Brain Potentials During Selective Attention, Memory Search, and Mental Rotation

Event-related potentials were measured in a task that combined the classic selective attention paradigm with memory search and mental rotation paradigms. Subjects were required to attend to stimulus letters in one color and to ignore stimuli in a different color. Within the attended category subjects searched for target letters from a prememorized set (the memory set) and indicated whether they were presented normally or in mirror-image. Letters in all stimulus categories were presented randomly in either their upright position or rotated over 60 degrees, 120 degrees, or 180 degrees. The event-related potentials showed that the earliest effect of attending to color was a positivity at the anterior electrodes and a negativity at Oz (onset about 150 ms), followed by the enhancement of a central N2 component (N2b, onset about 220 ms). The early effect is thought to reflect selective processing of elementary stimulus features, the later N2b the covert orienting of attention. A later, prolonged central negativity elicited by attended stimuli covaried with the duration of the memory search process. Target detection resulted in a parietal P3b component, and also in an earlier negativity (in the range approximately 200-300 ms) to both attended and unattended targets, suggesting an early preattentive target classification. There were two effects of the rotation of stimuli. First there was an early occipital effect (about 200-300 ms), irrespective of attention and stimulus categories. It was argued that this effect reflected the preattentive identification of the orientation of the stimulus letters. A later (onset about 350-400 ms) parietal effect consisted of an increase in negativity as a function of the angle over which letters were rotated. This prolonged negativity had a later onset latency than the search-related negativity, and was restricted to the event-related potentials to target letters in the attended input channel. It was suggested that this component is the manifestation of a mental rotation process. It was argued that rotation-related negativity and memory search-related activity reflect operations in distinct working memory subsystems.     

Attention to Color: An Analysis of Selection, Controlled Search, and Motor Activation, Using Event-Related Potentials

In this study the organization of information processing in a selective search task was examined by analyzing event-related potentials. This task consisted of searching for target letters in a relevant (attended) color. The ERPs revealed two different effects of attention: an early occipital negativity (+/- 150 ms) reflecting feature-specific attention, and a later, central N2b component (+/- 240 ms) reflecting covert orienting of attention. A later, prolonged negativity (search-related negativity) (+/- 300 ms), maximal at Cz, was related to controlled search to letters in the attended color. Detection of relevant targets resulted in a parietal P3b component. Depending on stimulus presentation conditions an earlier response to both attended and unattended targets was found: an N2 component (+/- 250 ms). In these same conditions, C'3-C'4 asymmetries (Corrected Motor Asymmetries--CMA) suggested motor activation at +/- 300 ms, in the same time range as search-related negativity. It was argued that N2 and CMA suggest the existence of a preattentive target detection system, operating in parallel with a slower serial attentive system, as reflected by N2b and search negativity.     

Cognitive Event-Related Potential Components During Continuous Recognition Memory for Pictures

Event-related brain potentials (ERPs) were recorded from 28 young adult subjects during a continuous recognition memory paradigm, with pictures as stimuli. Subjects were required to determine on each trial if the picture was "new" (never before presented) or "old" (seen previously). To assess differences between primary and secondary memory, old items were presented after lags of 2, 8, and 32 intervening pictures (equiprobable) following their first presentation. The results suggested that a negativity (Cz maximum) at 300 ms was the most likely candidate for the brain event reflecting the retrieval of the item from memory. Old/new effects were modulated by two types of activity, both of which were larger in the ERPs elicited by new items. The earlier of these, possibly similar to the N400, had a duration from about 250-600 ms, began with the N300 deflection, and lasted until "P300" began to decrement. The other was positive, resembled "positive slow wave," onset as P300 began to decrement, and lasted until the end of the recording epoch. There were no consistent effects of item lag on the behavioral data or on any of the ERP components, suggesting that for pictorial stimuli, the distinction between the two types of memory store, primary (i.e., immediate memory) and secondary (newly learned information), may not be relevant. In consonance with the lack of lag effects, it was suggested that the lack of a robust subsequent memory effect on the ERP waveform could have been due to the use of pictures, which may have required less elaborative processing in order to be encoded at input.     

