Electrophysiological correlates of rapid spatial orienting towards fearful faces

We investigated the spatio-temporal dynamic of attentional bias towards fearful faces. Twelve participants performed a covert spatial orienting task while recording visual event-related brain potentials (VEPs). Each trial consisted of a pair of faces (one emotional and one neutral) briefly presented in the upper visual field, followed by a unilateral bar presented at the location of one of the faces. Participants had to judge the orientation of the bar. Comparing VEPs to bars shown at the location of an emotional (valid) versus neutral (invalid) face revealed an early effect of spatial validity: the lateral occipital P1 component (approximately 130 ms post-stimulus) was selectively increased when a bar replaced a fearful face compared to when the same bar replaced a neutral face. This effect was not found with upright happy faces or inverted fearful faces. A similar amplification of P1 has previously been observed in electrophysiological studies of spatial attention using non-emotional cues. In a behavioural control experiment, participants were also better at discriminating the orientation of the bar when it replaced a fearful rather than a neutral face. In addition, VEPs time-locked to the face-pair onset revealed a C1 component (approximately 90 ms) that was greater for fearful than happy faces. Source localization (LORETA) confirmed an extrastriate origin of the P1 response showing a spatial validity effect, and a striate origin of the C1 response showing an emotional valence effect. These data suggest that activity in primary visual cortex might be enhanced by fear cues as early as 90 ms post-stimulus, and that such effects might result in a subsequent facilitation of sensory processing for a stimulus appearing at the same location. These results provide evidence for neural mechanisms allowing rapid, exogenous spatial orienting of attention towards fear stimuli.     

Electrophysiological evidence for temporal dissociation between spatial attention and sensory competition during human face processing

Scalp event-related potential (ERP) studies in humans indicate that face processes taking place between 130 and 170 ms after stimulus onset at posterior sites (N170) are strongly reduced when another face stimulus is processed concurrently or has been presented shortly before for a prolonged period. These observations suggest that neural representations of individual faces compete in the occipitotemporal cortex as early as 130 ms. Here, we tested the respective role of spatial attention and sensory competition in accounting for the amplitude reduction of the N170 during concurrent face stimulation. ERPs time locked to a lateralized face stimulus were recorded while subjects were fixating either a face or a controlled scrambled-face stimulus (context factor) and were engaged in either a high- or a low-attentional load task at fixation (task factor). The N170 amplitude to the lateralized face stimulus was reduced both when the central stimulus was a face compared with a scrambled face and when the attentional load at fixation was high. However, these effects of context and task factors were largely additive. Most importantly, spatial attention modulated visual processes as early as 80 ms after stimulus onset, whereas sensory competition effects started at about 130 ms. These results provide strong evidence that the N170 in response to faces is modulated by spatial attention, and also that spatial attention and sensory competition do not reflect the same mechanisms of early selection of visual information in the extrastriate cortex.     

Attention modulates gamma-band oscillations differently in the human lateral occipital cortex and fusiform gyrus

We studied the existence, localization and attentional modulation of gamma-band oscillatory activity (30-130 Hz) in the human intracranial region. Two areas known to play a key role in visual object processing: the lateral occipital (LO) cortex and the fusiform gyrus. These areas consistently displayed large gamma oscillations during visual stimulus encoding, while other extrastriate areas remained systematically silent, across 14 patients and 291 recording sites scattered throughout extrastriate visual cortex. The lateral extent of the responsive regions was small, in the range of 5 mm. Induced gamma oscillations and evoked potentials were not systematically co-localized. LO and the fusiform gyrus displayed markedly different patterns of attentional modulation. In the fusiform gyrus, attention enhanced stimulus-driven gamma oscillations. In LO, attention increased the baseline level of gamma oscillations during the expectation period preceding the stimulus. Subsequent gamma oscillations produced by attended stimuli were smaller than those produced by unattended, irrelevant stimuli. Attentional modulations of gamma oscillations in LO and the fusiform gyrus were thus very different, both in their time-course (preparatory period and/or stimulus processing) and direction of modulation (increase or decrease). Our results thus suggest that the functional role of gamma oscillations depends on the area in which they occur.     

