visual areas. Third, we showed that individual differences in the measured conflict-related EEG activity, particularly the midfrontal N2, theta power, and global theta connectivity, predict the behavioral efficiency in conflict resolution.

Lateralization of spatial rather than temporal attention underlies the left hemifield advantage in rapid serial visual presentation.

In bilateral rapid serial visual presentation (RSVP), the second of two targets, T1 and T2, is better identified in the left visual field (LVF) than in the right visual field (RVF). This LVF advantage may reflect hemispheric asymmetry in temporal attention or/and in spatial orienting of attention. Participants performed two tasks: the "standard" bilateral RSVP task (Exp.1) and its unilateral variant (Exp.1 & 2). In the bilateral task, spatial location was uncertain, thus target identification involved stimulus-driven spatial orienting. In the unilateral task, the targets were presented block-wise in the LVF or RVF only, such that no spatial orienting was needed for target identification. Temporal attention was manipulated in both tasks by varying the T1-T2 lag. The results showed that the LVF advantage disappeared when involvement of stimulus-driven spatial orienting was eliminated, whereas the manipulation of temporal attention had no effect on the asymmetry. In conclusion, the results do not support the hypothesis of hemispheric asymmetry in temporal attention, and provide further evidence that the LVF advantage reflects right hemisphere predominance in stimulus-driven orienting of spatial attention. These conclusions fit evidence that temporal attention is implemented by bilateral parietal areas and spatial attention by the right-lateralized ventral frontoparietal network.

A right hemisphere advantage at early cortical stages of processing alphanumeric stimuli. Evidence from electrophysiology.

This study investigates hemispheric asymmetry evoked by non-target alphanumeric stimuli in a bilateral rapid serial visual presentation (RSVP) task. Our indicators of asymmetry are shorter latencies and larger amplitudes of the right hemisphere (RH) P1 and N1 components of visual evoked potentials (VEPs). This VEP asymmetry might reflect either a RH advantage, possibly in early perceptual processing, or for familiar stimuli, or for directing attention, or might be a paradoxical reflection of left hemisphere specialization in letter processing. Experiment 1 showed that the VEP asymmetry decreased, though remained present, with unfamiliar stimuli (Tibetan letters), as compared to familiar stimuli (Latin letters and Arabic digits). Experiment 2 showed that while leftward and rightward attentional biases affected the relation between hemispheres contra- and ipsilateral to attended visual fields, the VEP asymmetry remained independent of attention. As the most parsimonious explanation, the primary cause of the VEP asymmetry seems to be a general predominance of the RH in early perceptual processing.

Bias for the left visual field in rapid serial visual presentation: effects of additional salient cues suggest a critical role of attention.

Everyday experience suggests that people are equally aware of stimuli in both hemifields. However, when two streams of stimuli are rapidly presented left and right, the second target (T2) is better identified in the left hemifield than in the right hemifield. This left visual field (LVF) advantage may result from differences between hemifields in attracting attention. Therefore, we introduced a visual cue shortly before T2 onset to draw attention to one stream. Thus, to identify T2, attention was correctly positioned with valid cues but had to be redirected to the other stream with invalid ones. If the LVF advantage is caused by differences between hemifields in attracting attention, invalid cues should increase, and valid cues should reduce the LVF advantage as compared with neutral cues. This prediction was confirmed. ERP analysis revealed that cues evoked an early posterior negativity, confirming that attention was attracted by the cue. This negativity was earlier with cues in the LVF, which suggests that responses to salient events are faster in the right hemisphere than in the left hemisphere. Valid cues speeded up, and invalid cues delayed T2-evoked N2pc; in addition, valid cues enlarged T2-evoked P3. After N2pc, right-side T2 evoked more sustained contralateral negativity than left T2, least long-lasting after valid cues. Difficulties in identifying invalidly cued right T2 were reflected in prematurely ending P3 waveforms. Overall, these data provide evidence that the LVF advantage is because of different abilities of the hemispheres in shifting attention to relevant events in their contralateral hemifield.

Does level of processing affect the transition from unconscious to conscious perception?

Recently, Windey, Gevers, and Cleeremans (2013) proposed a level of processing (LoP) hypothesis claiming that the transition from unconscious to conscious perception is influenced by the level of processing imposed by task requirements. Here, we carried out two experiments to test the LoP hypothesis. In both, participants were asked to classify briefly presented pairs of letters as same or different, based either on the letters' physical features (a low-level task), or on a semantic rule (a high-level task). Stimulus awareness was measured by means of the four-point Perceptual Awareness Scale (PAS). The results showed that low or moderate stimulus visibility was reported more frequently in the low-level task than in the high-level task, suggesting that the transition from unconscious to conscious perception is more gradual in the former than in the latter. Therefore, although alternative interpretations remain possible, the results of the present study fully support the LoP hypothesis.

