Event-related Potential Study of Novelty Processing Abnormalities in Autism

To better understand visual processing abnormalities in autism we studied the attention orienting related frontal event potentials (ERP) and the sustained attention related centro-parietal ERPs in a three stimulus oddball experiment. The three stimulus oddball paradigm was aimed to test the hypothesis that individuals with autism abnormally orient their attention to novel distracters as compared to controls. A dense-array 128 channel EGI electroencephalographic (EEG) system was used on 11 high-functioning children and young adults with autism spectrum disorder (ASD) and 11 age-matched, typically developing control subjects. Patients with ASD showed slower reaction times but did not differ in response accuracy. At the anterior (frontal) topography the ASD group showed significantly higher amplitudes and longer latencies of early ERP components (e.g., P100, N100) to novel distracter stimuli in both hemispheres. The ASD group also showed prolonged latencies of late ERP components (e.g., P2a, N200, P3a) to novel distracter stimuli in both hemispheres. However, differences were more profound in the right hemisphere for both early and late ERP components. Our results indicate augmented and prolonged early frontal potentials and a delayed P3a component to novel stimuli, which suggest low selectivity in pre-processing and later-stage under-activation of integrative regions in the prefrontal cortices. Also, at the posterior (centro-parietal) topography the ASD group showed significantly prolonged N100 latencies and reduced amplitudes of the N2b component to target stimuli. In addition, the latency of the P3b component was prolonged to novel distracters in the ASD group. In general, the autistic group showed prolonged latencies to novel stimuli especially in the right hemisphere. These results suggest that individuals with autism over-process information needed for the successful differentiation of target and novel stimuli. We propose the potential application of ERP evaluations in a novelty task as outcome measurements in the biobehavioral treatment (e.g., EEG biofeedback, TMS) of autism.     

Amplitude modulation of steady-state visual evoked potentials by event-related potentials in a working memory task

Previous studies have shown that the amplitude and phase of the steady-state visual-evoked potential (SSVEP) can be influenced by a cognitive task, yet the mechanism of this influence has not been understood. As the event-related potential (ERP) is the direct neural electric response to a cognitive task, studying the relationship between the SSVEP and ERP would be meaningful in understanding this underlying mechanism. In this work, the traditional average method was applied to extract the ERP directly, following the stimulus of a working memory task, while a technique named steady-state probe topography was utilized to estimate the SSVEP under the simultaneous stimulus of an 8.3-Hz flicker and a working memory task; a comparison between the ERP and SSVEP was completed. The results show that the ERP can modulate the SSVEP amplitude, and for regions where both SSVEP and ERP are strong, the modulation depth is large.     

Functional brain organization of global and local visual perception: an ERP study

Adult subjects were asked to recognize a hierarchical visual stimulus (a letter) while their attention was drawn to either the global or local level of the stimulus. Event-related potentials (ERP) and psychophysical indices (reaction time and percentage of correct responses) were measured. An analysis of psychophysical indices showed the global level precedence effect, i.e., the increase in a small letter recognition time when this letter is a part of incongruent stimulus. An analysis of ERP components showed level-related (global vs. local) differences in the timing and topography of the brain organization of perceptual processing and regulatory mechanisms of attention. Visual recognition at the local level was accompanied by (1) stronger activation of the visual associative areas (Pz and T6) at the stage of sensory features analysis (P1 ERP component), (2) involvement mainly of inferior temporal cortices of the right hemisphere (T6) at the stage of sensory categorization (P2 ERP component), and (3) involvement of prefrontal cortex of the right hemisphere at the stage of the selection of the relevant features of the target (N2 ERP component). Visual recognition at the global level was accompanied by (1) pronounced involvement of mechanisms of early sensory selection (N1 ERP component), (2) prevailing activation of parietal cortex of the right hemisphere (P4) at the stage of sensory categorization (P2 ERP component) as well as at the stage of the target stimulus identification (P3 ERP component). It is suggested that perception at the global level of the hierarchical stimulus is related primarily to the analysis of the spatial features of the stimulus in the dorsal visual system whereas the perception at the local level primarily involves an analysis of the object-related features in the ventral visual system.     

