Brain mechanisms of involuntary visuospatial attention: an event-related potential study

The brain mechanisms mediating visuospatial attention were investigated by recording event-related potentials (ERPs) during a line-orientation discrimination task. Nonpredictive peripheral cues were used to direct participant's attention involuntarily to a spatial location. The earliest attentional modulation was observed in the P1 component (peak latency about 130 ms), with the valid trials eliciting larger P1 than invalid trials. Moreover, the attentional modulations on both the amplitude and latency of the P1 and N1 components had a different pattern as compared to previous studies with voluntary attention tasks. In contrast, the earliest visual ERP component, C1 (peak latency about 80 ms), was not modulated by attention. Low-resolution brain electromagnetic tomography (LORETA) showed that the earliest attentional modulation occurred in extrastriate cortex (middle occipital gyrus, BA 19) but not in the primary visual cortex. Later attention-related reactivations in the primary visual cortex were found at about 110 ms after stimulus onset. The results suggest that involuntary as well as voluntary attention modulates visual processing at the level of extrastriate cortex; however, at least some different processes are involved by involuntary attention compared to voluntary attention. In addition, the possible feedback from higher visual cortex to the primary visual cortex is faster and occurs earlier in involuntary relative to voluntary attention task.     

Neurophysiological correlates of cognitive development: changes in long-latency event-related potentials from childhood to adulthood.

Event-related brain potentials (ERPs) to tachistoscopically flashed slides were recorded from subjects ranging from 6 to 36 years of age. Subjects counted the number of presentations of target slides (P = 0.12) randomly interposed in sequences of background slides (either P = 0.76 or P = 0.88). Also interposed (P = 0.12) in these sequences of targets and backgrounds were one of two types of slides: those (termed dims) bearing any one of the letters from C to Z or those bearing 'novel' patterns, each consisting of a different, quasi-random, 'unrecognizable' color pattern. Targets and backgrounds elicited N1, P2, N2 and P3 waves in subjects of every age tested, suggesting that the mode of processing such explicitly important events and the neuronal substrate for such processing, is similar in children and adults. However, P3 latencies and reaction times in response to these events decreased with age, suggesting that the speed of such processing increases with age. On the other hand, dims and novels elicited ERPs in young children and adults characterized by different late waves. ERPs in 6--8 year olds were characterized by Nc waves (ca. 410 msec and 30 muV) and Pc waves (ca. 900 msec and 30 muV), while ERPs in adults were characterized by P3 waves (ca. 420 msec and 15 muV. The transition from the childhood to adulthood wave form occured in the mid-teens. It is suggested that these differences in ERP wave forms reflect differences in the way children and adults categorize events.     

The influence of caffeine on spatial-selective attention: an event-related potential study.

OBJECTIVES: Following the indications of previous studies that caffeine might have a specific effect on the processing of spatial information compared with other types of information, the present study investigated the influence of caffeine on an often used spatial-selective attention task. METHODS: Event-related potentials (ERPs) were recorded from 11 participants under conditions of caffeine (250 mg) and placebo. RESULTS: Spatial-selective attention effects were reflected in the ERPs as more positive going occipital P1 and broadly distributed P2 components, and more negative going occipital-temporal N1 and broadly distributed N2 components. A treatment effect was found as a more positive going frontal P2 component in the caffeine condition, whereas interactions between treatment and attention were observed for P2 and N2 components, but not for P1 and N1 components. CONCLUSIONS: This pattern of results suggests that caffeine has no specific influence on spatial-selective attention, but rather, has a more general facilitating effect on perceptual processing, as well as a possible effect on the frontal control mechanisms, i.e. focusing attention and increasing selectivity.     