Different subjective awareness measures demonstrate the influence of visual identification on perceptual awareness ratings.

We compare four subjective awareness measures in the context of a visual identification task and investigate quantitative differences in terms of scale use and correlation with task performance. We also analyse the effect of identification task decisions on subsequent subjective reports. Results show that awareness ratings strongly predict accuracy for all scale types, although the type of awareness measure may influence the reported level of perceptual awareness. Surprisingly, the overall relationship between awareness ratings and performance was weaker when participants rated their awareness before providing identification responses. Furthermore, the Perceptual Awareness Scale was most exhaustive only when used after the identification task, whereas confidence ratings were most exhaustive when used before the identification task. We conclude that the type of subjective measure applied may influence the reports on visual awareness. We also propose that identification task decisions may affect subsequent awareness ratings.

Does CUD measure interhemispheric transfer time? The allocation of attention influences the Poffenberger effect.

A simple reaction time (SRT) difference between responses to visual stimuli presented ipsilaterally and contralaterally to the responding hand, known as the Poffenberger effect or the crossed-uncrossed difference (CUD), has been commonly interpreted as a measure of interhemispheric transfer time (IHTT). However, the validity of this interpretation and the measure's reliability have been debated. The present study aimed at obtaining reliable evidence of the influence of spatial attention on the CUD, which would provide an argument against the classical interpretation of CUD. To meet the high statistical power requirements, over 100 thousand SRTs in total were collected from 12 participants. The task had three stimulus presentation conditions differing in the degree of stimulus location uncertainty: blocked (no uncertainty), randomized (full uncertainty), and mixed (25% uncertainty). The results showed robust effects of location uncertainty, proving spatial attention's contribution to the CUD. Further, we observed a strong visual-field asymmetry reflecting the right hemisphere specialization in target detection and spatial reorienting. Lastly, despite exceptional reliability of the component SRT measures, the CUD reliability was still too low to justify using this measure as an index of individual differences.

Posterior theta activity reveals an early signal of self-face recognition.

Self-related visual information, especially one's own face and name, are processed in a specific, prioritized way. However, the spatio-temporal brain dynamics of self-prioritization have remained elusive. Moreover, it has been unclear whether this prioritization is an effect of enhancement and amplification, or rather a facilitating automatization of processing self-referential information. In this EEG study, 25 married women (who changed their surnames after marriage, so that their past and present surnames could be used as stimuli) performed a detection task with faces and names from five categories: self, self from the past, friend, famous, and unknown person. The aim was to determine the temporal and spatial characteristics of early electrophysiological markers of self-referential processing. We report results of event-related component (ERP) and time-frequency analyses. In the ERPs, the earliest self-relevance effect was displayed only 300 ms after stimulus onset in the midfrontal N2, and later in the parietal P3b, independently of the stimulus type. No self-relevance effect was found on the N170 component. However, local theta power at the occipito-temporal (visual) areas and inter-regional theta phase coherence between the visual and midfrontal areas showed that self-relevance differentiation of faces began already about 100-300 ms after stimulus onset. No such early effects were found for names. The results are discussed in terms of the time-course, functional localization, stimulus-specificity, and automatization of self-prioritization.

Air pollution and attention in Polish schoolchildren with and without ADHD.

BACKGROUND: Development and functioning of attention-a key component of human cognition-can be affected by environmental factors. We investigated whether long- and short-term exposure to particulate matter with aerodynamic diameter < 10 µm (PM10) and nitrogen dioxide (NO2) are related to attention in 10- to 13-year-old children living in Polish towns recruited in the NeuroSmog case-control study. METHODS: We investigated associations between air pollution and attention separately in children with attention deficit hyperactivity disorder (ADHD, n = 187), a sensitive, at-risk population with impaired attention and in population-based typically developing children (TD, n = 465). Alerting, orienting, and executive aspects of attention were measured using the attention network test (ANT), while inhibitory control was measured with the continuous performance test (CPT). We assessed long-term exposure to NO2 and PM10 using novel hybrid land use regression (LUR) models. Short-term exposures to NO2 and PM10 were assigned to each subject using measurements taken at the air pollution monitoring station nearest to their home address. We tested associations for each exposure-outcome pair using adjusted linear and negative binomial regressions. RESULTS: We found that long-term exposures to both NO2 and PM10 were associated with worse visual attention in children with ADHD. Short-term exposure to NO2 was associated with less efficient executive attention in TD children and more errors in children with ADHD. It was also associated with shorter CPT response times in TD children; however, this effect was accompanied by a trend towards more CPT commission errors, suggestive of more impulsive performance in these subjects. Finally, we found that short-term PM10 exposure was associated with fewer omission errors in CPT in TD children. CONCLUSIONS: Exposure to air pollution, especially short-term exposure to NO2, may have a negative impact on attention in children. In sensitive populations, this impact might be different than in the general population.