Event-Related Potentials During Memory Search and Selective Attention to Letter Size and Conjunctions of Letter Size and Color

The analysis of event-related potentials was used in two experiments to investigate the structure of information processing in a task in which subjects selectively attended to letter size (Experiment 1) or a conjunction of letter size and color (Experiment 2) and searched for target letters within the attended stimulus category. The event-related potentials showed that selective attention to letter size resulted in the enhancement of a central N2b component (onset about 200 ms), which was assumed to reflect feature nonspecific orienting of attention. When attention was directed to conjunctions of letter size and color an earlier effect was found (onset about 150 ms) consisting of positivity at the anterior electrodes and negativity at OZ. This earlier effect was assumed to reflect feature-specific selective processing. Although the early effect showed a hierarchical pattern of results, in which the effect of attending to size was contingent on the relevance of the color attribute, the N2b showed a more independent pattern of results, in which the relevance of either the color or the size attribute resulted in an enhancement of this component, independent of the relevance of the other attribute. An increase in the duration of the memory search process resulted in a prolonged negativity with an onset of about 200 ms which was maximal at CZ. In both experiments the initial phase of this negativity was also found in the event-related potentials to the unattended stimulus categories, suggesting that the search process was initiated nonselectively and terminated after the selection cues were identified. Detection of attended target letters resulted in a parietal P3b component. In both experiments there was an earlier effect discriminating targets and nontargets in the range 200-300 ms.     

Unnoticed regularity violation elicits change-related brain activity

Event-related brain electric activity (ERP) was investigated to unnoticed visual changes. The orientation of grid elements (vertical or horizontal) changed after the presentation of 10–15 identical stimuli. The grid patterns were task irrelevant, and were presented in the background of a shape discrimination task. During the first half of the session, participants were unaware of the stimulus change. However, in comparison to the ERPs to the fifth identical stimuli, stimulus change elicited posterior negativities in the 270–375 ms range (visual mismatch negativity, vMMN). With participants instructed on the stimulus change, negativities emerged with earlier onset and with wider distribution. When stimulus change was preceded by only two identical stimuli, there were no such ERP effects. As the results show, a longer sequence of identical unattended stimuli may establish the memory representation of stimulus regularity, and violation of regularity is indicated by posterior negative ERP components (vMMN).     

Event-related potentials before saccades and antisaccades and their relation to reaction time

In the present study, reaction time (RT) was measured in 12 healthy subjects in a saccade and antisaccade task while recording electroencephalographic activity (EEG) from 62 electrodes on the scalp. Event-related potentials averaged both on target appearance and on saccade onset were larger in amplitude (increased negativity) for the antisaccade task compared to the saccade task. The relation of RT variability to EEG amplitude was studied by averaging stimulus-aligned and movement-aligned individual trials for each subject into four RT quartile groups. The analysis showed a relation of EEG amplitude to RT for both saccades and antisaccades. More specifically, the ERP negativity at 100–120 ms after stimulus onset in the saccade task and at 160–200 ms after stimulus onset in the antisaccade task for stimulus-aligned ERPs decreased monotonically with increasing RT as would be expected if this signal would be related to the eye movement preparation processes. This was much more pronounced and wide spread for the antisaccades than for visually triggered saccades. The peak negativity before movement onset for movement-aligned ERPs also covaried with RT suggesting no relation of this activity to movement preparation processes. This study then confirmed that only a particular ERP signal variation was related to the saccadic eye movement preparatory processes while other components of the ERP have no specific relation to the movement preparation. This particular signal was more prominent for antisaccades compared to visually triggered saccades supporting previous evidence for the cortical involvement in the preparation of these voluntary eye movements. In conclusion, this study validates the use of ERPs in the study of the planning and execution of saccadic eye movements.     