Characteristics of event-related potentials in response to symbolical and alphabetical stimulation matrices used in a P300-based brain-computer interface

In order to create a P300-based brain-computer interface (BCI) (the so-called Farwell-Donchin paradigm, FD) with a symbol matrix used as a stimulus, we compared characteristics of event-related potentials (ERPs) in response to stimulation by 6 × 6 matrices composed of either pictogram symbols or Cyrillic alphabet characters. Nine healthy adults were examined in 18 experiments, during which 28-channel EEGs were recorded in the course of stimulation with matrices of these two types. The obtained ERP data, i.e., amplitudes and peak latencies of the ERP components N1, P3 (with the P3a and P3b sub-components), and N4 were compared and analyzed for different types of stimulation matrices. In at least seven out of nine subjects, P3a, P3b, and N4 ERP amplitudes were larger in response to the symbol matrix than to the character matrix, while N1 amplitudes were larger for the character matrix. For N1 and P3a, the ERP latencies were shorter for the symbol matrix, while for P3b and N4, they were longer for the character matrix. The topography of differential ERP responses to the two types of stimuli was analyzed using a series of paired t-tests. Differences of ERP component amplitudes were determined individually for each of the 28 channels; next, for each site, absolute t-test values were summed for all nine subjects. For all ERP components studied, the t-test for peak amplitudes in response to target and non-target letters identified two separate areas with distinct lateralization. ERP responses to target and non-target symbols differed most in transversely extended areas. Finally, the yield surface of differential response to target letters and target symbols had a complex topography.     

Erratum to: Functional Brain Organization of Global and Local Visual Perception: An Event-Related Potentials Study

Adult subjects were asked to recognize a hierarchical visual stimulus (a letter) while their attention was drawn to either the global or local level of the stimulus. Event-related potentials (ERP) and behavioral indices (reaction time and percentage of correct responses) were measured. An analysis of behavioral indices showed the global level precedence effect, i.e. the increase in a small letter recognition time when this letter is a part of incongruent stimulus. An analysis of ERP components showed level-related (global vs. local) differences in the timing and topography of the brain organization of perceptual processing and regulatory mechanisms of attention. Visual recognition at the local level was accompanied by (1) stronger activation of the visual associative areas (P _z and T ₆) at the stage of sensory features analysis (P1 ERP component), (2) involvement mainly of inferior temporal cortices of the right hemisphere (T ₆) at the stage of sensory categorization (P2 ERP component), and (3) involvement of prefrontal cortex of the right hemisphere at the stage of selection of the relevant features of the target (N2 ERP component). Visual recognition at the global level was accompanied by (1) pronounced involvement of mechanisms of early sensory selection (N1 ERP component), (2) prevailing activation of parietal cortex of the right hemisphere (P ₄) at the stage of sensory categorization (P2 ERP component) as well as at the stage of the target stimulus identification (P3 ERP component). We suggested that perception of the hierarchical stimulus at the global level is related primarily to the analysis of its spatial features in the dorsal visual system whereas the perception at the local level primarily involves an analysis of the object-related features in the ventral visual system.     

Role of the Frontal Cortical Areas in the Analysis of Visual Stimuli at Conscious and Unconscious Levels

Event-related potentials (ERP) in response to complex target stimuli, which consisted of a central recognizable picture and a lateral masked image (analyzed at the unconscious level) were recorded in adult subjects and seven-year-old children. ERP components N200, N300, and P400/N400 had different topography and were differently pronounced in adults and children. In adult subjects, the N200 component that reflects the processing of a sensory stimulus was recorded in the temporo-parieto-occipital and occipital areas. In children, N200 was recorded in the caudal regions and the frontal areas of the cortex. Analysis of different waveforms obtained by subtraction of the ERP to the central stimulus from the ERP to the complex stimulus showed that unconscious stimulus processing in adult subjects is not reflected in the ERP structure. In children, an unconsciously processed image incorporated into a complex stimulus evokes processing negativity in the occipital and frontal cortical areas. Comparison of ERP in groups of children divided by their reflectivity/impulsivity showed that, predominantly, the left frontal area is involved in image analysis at the unconscious level in reflective children and, predominantly, the right frontal area participates in unconscious image analysis in impulsive children. It is suggested that the perfection of the visual recognition of a target stimulus, which contains additional unconsciously processed information, consists in growth of the involvement of the left-hemispheric mechanisms (with respective growth of significance of the left-hemispheric mechanisms) and in a decrease in the role of the frontal areas in analysis of sensory information.     