Hemispheric asymmetry in the efficiency of attentional networks.

Despite the fact that hemispheric asymmetry of attention has been widely studied, a clear picture of this complex phenomenon is still lacking. The aim of the present study was to provide an efficient and reliable measurement of potential hemispheric asymmetries of three attentional networks, i.e. alerting, orienting and executive attention. Participants (N=125) were tested with the Lateralized Attention Network Test (LANT) that allowed us to investigate the efficiency of the networks in both visual fields (VF). We found a LVF advantage when a target occurred in an unattended location, which seems to reflect right hemisphere superiority in control of the reorienting of attention. Furthermore, a LVF advantage in conflict resolution was observed, which may indicate hemispheric asymmetry of the executive network. No VF effect for alerting was found. The results, consistent with the common notion of general right hemisphere dominance for attention, provide a more detailed account of hemispheric asymmetries of the attentional networks than previous studies using the LANT task.

Subjective measures of consciousness in artificial grammar learning task.

Consciousness can be measured in various ways, but different measures often yield different conclusions about the extent to which awareness relates to performance. Here, we compare five different subjective measures of awareness in the context of an artificial grammar learning task. Participants (N=217) expressed their subjective awareness of rules using one of five different scales: confidence ratings (CRs), post-decision wagering (PDW), feeling of warmth (FOW), rule awareness (RAS), and continuous scale (SDS). All scales were equally sensitive to conscious knowledge. PDW, however, was affected by risk aversion, and both RAS and SDS applied different minimal criteria for rule awareness. CR seems to capture the largest range of consciousness, but failed to indicate unconscious knowledge with the guessing criterion. We close by discussing the theoretical implications of scale sensitivity and propose that CR's unique features enable (in conjunction with RAS and FOW) a finer assessment of subjective states of awareness.

Phasic alerting facilitates endogenous orienting of spatial attention: Evidence from event-related lateralizations of the EEG.

Alerting has been hypothesized to affect spatial orienting either by accelerating the speed of attentional shift toward the cued target location (the accelerating hypothesis) or by enhancing the orienting effect without changing its time course (the enhancing hypothesis). To investigate the neural underpinnings of the effect of phasic alerting on endogenous orienting, we recorded the electroencephalogram (EEG) in a variant of the spatial cueing task with a tone presented 100 ms before the cue as a phasic alerting signal, and calculated cue-evoked event-related lateralizations (ERLs) providing a precise assessment of preparatory visuospatial attention. Behavioral results showed that the spatial orienting effect was increased under the phasic alerting condition, as expected. The EEG results showed that an orienting-related ERL component called a late directing attention positivity (LDAP) had shorter onset latency and larger amplitude in the alerting condition than in the no-alerting (no-tone) condition. In conclusion, phasic alerting seems to both accelerate and enhance orienting-related preparatory modulations within the ventral visual stream.

The response relevance of visual stimuli modulates the P3 component and the underlying sensorimotor network.

The functional meaning and neural basis of the P3b component of ERPs are still under debate. One of the main issues is whether P3b reflects only stimulus-related processes (stimulus evaluation hypothesis) or response-related processes as well (stimulus-response or S-R link activation hypothesis). Here, we conducted an EEG experiment examining whether P3b may indeed reflect an S-R link activation, followed by an fMRI experiment in which we explored the brain areas and functional connectivity possibly constituting the neural basis of these sensorimotor links. In both experiments, two successive visual stimuli, S1 and S2, were presented with a 1 sec interval, and responses were defined either by S1 or S2, while participants responded only after S2 onset. The obtained EEG results suggest that P3b may be interpreted in terms of the S-R link activation account, although further studies are needed to disentangle P3-related activity from overlapping anticipatory activity. The obtained fMRI results showed that processing of the relevant S1 involved activation of a distributed postero-anterior sensorimotor network, and increased strength of functional connectivity within this network. This network may underlie activation of the S-R links, thus possibly also the P3b component, forming a bridging step between sensory encoding and response execution.

On the locus of the effect of alerting on response conflict: An event-related EEG study with a speed-accuracy tradeoff manipulation.