An electrophysiological investigation of the spatial distribution of attention to colored stimuli in focused and divided attention conditions

In the present experiment ERPs were recorded to colored stimulus bars (red or blue) which were randomly presented at one out of eight different spatial locations on a visual display. The locations were situated on a hemi-circle around the fixation point, with four locations lying in each visual half-field. The subjects were instructed to attend to one stimulus location (the most peripheral left or right location: the relevant location) in a focused attention condition and to all four locations within the same visual half-field in a divided attention condition, and were instructed to respond to stimulus bars in one color presented at the relevant location(s). In the focused attention condition, spatial attention resulted in early positivity in the P1 latency range (ca. 100-175 ms), followed by a prolonged negativity in the N1, P2, and N2 latency range (ca. 175-350 ms). These effects generalized to locations in the same visual half-field as the relevant location. The effect of attending and responding to the target color consisted of a number of different effects. An early anterior positivity, occipital negativity was observed for the relevant location and for locations in the same visual half-field as the relevant location, but not for the locations in the opposite visual field. A later central negativity (N2b) appeared to be confined to the relevant location and one location adjacent to it. Finally, a late parietal positivity (P3b) was exclusively evoked by target stimuli at the relevant location. In the divided attention condition, the ERPs evoked by stimuli presented at each of the four locations within the relevant visual half-field showed increased early positivity (enhancing P1 amplitude) as compared to the ERPs to stimuli in the opposite (irrelevant) visual half-field. The early color selection effect was also found to all stimuli within the relevant half-field, but the N2b component was evoked both by stimuli within the relevant half-field and by stimuli at the location in the irrelevant half-field closest to the midline. The P3b was present to target stimuli at the three most lateral positions within the relevant half-field but it was absent to the relevant location closest to the midline. The data suggested that the attentional spotlight encompassed at least approximately 2 degrees in the focused attention condition, and an efficient selection by visual half-field in the divided attention situation.(ABSTRACT TRUNCATED AT 400 WORDS)     

TMS of the right angular gyrus modulates priming of pop-out in visual search: combined TMS-ERP evidence

During priming of pop-out, performance at discriminating a pop-out feature target in visual search is affected by whether the target on the previous trial was defined by the same feature as on the upcoming trial. Recent studies suggest that priming of pop-out relies on attentional processes. With the use of simultaneous, combined transcranial magnetic stimulation and event-related potential recording (TMS-ERP), we tested for any critical role of the right angular gyrus (rANG) and left and right frontal eye fields (FEFs)-key attentional sites-in modulating both performance and the ERPs evoked by such visual events. Intertrial TMS trains were applied while participants discriminated the orientation of a color pop-out element in a visual search array. rANG TMS disrupted priming of pop-out, reducing reaction time costs on switch trials and speeding responses when the color of the pop-out target switched. rANG TMS caused a negativity in the ERP elicited in response to the visual stimulus array, starting 210 ms after stimulus onset. Both behavioral and ERP effects were apparent only after rANG TMS, on switch trials, and when the target in the visual search array was presented in the left visual field, with no effects after left or right FEF TMS. These results provide evidence for an attentional reorienting mechanism, which originates in the rANG and is modulated by the implicit memory of the previous trial. The rANG plays a causal role on switch trials during priming of pop-out by interacting with visual processing, particularly in the ipsilateral hemisphere representing the contralateral hemifield.     

Channel probability and Nd: an event-related potential sign of attention strategies

Event-related potentials were recorded in a two-channel selective attention paradigm designed to assess the effects of channel probability on processing negativity. For channel probabilities of 70% or greater, the subtraction Nd for standard tones was greatly reduced or absent. This change was due to apparent processing negativity in the waveform of the irrelevant standards at high channel probabilities. Deviant long tones showed similar early Nds to standard tones, and also developed late attention-related negativity commencing at about 500 ms from stimulus onset. It was concluded that there was no evidence that processing negativity to the relevant tones, defined as a negative component associated with selective attention, is affected by channel probability. Changes in Nd amplitude as the channel probabilities became more unequal were attributed to changes in strategies used by subjects, which primarily altered ERPs associated with the irrelevant tones.     