P300 assessment of early Alzheimer's disease

The P300 (P3) event-related brain potential (ERP) was elicited in 16 demented patients presumed to be in the early stages of Alzheimer's disease and 16 normal control subjects well matched for age, sex, education and occupational level. All subjects performed a simple auditory discrimination task in which a target tone was presented randomly on 20% of the trials. P3 amplitude was smaller and peak latency longer for the Alzheimer patients compared to control subjects. A second ERP task also was administered in which the target tone occurred 50% of the time. Analysis of the target/standard tone presentation sequences indicated that the Alzheimer patient group demonstrated less amplitude difference between the target and standard sequences and longer overall latencies compared to the control group. The results that the P3 ERP component from auditory stimuli can provide useful information about Alzheimer's disease during its early stages.     

Heritable features of the auditory oddball event-related potential: peaks,latencies, morphology and topography

Baseline auditory ERP data from a larger study of the genetic determinants of the response to alcohol were collected from 59 monozygotic (MZ) twin pairs and from 39 same-sex dizygotic (DZ) twin pairs who drank socially. Three methods for measuring genetic influence on the ERPs were applied. First, based on maximum-likelihood estimates, the heritability of conventional peak amplitude and latency of N1 and P3 components was computed for each of 16 lead locations using tests of the significance of heritability based on intraclass correlations. P3 amplitude provided the strongest results, distributed symmetrically over caudal leads, and implied gene dominance as the mode of genetic transmission for the P3 component. A substantial genetic influence on N1 latency suggested a mixture of additive and dominance effects in the left fronto-temporal region. N1 amplitude measures trended towards significant heritability, but none was observed for P3 latency. The second method used the maximum of the cross-correlation function to compare wave form shape in a lead-by-lead analysis of data from cotwins. Genetic influence was apparent in both target and non-target ERP responses, with a fronto-central topography of significant results. The third method reduced all spatial and temporal ERP differences from a pair of twins to a single scalar number for each response. Distributions of this global measure revealed significant genetic influence on both non-target and target ERPs. A post hoc analysis of the effect of gender on the heritability of N1 or P3 peaks and latencies revealed no statistically significant observations in this sample of young adult twins.     

Pain-related and cognitive components of somatosensory evoked potentials following CO2 laser stimulation in man

We recorded cortical potentials evoked by painful CO₂ laser stimulation (pain SEP), employing an oddball paradigm in an effort to demonstrate event-related potentials (ERP) associated with pain. In 12 healthy subjects, frequent (standard) pain stimuli (probability 0.8) were delivered to one side of the dorsum of the left hand while rare (target) pain stimuli (probability 0.2) were delivered to the other side of the same hand. Subjects were instructed to perform either a mental count or button press in response to the target stimuli. Two early components (N2 and P2) of the pain SEP demonstrated a Cz maximal distribution, and showed no difference in latency, amplitude or scalp topography between the oddball conditions or between response tasks. In addition, another positive component (P3) following the P2 was recorded maximally at Pz only in response to the target stimuli with a peak latency of 593 msec for the count task and 560 msec for the button press task. Its scalp topography was the same as that for electric and auditory P3. The longer latency of pain P3 can be explained not only by its slower impulse conduction but also by the effects of task difficulty in the oddball paradigm employing the pain stimulus compared with electric and auditory stimulus paradigms. It is concluded that the P3 for the pain modality is mainly related to a cognitive process and corresponds to the P3 of electric and auditory evoked responses, whereas both N2 and P2 are mainly pain-related components.     

Functional brain organization of global and local visual perception: Analysis of event-related potentials