The present study investigated the locus of the effect of alerting on response conflict, and examined whether this effect may be interpreted as an alerting-triggered imbalance of speed-accuracy tradeoff (SAT). Participants performed the flanker task with an alerting tone in half of the trials and SAT manipulation. Behavioral results showed the usual increase of the conflict effect (incongruent-congruent flankers) in the alerting trials. This interaction was not affected by SAT manipulation in response times, although accuracy emphasis abolished alerting effects in error rates. Event-related potential (ERP) results showed that alerting increased the conflict by facilitating the selection and activation of stimulus-response links, reflected in modulations of the P3b component, and by enhancing the activation of incorrect response evoked by incongruent flankers, reflected in an increased initial incorrect activation in the lateralized readiness potential (LRP). Time-frequency analysis showed that the alerting-triggered increase of conflict entailed stronger response of executive mechanisms, reflected in a larger conflict-related midfrontal theta-band power. These EEG effects were not affected by SAT manipulation. In conclusion, alerting affects both the emergence of conflict and conflict control, and this alerting-conflict interaction could not be explained in terms of SAT.

Differential effects of phasic and tonic alerting on the efficiency of executive attention.

The study examined how alerting and executive attention interact in a task involving conflict resolution. We proposed a tentative scenario in which an initial exogenous phasic alerting phase is followed by an endogenous tonic alerting phase, and hypothesized that these two processes may have distinct effects on conflict resolution. Phasic alerting was expected to increase the conflict, whereas tonic alerting was expected to decrease the conflict. Three experiments were conducted using different variants of the flanker task with visual alerting cues and varied cue-target intervals (SOA), to differentiate between effects of phasic alerting (short SOA) and tonic alerting (long SOA). The results showed that phasic alerting consistently decreased the efficiency of conflict resolution indexed by response time and accuracy, whereas tonic alerting increased the accuracy of conflict resolution, but at a cost in the speed of processing the conflict. The third experiment additionally showed that the effects of phasic alerting may be modulated by the psychophysical strength of alerting cues. Discussed are possible mechanisms that could account for the observed interactions between alerting and conflict resolution, as well as some discrepancies between the current and previous studies.

Biased odds for heads or tails: Outcome-evoked P3 depends on frequencies of guesses.

Events that had to be predicted evoke large P3 components of the event-related EEG potential. There is conflicting evidence whether these P3s are moderated by participants' preceding guesses. In the present study, participants made one prediction frequently and the other rarely because one stimulus was presented frequently and the other rarely. Thereby, effects on stimulus-evoked P3s of both guess frequency and stimulus frequency could be tested. Indeed, P3s were not only larger with rare than frequent stimuli but also larger after rare than frequent guesses. This result pattern may have additionally been affected by expectancies for payoff. In any case, the modification of outcome-evoked P3 by what had been guessed may reflect that each of the four guess-stimulus combinations is encoded as a separate event category. In terms of the stimulus-response link hypothesis of P3b, it is suggested that P3s are evoked by these events because internal responses (right or wrong) are associated to each of these event categories and need to be reactivated with rare guess-stimulus combinations.

Limits of control: the effects of uncontrollability experiences on the efficiency of attentional control.

Two experiments were conducted to explore the effects of experiencing uncontrollability on the efficiency of attentional control. The experience of uncontrollability was induced either by unsolvable tasks (Experiment 1) or by tasks in which non-contingent feedback was provided (Experiment 2). A version of the Attentional Network Test-Interactions with an additional measure of vigilance (ANTI-V) was used to evaluate the efficiency of the attentional networks (i.e., alerting, orienting, and executive). Results of both experiments revealed a decreased efficiency of executive attention in participants who experienced stable control deprivation but no negative effects in participants who were able to restore their sense of previously deprived control. Additionally, when participants were asked to perform unsolvable tasks and did not receive feedback (Experiment 1), detrimental effects on the orienting network and vigilance were observed. The motivational and cognitive mechanisms underlying the effects of various uncontrollability experiences on conflict resolution and attentional control are discussed.

Testing the S-R link hypothesis of P3b: The oddball effect on S1-evoked P3 gets reduced by increased task relevance of S2.

We had previously reported that the oddball effect on the P3b EEG potential evoked by infrequent vs. frequent S1 presented in a sequence of two stimuli, S1 and S2, gets reduced in a "combination task". In this task, responses were determined by the combinations of S1 and S2 rather than by S1 only. We had attributed this reduction of the oddball effect to increased task difficulty. The present study investigated possible reasons for this reduction of S1-evoked P3b in more detail, by making the combination task easier in several respects: allowing for forming associations from S1 to responses (Experiment 1), reducing the complexity of stimulus-response (S-R) mappings (Experiment 2), and decreasing S2 relevance in defining responses (Experiment 3). The results showed that only S2 relevance affected the oddball effect on S1-evoked P3b. Namely, when S2 attained some relevance by inducing a go/no-go decision for S1-defined responses, the oddball effect on S1-evoked P3b was intermediate between the large effect in the simple oddball task and the small effect in the combination task. The results may be explained in terms of the S-R link hypothesis of P3b which interprets P3b as reflecting reactivation of well-established S-R links.