A Topographic Study of ERPs Elicited by Visual Feature Discrimination

The functional properties and topographic distribution of event-related potential (ERP) components elicited by visual discrimination of orientation, spatial frequency, spatial location, and color were investigated. ERPs were recorded from 28 electrode sites from 16 adult subjects. Five ERP components were measured: N1 (peak latency = 160 ms), P2 (250 ms), anterior N2 (260 ms), posterior N2 (280 ms), and P3 (400 ms). N1 and P2 were more negative when a stimulus was a target, showing the selection negativity effect. Feature-specific effects on component amplitude or topography varied by component. N1 and P2 were sensitive to stimulus orientation and location. Anterior or posterior N2 was sensitive to orientation, spatial frequency, and location. P3 varied with orientation, but not with other stimulus features. Cross-task comparisons of ERPs to vertical line segments in the color, orientation, and location discrimination tasks indicated that P2 and N2, but not N1 and P3, were sensitive to changes in task-demand. These data provide topographic evidence that ERP components in the 160-400 ms time domain can be differentiated on the basis to processing of specific visual features, and reflect neurophysiologically distinct visual pathways in the human cortex.     

Visual event-related brain potentials in 4-month-old infants at risk for neurodevelopmental impairments

The recording of event-related potentials (ERPs) is an electrophysiologic technique that has been used to evaluate the functional maturation of neural pathways responsible for recognition memory systems in infants and children. The purpose of this study was to evaluate ERP correlates of visual recognition memory in 4-month-old infants at risk for later cognitive impairments. We compared ERPs using a test of shape recognition at 4 months of age (adjusted for prematurity) in 16 high-risk, neonatal intensive care unit (NICU) survivors and 16 healthy full-term infants. ERPs were recorded while infants were familiarized with one stimulus (a red cross, 15 trials), then tested with 60 trials of this familiar stimulus and a novel stimulus (a red corkscrew). Both the NICU and control groups' ERPs demonstrated evidence of differential processing of the two stimuli, but the NICU groups' ERP patterns were distinctly different from those of the control group. In the NICU group, the novel stimulus elicited parietal positivity at 1000–1700 ms poststimulus, whereas in the control group the novel stimulus elicited occipital and frontal negativity at 500–1700 ms poststimulus. The ERP pattern demonstrated by the NICU group was atypical as it has not been previously described in healthy infants. The results of the study indicate that the ERP technique can be used to demonstrate altered patterns of neural activity during tasks of visual recognition memory in high-risk infants. We speculate that the atypical ERP patterns described in this study may indicate that patterns of synaptic organization were altered by neonatal events. © 1997 John Wiley & Sons, Inc. Dev Psychobiol 30 : 11–28, 1997     

Modification of sudden onset auditory ERP by involuntary attention to visual stimuli.

To investigate the cross-modal nature of the exogenous attention system, we studied how involuntary attention in the visual modality affects ERPs elicited by sudden onset of events in the auditory modality. Relatively loud auditory white noise bursts were presented to subjects with random and long inter-trial intervals. The noise bursts were either presented alone, or paired with a visual stimulus with a visual to auditory onset asynchrony of 120 ms. In a third condition, the visual stimuli were shown alone. All three conditions, auditory alone, visual alone, and paired visual/auditory, were randomly inter-mixed and presented with equal probabilities. Subjects were instructed to fixate on a point in front of them without task instructions concerning either the auditory or visual stimuli. ERPs were recorded from 28 scalp sites throughout every experimental session. Compared to ERPs in the auditory alone condition, pairing the auditory noise bursts with the visual stimulus reduced the amplitude of the auditory N100 component at Cz by 40% and the auditory P200/P300 component at Cz by 25%. No significant topographical change was observed in the scalp distributions of the N100 and P200/P300. Our results suggest that involuntary attention to visual stimuli suppresses early sensory (N100) as well as late cognitive (P200/P300) processing of sudden auditory events. The activation of the exogenous attention system by sudden auditory onset can be modified by involuntary visual attention in a cross-model, passive prepulse inhibition paradigm.     