The orienting of visuospatial attention: an event-related brain potential study

This study investigated the electrophysiological correlates of shifting, maintaining, and relaxing the focus of attention, using a symbolic cuing task. Cues and imperative stimuli were presented in rapid succession, and the ADJAR procedure was used to remove the contribution of event-related potential (ERP) activity associated with the imperative stimulus from the cue-related ERP waveforms. Initial analyses, comparing left and right attention-directing cues, replicated previous findings of early directing attention negativity (EDAN) and anterior directing attention negativity (ADAN) effects. To isolate ERP activity that is common to leftward and rightward attention shifts, the combined ERP activity elicited by attention-directing cues was compared to the ERP activity elicited by non-informative cues. This analysis revealed a strong and broadly distributed early positivity followed by a sustained central negativity, possibly reflecting the controlled orienting and subsequent maintenance of attentional focus. Finally, imperative stimuli preceded by non-informative cues were characterized by an enhanced posterior P2 effect, with a scalp distribution indicative of generators in visual areas. This result suggests a relatively late (re)activation in visual areas associated with the processing of stimuli that had not been cued in advance.     

Visual distraction: a behavioral and event-related brain potential study in humans

Recent studies reported that the detection of changes in the visual stimulation results in distraction of cognitive processing. From event-related brain potentials it was argued that distraction is triggered by the automatic detection of deviants. We tested whether distraction effects are confined to the detection of a deviation or can be triggered by changes per se, namely by rare stimuli that were not deviant with respect to the stimulation. The results obtained comparable early event-related brain potential effects for rare and deviant stimuli, suggesting an automatic detection of these changes. In contrast, behavioral distraction and attention-related event-related brain potential components were confined to deviant stimuli. This finding suggests that deviancy from a given standard adds a genuine contribution to distraction.     

Auditory distraction: event-related potential and behavioral indices.

OBJECTIVE: The aim of this study was to illuminate behavioral and event-related potential (ERP) effects of attentional orienting and reorienting obtained in a newly developed auditory distraction paradigm, to provide more precise indicators about the neural generators of the ERP effects using scalp current density (SCD) analysis, and to evaluate the stability of the distraction effects. METHODS: In two sessions separated by 25 days, 10 subjects were presented with tones being of short (200 ms) and long (400 ms) duration equiprobably; tones were of high-probability standard or of low-probability deviant frequency. In Distraction condition, subjects had to behaviorally discriminate short from long tones. In Ignore condition, subjects were reading a book. Behavioral performance and multi-channel EEG were recorded. RESULTS: Task-irrelevant frequency deviations prolonged reaction times in the duration discrimination task by more than 35 ms and elicited the MMN and P3a components of the event-related potential. The P3a was followed by a negative deflection called RON (reorienting negativity). P3a and RON were absent in Ignore condition. All effects were found to be highly stable between sessions (product-moment correlations between 0.76 and 0.90). SCD analysis suggested frontal generators for P3a and for RON. CONCLUSIONS: It is demonstrated that small frequency deviations may yield distinct distraction effects in a tone duration discrimination task on a behavioral and on an electrophysiological level. Results support the hypothesis that frontal areas are involved in the exogenous orienting of attention (P3a) and in the reorienting of attention (RON). Due to the high stability of the deviance-related behavioral and ERP effects, this distraction paradigm may be utilized for clinical research.     

The development of auditory evoked dipole source activity from childhood to adulthood.

OBJECTIVE: Multi-channel recordings show that observed developmental changes of late auditory evoked potentials (LAEP) depend on the location of the scalp electrode. These findings suggest that different LAEP generators have a distinct developmental course. The goal of this study was to investigate the maturational process of cortical LAEP generators. METHODS: Eighty-seven healthy children and adolescents with normal hearing, ages 5-16 years, and 21 adults, ages 20-30 years, participated in the study. Pure tone LAEP were recorded from 21 derivations. Dipole source analysis was performed by means of brain electric source analysis (BESA). Peak latencies and amplitudes of dipole source activity were estimated. RESULTS: While the number, location, and direction of dipole sources were similar in children and adults, the course of their activity differed greatly. The latencies shortened and the amplitudes decreased during development. In adolescence a new component appeared in the activity of the tangential dipole, which reflects the generators in the supra-temporal plane. The variability of parameters was greater in children than in adults. CONCLUSIONS: Since the dipole source activity of LAEP in childhood differs considerably from that in adulthood, dipole source analysis could be a useful tool for studying both normal and disturbed maturation of the auditory perceptual function.     