Event-related potentials (ERP) of the brain and psychometric indices (reaction time and percentage of correct responses) were studied in adult subjects during recognizing hierarchical visual stimuli (letters), while the subject’s attention was drawn to either the global or the local level of the stimulus. The psychophysical indices demonstrated the global precedence effect, i.e., an increased recognition time of a small letter, which was a part of an incongruent stimulus. The ERP component analysis demonstrated that differences in the regulatory mechanisms of attention and timing and topography of brain organization during processing of visual information depended on the level of recognizing the hierarchical stimulus (global vs. local). Visual recognition at the local level was accompanied by a stronger activation of visual associative areas (P _z and T ₆) at the stage of sensory feature analysis (P1 ERP component), as well as by the predominant involvement of the temporal inferior cortex of the right hemisphere (T ₆) at the stage of sensory categorization (the P2 ERP component) and of the frontal cortex of the right hemisphere at the stage of selection for the relevant target features (the N2 ERP component). Visual recognition at the global level was accompanied by significant involvement of the early sensory selection (the N1 ERP component) and predominant activation of the parietal cortex of the right hemisphere (P ₄) at the stage of sensory categorization (the P2 ERP component), as well as at the stage of identification of the target stimulus (the P3 ERP component). Perception of a stimulus at the global level is assumed to depend mostly on the analysis of its spatial features in the dorsal visual system, whereas perception at the local level involves analysis of the object-related features in the ventral visual system.     

Analysis of the rhodopsin cycle in limulus ventral photoreceptors using the early receptor potential

The early receptor potential (ERP) was recorded intracellularly from Limulus ventral photoreceptors. The ERP in cells dissected under red light was altered by exhaustive illumination. No recovery to the original wafeform was observed, even after 1 h in the dark. The ERP waveform could be further altered by chromatic adaptation or by changes in pH. The results indicate that at pH 7.8 there are two interconvertible pigment states with only slightly different lambdamax, whereas at pH 9.6 there are two interconvertible states with very different lambdamax. Under all conditions studied the ERPs were almost identical with those previously obtained in squid retinas. This strongly suggests that light converts Limulus rhodopsin to a stable photoequilibrium mixture of rhodopsin to a stable photoequilibrium mixture of rhodopsin and metarhodopsin and that, as in squid, the lambdamax of metarhodopsin depends on pH. This conversion at pH 7.8 is associated with a small (0.7 log unit) decrease in the maximum sensitivity of the late receptor potential. Thus the component of adaptation linked to changes in rhodopsin concentration is unimportant in comparison to the "neural" component.     

Topography of the N400: brain electrical activity reflecting semantic expectancy

When subjects read an semantically unexpected word, the brain electrical activity shows a negative deflection at about 400 msec in comparison with the response to an expected word. In order to study the brain systems related to this effect we mapped it with a dense (64-channel) electrode array and two reference-independent measures, one estimating the average potential gradients and the other radial current density. With these measures, the event-related brain potential (ERP) begins at about 70 msec with the P1, reflecting bilateral current sources over occipitoparietal areas. A strongly left-lateralized N1 then follows, peaking at about 180 msec, accompanied by an anterior positivity, the P2. A separate posterior positive pattern then emerges that seems to repeat the topography of the P1. Next, at about 350 msec, the ERP for the congruous word develops a P300 or LPC, characterized by a diffuse positivity over the superior surface of the head and several negativities over inferior regions. This superior source/inferior sink pattern of the LPC is greater over the left hemisphere. In contrast, the ERP for the incongruous word in this interval displays the N400 as a period in which topographic features are absent. At about 400 msec the ERP for the incongruous word begins to develop an LPC, which then remains relatively symmetric over the two hemispheres.     

Early Receptor Potential Evidence for the Existence of Two Thermally Stable States in the Barnacle Visual Pigment

The early receptor potential (ERP) in the barnacle photoreceptor is shown by intracellular recording to exhibit a strong dependence on the color of the stimulus and of the preceding adaptation. The adaptation effects appear to be stable for at least 3 h in the dark. Most strikingly, the ERP is positive after red adaptation and mainly negative after blue adaptation. The simplest hypothesis which accounts for these observations is that two thermally stable pigment states with different absorption spectra contribute to the ERP. All ERP responses appear to be consistent with the sums of different ratios of the ERP's of the two pure states. The relative populations of the two states are shown to vary reciprocally, suggesting that the two are states of the same closed pigment cycle. Both states have approximately Dartnall nomogram-shaped absorption spectra, one peaked near 495 nm, and the other near 532 nm.     