Brain potentials index executive functions during random number generation

The generation of random sequences is considered to tax different executive functions. To explore the involvement of these functions further, brain potentials were recorded in 16 healthy young adults while either engaging in random number generation (RNG) by pressing the number keys on a computer keyboard in a random sequence or in ordered number generation (ONG) necessitating key presses in the canonical order. Key presses were paced by an external auditory stimulus to yield either fast (1 press/800 ms) or slow (1 press/1300 ms) sequences in separate runs. Attentional demands of random and ordered tasks were assessed by the introduction of a secondary task (key-press to a target tone). The P3 amplitude to the target tone of this secondary task was reduced during RNG, reflecting the greater consumption of attentional resources during RNG. Moreover, RNG led to a left frontal negativity peaking 140 ms after the onset of the pacing stimulus, whenever the subjects produced a true random response. This negativity could be attributed to the left dorsolateral prefrontal cortex and was absent when numbers were repeated. This negativity was interpreted as an index for the inhibition of habitual responses. Finally, in response locked ERPs a negative component was apparent peaking about 50 ms after the key-press that was more prominent during RNG. Source localization suggested a medial frontal source. This effect was tentatively interpreted as a reflection of the greater monitoring demands during random sequence generation.     

Masked repetition priming and event-related brain potentials: a new approach for tracking the time-course of object perception

This study reports a new approach to studying the time-course of the perceptual processing of objects by combining for the first time the masked repetition priming technique with the recording of event-related potentials (ERPs). In a semantic categorization task ERPs were recorded to repeated and unrelated target pictures of common objects that were immediately preceded by briefly presented pattern masked prime objects. Three sequential ERP effects were found between 100 and 650 ms post-target onset. These effects included an early posterior positivity/anterior negativity (N/P190) that was suggested to reflect early feature processing in visual cortex. This early effect was followed by an anterior negativity (N300) that was suggested to reflect processing of object-specific representations and finally by a widely distributed negativity (N400) that was argued to reflect more domain general semantic processing.     

Preferred EEG brain states at stimulus onset in a fixed interstimulus interval equiprobable auditory Go/NoGo task: A definitive study

This study examined the occurrence of preferred EEG phase states at stimulus onset in an equiprobable auditory Go/NoGo task with a fixed interstimulus interval, and their effects on the resultant event-related potentials (ERPs). We used a sliding short-time FFT decomposition of the EEG at Cz for each trial to assess prestimulus EEG activity in the delta, theta, alpha and beta bands. We determined the phase of each 2 Hz narrow-band contributing to these four broad bands at 125 ms before each stimulus onset, and for the first time, avoided contamination from poststimulus EEG activity. This phase value was extrapolated 125 ms to obtain the phase at stimulus onset, combined into the broad-band phase, and used to sort trials into four phase groups for each of the four broad bands. For each band, ERPs were derived for each phase from the raw EEG activity at 19 sites. Data sets from each band were separately decomposed using temporal Principal Components Analyses with unrestricted VARIMAX rotation to extract N1-1, PN, P2, P3, SW and LP components. Each component was analysed as a function of EEG phase at stimulus onset in the context of a simple conceptualisation of orthogonal phase effects (cortical negativity vs. positivity, negative driving vs. positive driving, waxing vs. waning). The predicted non-random occurrence of phase-defined brain states was confirmed. The preferred states of negativity, negative driving, and waxing were each associated with more efficient stimulus processing, as reflected in amplitude differences of the components. The present results confirm the existence of preferred brain states and their impact on the efficiency of brain dynamics in perceptual and cognitive processing.     

Spatio-temporal Modeling of Evoked Brain Activity During Memory Encoding and Target Comparison in Visual Tasks

In this paper, the temporal pattern of activity and approximate locations of brain areas related to selective attention and visual working memory processes were studied with event related potential (ERP) recordings in healthy humans. Three experimental series included pairs of the following conditions: Face comparison (familiar faces), Pattern comparison (abstract dot patterns), and Passive viewing. Participants compared pairs of consecutive targets presented in composite images on a computer screen. Spatio-temporal multiple dipole models were developed for 128-channel ERPs. Keeping dipole locations and orientations constant, we compared the source activities for ERPs recorded (1) in different tasks for task-specificity of activations, (2) after the first and second stimuli in the pair, i.e. on the encoding and comparison stages of the task, and (3) after the first stimulus in different series to compare encoding in different conditions. Sources located in the inferotemporal brain areas, especially in the left hemisphere, showed increased activity after 200 ms from the first stimulus onset that may indicate encoding into visual working memory. The anterior sources, located near midline and showing activity around or after 300 ms, presumably reflect non-specific memory processes and attentional control. Major task-specific differences were observed in the temporo-parieto-occipital region in 250–500 ms.     