Auditory distraction by duration and location deviants: a behavioral and event-related potential study

Auditory distractibility was investigated using four noise stimuli that differed in their duration and/or sound source. In the duration-task/location-deviant condition, participants were asked to discriminate between equiprobable short and long stimuli. Mostly, stimuli were presented from one location (Standards), but, infrequently, a stimulus was presented from another location (Deviant). In the location-task/duration-deviant condition, participants had to discriminate between stimuli presented equiprobably from the speaker in front of them or to their left. Here, most stimuli were of equal duration (Standards), but, infrequently, a stimulus duration changed (Deviant). The rare deviations in location and duration were irrelevant for the actual task. Whether they affected processes related to the actual task was assessed with performance- and event-related potential (ERP) measures. In both conditions, responses to Deviants were slowed compared to responses to Standards. Deviants elicited ERP components mismatch negativity (MMN), P3a and reorienting negativity (RON). These results show that the processing of both a sound's duration and a sound's location can be distracted by rare, but irrelevant, changes in its location and duration, respectively. Behavioral distraction effects were markedly smaller with duration Deviants. It is suggested that duration Deviants interfere with task-related processing at later stages than location Deviants, as the processing of task-relevant information (i.e. stimulus location) commences before deviation in the location-task/duration-deviant condition occurs. Interestingly, distraction effects also prevail in the first Standard stimulus after a Deviant, as indicated by the prolonged response times and late negativity in the ERPs.     

Neural mechanisms of voluntary and involuntary recall: a PET study

Neuropsychological and neuroimaging studies on episodic memory retrieval have primarily focused on volitional memory tasks. However, some conscious memories arise involuntarily, i.e. without a strategic retrieval attempt, yet little is known about the neural network underlying involuntary episodic memory. The aim of this study was to determine whether voluntary and involuntary recall are mediated by separate cortical networks. We used positron emission tomography (PET) to measure changes in regional cerebral blood flow (rCBF) in 12 healthy subjects during voluntary and involuntary cued recall of pictures and a control condition with no episodic memory requirements. Involuntary recall was elicited by using an incidental memory task. Compared to the control condition, voluntary and involuntary recall were both associated with significant regional cerebral blood flow (rCBF) increases in posterior cingulate gyrus (PCG; BA 23), left precuneus (BA 7), and right parahippocampal gyrus (BA 35/36). In addition, rCBF in right dorsolateral prefrontal cortex (PFC; BA 8/9) and left precuneus (BA 7) was significantly larger during voluntary compared to involuntary recall, while rCBF was enhanced in left dorsolateral PFC (BA 9) during involuntary recall. The findings corroborate an association of the right PFC with a strategic component of episodic memory retrieval. Moreover, they show for the first time that it is possible to activate the medial temporal lobe, the PCG, and the precuneus, regions normally associated with retrieval success, without this strategic element. The relatively higher activity in precuneus during voluntary compared to involuntary recall suggests that activity in this region co-varies not only with retrieval success but also with retrieval intentionality.     

Auditory distraction with different presentation rates: an event-related potential and behavioral study.

OBJECTIVE: The present study addresses the question of whether behavioral and electrophysiological effects obtained with the auditory distraction paradigm proposed by Schr?ger et al. [Clin Neurophysiol 2000;111:1450] depend on the timing of stimulus occurrence. METHODS: Subjects had to discriminate the duration of tones. Occasionally, task-irrelevant frequency changes were used as distractors. In 3 experiments the stimulus onset asynchrony (SOA) was manipulated: In Experiment 1, SOAs of 1500, 2000 and 3000 ms were used in separate blocks; in Experiment 2, 5 different SOA of lengths between 1300 and 2500 ms were used within the blocks; in Experiment 3, a constant SOA of 1400 ms was compared with a SOA of random lengths between 1300 and 1500 ms. Performance data was analyzed for distraction effects. Event-related potentials (ERPs) were examined for deviance-related components, i.e. the mismatch negativity (MMN), the P3a and the re-orienting negativity (RON). RESULTS: Behavioral distraction effects were obtained in all experimental conditions. The electrophysiological data show MMN, P3a and RON for the deviating tones in all experimental conditions as well. CONCLUSIONS: The behavioral and electrophysiological deviance-related effects were not sensitive to the SOA manipulations. Therefore, the timing of the presentation can be adjusted to the proficiency level of the population to be tested without loosing the distraction effects.     