Volatile anesthetics enhance flash-induced gamma oscillations in rat visual cortex

BACKGROUND: The authors sought to understand neural correlates of anesthetic-induced unconsciousness. Cortical gamma oscillations have been associated with neural processes supporting conscious perception, but the effect of general anesthesia on these oscillations is controversial. In this study, the authors examined three volatile anesthetics, halothane, isoflurane, and desflurane, and compared their effects on flash-induced gamma oscillations in terms of equivalent concentrations producing the loss of righting reflex (1 minimum alveolar concentration for the loss of righting [MAC(LR)]). METHODS: Light flashes were presented every 5 s for 5 min, and event-related potentials were recorded from primary visual cortex of 15 rats with a chronically implanted bipolar electrode at increasing anesthetic concentrations (0-2.4 MAC(LR)). Early cortical response was obtained by averaging poststimulus (0-100 ms) potentials filtered at 20-60 Hz across 60 trials. Late (100-1,000 ms) gamma power was calculated using multitaper power spectral technique. Wavelet decomposition was used to determine spectral and temporal distributions of gamma power. RESULTS: The authors found that (1) halothane, isoflurane, and desflurane enhanced the flash-evoked early cortical response in a concentration-dependent manner; (2) the effective concentration for this enhancement was the lowest for isoflurane, intermediate for halothane, and the highest for desflurane when compared at equal fractions of the concentration that led to a loss of righting; (3) the power of flash-induced late (> 100 ms) gamma oscillations was augmented at intermediate concentrations of all three anesthetic agents; and (4) flash-induced gamma power was not reduced below waking baseline even in deep anesthesia. CONCLUSIONS: These findings suggest that a reduction in flash-induced gamma oscillations in rat visual cortex is not a unitary correlate of anesthetic-induced unconsciousness.     

Neural sources of focused attention in visual search

Previous studies of visual search in humans using event-related potentials (ERPs) have revealed an ERP component called 'N2pc' (180-280 ms) that reflects the focusing of attention onto potential target items in the search array. The present study was designed to localize the neuroanatomical sources of this component by means of magnetoencephalographic (MEG) recordings, which provide greater spatial precision than ERP recordings. MEG recordings were obtained with an array of 148 magnetometers from six normal adult subjects, one of whom was tested in multiple sessions so that both single-subject and group analyses could be performed. Source localization procedures revealed that the N2pc is composed of two distinct neural responses, an early parietal source (180-200 ms) and a later occipito-temporal source (220-240 ms). These findings are consistent with the proposal that parietal areas are used to initiate a shift of attention within a visual search array and that the focusing of attention is implemented by extrastriate areas of the occipital and inferior temporal cortex.     

Electrophysiological studies of human face perception. I: Potentials generated in occipitotemporal cortex by face and non-face stimuli.

This and the following two papers describe event-related potentials (ERPs) evoked by visual stimuli in 98 patients in whom electrodes were placed directly upon the cortical surface to monitor medically intractable seizures. Patients viewed pictures of faces, scrambled faces, letter-strings, number-strings, and animate and inanimate objects. This paper describes ERPs generated in striate and peristriate cortex, evoked by faces, and evoked by sinusoidal gratings, objects and letter-strings. Short-latency ERPs generated in striate and peristriate cortex were sensitive to elementary stimulus features such as luminance. Three types of face-specific ERPs were found: (i) a surface-negative potential with a peak latency of approximately 200 ms (N200) recorded from ventral occipitotemporal cortex, (ii) a lateral surface N200 recorded primarily from the middle temporal gyrus, and (iii) a late positive potential (P350) recorded from posterior ventral occipitotemporal, posterior lateral temporal and anterior ventral temporal cortex. Face-specific N200s were preceded by P150 and followed by P290 and N700 ERPs. N200 reflects initial face-specific processing, while P290, N700 and P350 reflect later face processing at or near N200 sites and in anterior ventral temporal cortex. Face-specific N200 amplitude was not significantly different in males and females, in the normal and abnormal hemisphere, or in the right and left hemisphere. However, cortical patches generating ventral face-specific N200s were larger in the right hemisphere. Other cortical patches in the same region of extrastriate cortex generated grating-sensitive N180s and object-specific or letter-string-specific N200s, suggesting that the human ventral object recognition system is segregated into functionally discrete regions.     