Target and Non-Target Processing during Oddball and Cyberball: A Comparative Event-Related Potential Study

The phenomenon of social exclusion can be investigated by using a virtual ball-tossing game called Cyberball. In neuroimaging studies, structures have been identified which are activated during social exclusion. But to date the underlying mechanisms are not fully disclosed. In previous electrophysiological studies it was shown that the P3 complex is sensitive to exclusion manipulations in the Cyberball paradigm and that there is a correlation between P3 amplitude and self-reported social pain. Since this posterior event-related potential (ERP) was widely investigated using the oddball paradigm, we directly compared the ERP effects elicited by the target (Cyberball: “ball possession”) and non-target (Cyberball: “ball possession of a co-player) events in both paradigms. Analyses mainly focused on the effect of altered stimulus probabilities of the target and non-target events between two consecutive blocks of the tasks. In the first block, the probability of the target and non-target event was 33% (Cyberball: inclusion), in the second block target probability was reduced to 17%, and accordingly, non-target probability was increased to 66% (Cyberball: exclusion). Our results indicate that ERP amplitude differences between inclusion and exclusion are comparable to ERP amplitude effects in a visual oddball task. We therefore suggest that ERP effects–especially in the P3 range–in the Oddball and Cyberball paradigm rely on similar mechanisms, namely the probability of target and non-target events. Since the simulation of social exclusion (Cyberball) did not trigger a unique ERP response, the idea of an exclusion-specific neural alarm system is not supported. The limitations of an ERP-based approach will be discussed.     

Selective processing of multiple features in the human brain: effects of feature type and salience

Identifying targets in a stream of items at a given constant spatial location relies on selection of aspects such as color, shape, or texture. Such attended (target) features of a stimulus elicit a negative-going event-related brain potential (ERP), termed Selection Negativity (SN), which has been used as an index of selective feature processing. In two experiments, participants viewed a series of Gabor patches in which targets were defined as a specific combination of color, orientation, and shape. Distracters were composed of different combinations of color, orientation, and shape of the target stimulus. This design allows comparisons of items with and without specific target features. Consistent with previous ERP research, SN deflections extended between 160-300 ms. Data from the subsequent P3 component (300-450 ms post-stimulus) were also examined, and were regarded as an index of target processing. In Experiment A, predominant effects of target color on SN and P3 amplitudes were found, along with smaller ERP differences in response to variations of orientation and shape. Manipulating color to be less salient while enhancing the saliency of the orientation of the Gabor patch (Experiment B) led to delayed color selection and enhanced orientation selection. Topographical analyses suggested that the location of SN on the scalp reliably varies with the nature of the to-be-attended feature. No interference of non-target features on the SN was observed. These results suggest that target feature selection operates by means of electrocortical facilitation of feature-specific sensory processes, and that selective electrocortical facilitation is more effective when stimulus saliency is heightened.     

P300 from a single auditory stimulus

The P3(00) event-related brain potential (ERP) was elicited with auditory stimuli to compare 2 different discrimination tasks. The oddball paradigm presented both target and standard tones; the single-stimulus paradigm presented at target but no standard tone stimulus. Experiment 1 manipulated target stimulus probability (0.20, 0.50, 0.80) and produced highly similar P3 amplitude and latency results across probability levels for each paradigm. Experiment 2 factorially varied inter-stimulus interval (2 sec, 6 sec) and target stimulus probability (0.20, 0.80). P3 amplitude and latency were highly similar for both the oddball and single-stimulus procedures across all conditions.     

Antagonistic Components of the Late Receptor Potential in the Barnacle Photoreceptor Arising from Different Stages of the Pigment Process

The late receptor potential (LRP) recorded in barnacle photoreceptor cells exhibits, at high light levels, a strong dependence on the color of the stimulus and of the preceding adaptation. Most strikingly, red illumination of a cell previously adapted to blue light results in a depolarization which may last for up to 30 min after the light goes off, while blue illumination of a cell previously adapted to red light cuts short this extended depolarization or prevents its induction by a closely following red light. Comparison of the action spectra for the stimulus-coincident LRP and for the extended depolarization and its curtailment with those previously measured for the early receptor potential (ERP) confirms that these phenomena derive from the same bi-stable pigment as the ERP. The stimulus-coincident response and the extended depolarization appear to arise from substantial activation of the stable 532 nm state of the pigment, while activation of the stable 495 state depresses or prevents the extended depolarization and probably also depresses the stimulus-coincident response. Since either process can precede the other, with mutually antagonistic effects, one is not simply the reversal of the other; they must be based on separate mechanisms. Furthermore, comparison with ERP kinetics shows that both processes involve mechanisms additional to the pigment changes, as seen in the ERP. A model is proposed and discussed for the LRP phenomena and their dependences on wavelength, intensity, and duration of illumination based on excitor-inhibitor interactions.     