Intra-modal and cross-modal spatial attention to auditory and visual stimuli. An event-related brain potential study.

This study investigated cross-modal interactions in spatial attention by means of recording event-related brain potentials (ERPs). Noise bursts and light flashes were presented in random order to both left and right field locations separated by 60 degrees in free-field. One group of subjects was instructed to attend selectively to the noise bursts (attend-auditory group), and a second group attended only to the flashes (attend-visual group). On different runs attention was directed to either the right or left field stimuli of the designated modality. In the attend-auditory group, noise bursts at the attended location elicited a broad, biphasic negativity (Nd) beginning at 70 ms. The cross-modal spatial attention effect on the auditory ERPs in the attend-visual group was very similar in morphology, but the Nd was reduced in amplitude relative to the intra-modal effect. In the attend-visual group, flashes at the attended location elicited enhanced early (100-200 ms) and late (200-350 ms) ERP components relative to unattended-location flashes. The cross-modal effect in the attend-auditory group included small but significant enhancements of early components of the visual ERPs. It was concluded that spatial attention has a multi-modal organization such that the processing of stimuli at attended locations is facilitated at an early, sensory level, even for stimuli of an unattended modality.     

Auditory selective attention in middle-aged and elderly subjects: an event-related brain potential study.

Reaction times (RTs) and event-related brain potentials (ERPs) were recorded in middle-aged (MA) and elderly (ELD) subjects performing an auditory selective attention task. Subjects attended to tone bursts of a specified pitch and ear of delivery and responded to occasional longer duration target tones (75 vs. 25 msec). Infrequent novel stimuli (computer synthesized sounds and digitized environmental noises) were also included in the stimulus sequence. No significant age-related differences were found in the speed or accuracy of target detection. However, in both groups, RTs were delayed (by more than 300 msec) to targets that followed novel sounds. The prolongation was greater following novel sounds in the attended ear, particularly in the ELD group. The effects of selective attention on ERPs to standard tones were isolated as negative difference waves (Nds) by substracting ERPs to non-attended stimuli from ERPs to the same signals when attended. Nds had similar amplitudes, latencies of onset (60 msec), and distributions in ELD and MA groups. In both groups, Nd waves were more prominent following right ear stimulation, reflecting possible hemispheric asymmetries of generators in posterior temporal regions. The mismatch negativity (MMN) was isolated by subtracting ERPs to standard tones from ERPs to deviant stimuli. MMN amplitudes were reduced in the ELD group. There was also a significant change in MMN distribution with age: the MMN was larger over the right hemisphere for MA subjects but larger over the left for ELD subjects. Elderly subjects showed a trend toward smaller P3 amplitudes and delayed P3 latencies, but group differences did not reach statistical significance.(ABSTRACT TRUNCATED AT 250 WORDS)     

The development of preparation, conflict monitoring and inhibition from early childhood to young adulthood: a Go/Nogo ERP study

The present developmental study aimed to trace changes in response expectation, preparation, conflict monitoring and subsequent response inhibition from 6 years of age to adulthood. In two groups of children (6-7 and 9-10 years old) and young adults (19-23 years old), behavior and event-related brain activity (ERP) in a CPT-AX task was measured. Hits, false alarms, inattention and impulsivity scores and ERP measures of conflict monitoring and inhibition (Nogo-N2 and P3), cue-orientation and prestimulus target expectation (cue-P2 and P3) and response preparation (Contingent Negative Variation; CNV) were collected. Behavioral measures indicated that attention processes developed most strongly before age 10, whereas impulsive behavior only started to diminish after the age of 10. Nogo-N2 effects were largest and more widely distributed across fronto-parietal electrodes in 6-7-year olds and decreased linearly with age. Nogo-P3 effects showed an opposite pattern by being absent in the youngest children, starting to develop at age 9-10 and reaching maturity in young adulthood. These developmental behavioral and ERP results are supportive of links between Nogo-N2 and conflict monitoring and Nogo-P3 and response inhibition and suggest that both are liable to different developmental progress. Furthermore, enhanced cue-P3 activity in both 6-7 and 9-10-year olds was argued to reflect a higher level of Go-stimulus expectation, that might cause them to experience more conflict on subsequent Nogo-trials, when the 'not-expected' stimulus appears. On the other hand, young children's reduced preparatory CNV activity was interpreted as a sign of reduced response priming caused by yet immature fronto-parietal networks involved in motor regulation.     