Working memory constraints on syntactic processing: an electrophysiological investigation

Event-related potentials (ERPs) and reaction times (RTs) were used to study how the processing of sentences with morphosyntactic violations is constrained by working memory (WM) capacity. The available WM capacity was varied by three orthogonal manipulations: (1) syntactic complexity; (2) additional WM load; and (3) verbal WM span. The processing of the morphosyntactic violations was reflected in longer RTs in ungrammatical compared with grammatical sentences, and in an anterior negativity and a centroparietal positivity in the ERPs. While the behavioral grammaticality effect was not influenced by the WM manipulations, the ERP effects were. The amplitude of the anterior negativity was modulated by the combination of complexity and load, and by WM span. The onset of the centroparietal positivity was delayed in the high-load condition, and for the low-span group. ERPs over the course of the sentences showed a frontal negative slow wave under high WM load, largest for the low-span group. The finding that online syntactic processing is related to WM span and to additional WM load does not support the theory that there is a WM capacity specific for syntactic processing.     

The human brain processes visual changes that are not cued by attended auditory stimulation

Event-related potentials (ERPs) to visual stimuli were recorded from the scalp of eight adult humans performing a task in which they counted vowels from a heard story. In the oddball condition, a repeated (standard) light bar of 50 ms in duration was rarely (P = 0.1) replaced by a (deviant) one differing in orientation from the standard. In the control condition, standards were simply omitted from the series and only (alone-) deviants retained. In both conditions, visual stimuli were asynchronous with auditory-task-relevant stimuli. ERPs to deviants significantly differed in amplitude from those to standards in the midline electrodes centrally, parietally and occipitally at 160-200 ms from stimulus onset. Occipitally, such a difference was absent between ERPs to alone-deviants and those to standards. The occipital differential ERPs to deviants, which thus could be found only when standards were present in the series, are discussed in the context of the mismatch negativity (MMN).     

Stimulus presentation rate dissociates sequential effects in event-related potentials and reaction times

The present study explored the impact of the stimulus presentation rate on sequential effects in event-related potentials (ERPs) and reaction times (RTs). Random series of equiprobable tones were presented at interstimulus intervals (ISIs) of 1.3, 2.1, and 2.9s. Fast and accurate choice responses to the tones were required. Although sequential effects in RTs were stable across all ISIs, the common first-order repetition effect in P300 amplitude was only observed at the 1.3-s ISI and not at the slower presentation rates. This dissociation between the first-order effects in RTs and ERPs speaks against an explanation of both effects by a common expectancy mechanism. In addition, sequential effects were observed as early as about 100 ms after stimulus onset in the lateralized readiness potential. Together with similar sequential effects in P300 latency, this finding supports a continuous flow model of information processing.     

Event related potentials to emotional adjectives during reading

We investigated to what extent emotional connotation influences cortical potentials during reading. To this end, event-related potentials (ERPs) were recorded during reading of high arousal pleasant and unpleasant and low arousal neutral adjectives that were presented at rates of 1 Hz and 3 Hz. Enhanced processing of both pleasant and unpleasant emotional compared to neutral adjectives was first reflected in an amplified early posterior negativity (EPN) starting from 200 ms after word onset. Later potentials (>300 ms), as analyzed in the slower 1 Hz condition, revealed facilitated processing selectively for pleasant adjectives that were associated with a reduced N400 and an enhanced late positive potential (LPP). Pleasant adjectives were also better remembered in an incidental memory test. Thus, emotionally relevant adjectives are processed spontaneously and selectively. Initially, emotional arousal drives attention capture (EPN). Healthy subjects may have a natural bias toward pleasant information facilitating late ERPs (N400, LPP) to pleasant adjectives as well as their superior recall.