Competitive Semantic Memory Retrieval: Temporal Dynamics Revealed by Event-Related Potentials

Memories compete for retrieval when they are related to a common retrieval cue. Previous research has shown that retrieval of a target memory may lead to subsequent retrieval-induced forgetting (RIF) of currently irrelevant competing memories. In the present study, we investigated the time course of competitive semantic retrieval and examined the neurocognitive mechanisms underlying RIF. We contrasted two theoretical accounts of RIF by examining a critical aspect of this memory phenomenon, namely the extent to which it depends on successful retrieval of the target memory. Participants first studied category-exemplar word-pairs (e.g. Fruit—Apple). Next, we recorded electrophysiological measures of brain activity while the participants performed a competitive semantic cued-recall task. In this task, the participants were provided with the studied categories but they were instructed to retrieve other unstudied exemplars (e.g. Fruit—Ma__?). We investigated the event-related potential (ERP) correlates of retrieval success by comparing ERPs from successful and failed retrieval trials. To isolate the ERP correlates of continuous retrieval attempts from the ERP correlates of retrieval success, we included an impossible retrieval condition, with incompletable word-stem cues (Drinks—Wy__) and compared it with a non-retrieval presentation baseline condition (Occupation—Dentist). The participants’ memory for all the studied exemplars was tested in the final phase of the experiment. Taken together, the behavioural results suggest that RIF is independent of target retrieval. Beyond investigating the mechanisms underlying RIF, the present study also elucidates the temporal dynamics of semantic cued-recall by isolating the ERP correlates of retrieval attempt and retrieval success. The ERP results revealed that retrieval attempt is reflected in a late posterior negativity, possibly indicating construction of candidates for completing the word-stem cue and retrieval monitoring whereas retrieval success was reflected in an anterior positive slow wave.     

Topographic analysis of auditory event-related potentials associated with acoustic and semantic processing

The topography of auditory event-related potentials (ERPs) was examined during 3 kinds of tasks: selection of a specified real word or nonsense syllable from a list; simple detection of each of the same stimuli without discrimination; and classification of a set of words according to a specified semantic category. The potentials that were associated with the additional processing required by the discriminative tasks were disclosed by subtracting the wave forms obtained in the detection condition from those obtained during discriminative performance. Difference wave forms were also derived between the semantic classification and verbal discriminative ERP to delineate the changes associated with the extraction of word meaning.The topography of the ERP associated with stimulus detection was comparable to that found in previous studies of evoked potentials to non-speech stimuli. This distribution was consistent with 2 cortical generators, one within the supratemporal plane and the other on the lateral surface of the superior temporal gyrus. When discriminative performance was required on the basis of acoustic stimulus properties, the topography of the difference wave form that reflected this discriminative processing extended more posteriorly over temporal cortex. Semantic processing elicited a further posterior extension of ERP components by 330 msec after stimulus onset, as well as longer latency potentials that were not present in the verbal selection task. These differences imply that a more extensive portion of language cortex is engaged in semantic classification than in verbal identification.     

Electromagnetic field of mobile phones affects visual event related potential in patients with narcolepsy

The effects of the mobile phone (MP) electromagnetic fields on electroencephalography (EEG) and event-related potentials (ERP) were examined. With regard to the reported effects of MP on sleep, 22 patients with narcolepsy-cataplexy were exposed or sham exposed for 45 min to the MP (900 MHz, specific absorption rate 0.06 W/kg) placed close to the right ear in a double blind study. There were no changes of the EEG recorded after the MP exposure. A subgroup of 17 patients was studied on visual ERP recorded during the MP exposure. Using an adapted "odd-ball" paradigm, each patient was instructed to strike a key whenever rare target stimuli were presented. There were three variants of target stimuli (horizontal stripes in (i) left, (ii) right hemifields or (iii) whole field of the screen). The exposure enhanced the positivity of the ERP endogenous complex solely in response to target stimuli in the right hemifield of the screen (P < 0.01). The reaction time was shortened by 20 ms in response to all target stimuli (P < 0.05). In conclusion, the electromagnetic field of MP may suppress the excessive sleepiness and improve performance while solving a monotonous cognitive task requiring sustained attention and vigilance.