Auditory evoked potential development in early childhood: a longitudinal study.

Serial recordings of auditory evoked potentials (AEPs) to clicks were obtained using a vertex-mastoid derivation from 16 normal children during sleep over an age span from near birth to age 3. The AEP components studied were: N0 (38 +/- 10 msec), P1 (79 +/- 24 msec), N1 (109 +/- 39 msec), P2 (186 +/- 35 msec), N2 (409 +/- 97 msec), P3A (554 +/- 116 msec), P3B (757 +/- 121 msec) and P3 (728 +/- 128 msec). Amplitudes and latencies of the components were calculated and regressions of the measures on age were computed for the group as a whole, for each subject and for subsets of the data based on sleep stage, sex, order of stimulus presentation and a rearing/race factor. For the group as a whole the latencies of P1, P2, P3, and P3B decreased with age. The amplitudes of P1N1 and the N2P3 waves increased with age. Most change occurred during the first year of life. In general, the changes with age were also found to hold across all of the factors examined, although individuals varied widely in the degree to which they conformed to the trends found for the data as a whole. The amount contributed by each of the factors mentioned above to the total variance was estimated. The proportions varied for different EP components but, in general, age, sleep state, and subject factors other than rearing/race and sex accounted for most variance. One half to 5/6 of the unexplained variance in AEP latencies and amplitudes (i.e., that not due to age, sleep state, etc.) occurred across rather than within subjects. For both the group as a whole and for individual children, P2 and N2 latencies were found to exhibit the greatest stability across time. The results of the longitudinal study reported here were in good agreement with those of a previous study from this laboratory which utilized a cross-sectional design.     

Selective auditory attention in 3- to 5-year-old children: an event-related potential study

Behavioral and electrophysiological evidence suggests that the development of selective attention extends over the first two decades of life. However, much of this research may underestimate the attention abilities of young children. By providing strong, redundant attention cues, we show that sustained endogenous selective attention has similar effects on ERP indices of auditory processing in adults and children as young as 3 years old. All participants were cued to selectively attend to one of two simultaneously presented stories that differed in location (left/right), voice (male/female), and content. The morphology of the ERP waveforms elicited by probes embedded in the stories was very different for adults, who showed a typical positive-negative-positive pattern in the 300 ms after probe onset, and children, who showed a single broad positivity during this epoch. However, for 3- to 5-year-olds, 6- to 8-year-olds, and adults, probes in the attended story elicited larger amplitude ERPs beginning around 100 ms after probe onset. This attentional modulation of exogenously driven components was longer in duration for the youngest children. In addition, attended linguistic probes elicited a larger negativity 300-500 ms for all groups, indicative of additional attentional processing. These data show that with adequate cues, even children as young as 3 years old can selectively attend to one auditory stream while ignoring another and that doing so alters auditory sensory processing at an early stage. Furthermore, they suggest that the neural mechanisms by which selective attention affects auditory processing are remarkably adult-like by this age.     

The neuronal correlate of bidirectional synesthesia: a combined event-related potential and functional magnetic resonance imaging study

The neuronal correlate of a rare explicit bidirectional synesthesia was investigated with numerical and physical size comparison tasks using both functional magnetic resonance imaging and event-related potentials. Interestingly, although participant I.S. exhibited similar congruity effects for both tasks at the behavioral level, subsequent analyses of the imaging data revealed that different brain areas were recruited for each task, and in different time windows. The results support: (1) the genuineness of bidirectional synesthesia at the neuronal level, (2) the possibility that discrepancy in the neuronal correlates of synesthesia between previous studies might be task-related, and (3) the possibility that synesthesia might not be a unitary phenomenon.     

Progressive increase of frontostriatal brain activation from childhood to adulthood during event-related tasks of cognitive control

Higher cognitive inhibitory and attention functions have been shown to develop throughout adolescence, presumably concurrent with anatomical brain maturational changes. The relatively scarce developmental functional imaging literature on cognitive control, however, has been inconsistent with respect to the neurofunctional substrates of this cognitive development, finding either increased or decreased executive prefrontal function in the progression from childhood to adulthood. Such inconsistencies may be due to small subject numbers or confounds from age-related performance differences in block design functional MRI (fMRI). In this study, rapid, randomized, mixed-trial event-related fMRI was used to investigate developmental differences of the neural networks mediating a range of motor and cognitive inhibition functions in a sizeable number of adolescents and adults. Functional brain activation was compared between adolescents and adults during three different executive tasks measuring selective motor response inhibition (Go/no-go task), cognitive interference inhibition (Simon task), and attentional set shifting (Switch task). Adults compared with children showed increased brain activation in task-specific frontostriatal networks, including right orbital and mesial prefrontal cortex and caudate during the Go/no-go task, right mesial and inferior prefrontal cortex, parietal lobe, and putamen during the Switch task and left dorsolateral and inferior frontotemporoparietal regions and putamen during the Simon task. Whole-brain regression analyses with age across all subjects showed progressive age-related changes in similar and extended clusters of task-specific frontostriatal, frontotemporal, and frontoparietal networks. The findings suggest progressive maturation of task-specific frontostriatal and frontocortical networks for cognitive control functions in the transition from childhood to mid-adulthood.     

An event-related potential study of selective auditory attention in children and adults

In a dichotic listening paradigm, event-related potentials (ERPs) were recorded to linguistic and nonlinguistic probe stimuli embedded in 2 different narrative contexts as they were either attended or unattended. In adults, the typical N1 attention effect was observed for both types of probes: Probes superimposed on the attended narrative elicited an enhanced negativity compared to the same probes when unattended. Overall, this sustained attention effect was greater over medial and left lateral sites, but was more posteriorly distributed and of longer duration for linguistic as compared to nonlinguistic probes. In contrast, in 6- to 8-year-old children the ERPs were morphologically dissimilar to those elicited in adults and children displayed a greater positivity to both types of probe stimuli when embedded in the attended as compared to the unattended narrative. Although both adults and children showed attention effects beginning at about 100 msec, only adults displayed left-lateralized attention effects and a distinct, posterior distribution for linguistic probes. These results suggest that the attentional networks indexed by this task continue to develop beyond the age of 8 years.     

The development of functional connectivity within the dorsal striatum from early childhood to adulthood

Dorsal striatum, principally comprising of caudate and putamen, is well-known to support motor function but also various higher-order cognitive functions. This is enabled by developing short- and long-range connections to distributed cortical regions throughout the life span, but few studies have examined developmental changes from young children to adults in the same cohort. Here we investigated the development of dorsal-striatal network in a large (n = 476), single-site sample of healthy subjects 3–42 years of age in three groups (children, adolescence, adults). The results showed that the connectivity within the striatum and to sensorimotor regions was established at an early stage of life and remained strong in adolescence, supporting that sensory-seeking behaviours and habit formation are important learning mechanisms during the developmental periods. This connectivity diminished with age, as many behaviours become more efficient and automated. Adolescence demonstrated a remarkable transition phase where the connectivity to dorsolateral prefrontal cortex emerged but connectivity to the dorsomedial prefrontal and posterior brain, which belong to the ventral attentional and default mode networks, was only seen in adults. This prolonged maturation in between-network integration may explain the behavioural characteristics of adolescents in that they exhibit elaborated cognitive performance but also demonstrate high risk-taking behaviours.     

Distraction effects in vision: behavioral and event-related potential indices

Evidence is presented which shows that slight changes in serially presented visual input can be detected automatically and may result in behavioral distraction. In two experiments, reaction times on a two-alternative, forced-choice duration discrimination task were prolonged in trials where a task-irrelevant change in the location of the stimulus occurred. The P1 and N1 components of the event-related potential were enhanced in such trials. However, electrophysiological and behavioral effects were not affected when the duration discrimination was made more difficult and the N1 amplitude increase and RT prolongation were confined to situations where task-irrelevant location changes were infrequent. Thus, these effects most probably reflect pre-attentive change detection and subsequent distraction in vision, whereas the P1 effect is probably due to spatial eccentricity.