Large-scale visuomotor integration in the cerebral cortex

Efficient visuomotor behavior depends on integrated processing by the visual and motor systems of the cerebral cortex. Yet, many previous cortical neurophysiology studies have examined the visual and motor modalities in isolation, largely ignoring questions of large-scale cross-modal integration. To address this issue, we analyzed event-related local field potentials simultaneously recorded from multiple visual, motor, and executive cortical sites in monkeys performing a visuomotor pattern discrimination task. The timing and cortical location of four aspects of event-related activities were examined: stimulus-evoked activation onset, stimulus-specific processing, stimulus category-specific processing, and response-specific processing. Activations appeared earliest in striate cortex and rapidly thereafter in other visual areas. Stimulus-specific processing began early in most visual cortical areas, some at activation onset. Early onset latencies were also observed in motor, premotor, and prefrontal areas, some as early as in striate cortex, but these early-activating frontal sites did not show early stimulus-specific processing. Response-specific processing began around 150 ms poststimulus in widespread cortical areas, suggesting that perceptual decision formation and response selection arose through concurrent processes of visual, motor, and executive areas. The occurrence of stimulus-specific and stimulus category-specific differences after the onset of response-specific processing suggests that sensory and motor stages of visuomotor processing overlapped in time.     

Changes in auditory cortex parallel rapid perceptual learning

Learning perceptual skills is characterized by rapid improvements in performance within the first hour of training (fast perceptual learning) followed by more gradual improvements that take place over several daily practice sessions (slow perceptual learning). Although it is widely accepted that slow perceptual learning is accompanied by enhanced stimulus representation in sensory cortices, there is considerable controversy about the neural substrates underlying early and rapid improvements in learning perceptual skills. Here we measured event-related brain potentials while listeners were presented with 2 phonetically different vowels. Listeners' ability to identify both vowels improved gradually during the first hour of testing and was paralleled by enhancements in an early evoked response ( approximately 130 ms) localized in the right auditory cortex and a late evoked response ( approximately 340 ms) localized in the right anterior superior temporal gyrus and/or inferior prefrontal cortex. These neuroplastic changes depended on listeners' attention and were preserved only if practice was continued; familiarity with the task structure (procedural learning) was not sufficient. We propose that the early increases in cortical responsiveness reflect goal-directed changes in the tuning properties of auditory neurons involved in parsing concurrent speech signals. Importantly, the neuroplastic changes occurred rapidly, demonstrating the flexibility of human speech segregation mechanisms.     

Nouns and verbs in the intact brain: evidence from event-related potentials and high-frequency cortical responses.

Lesion evidence indicates that words from different lexical categories, such as nouns and verbs, may have different cortical counterparts. In this study, processing of nouns and verbs was investigated in the intact brain using (i) behavioral measures, (ii) stimulus-triggered event-related potentials and (iii) high-frequency electrocortical responses in the gamma band. Nouns and verbs carefully matched for various variables, including word frequency, length, arousal and valence, were presented in a lexical decision task while electrocortical responses were recorded. In addition, information about cognitive processing of these stimuli was obtained using questionnaires and reaction times. As soon as approximately 200 ms after stimulus onset, event-related potentials disclosed electrocortical differences between nouns and verbs over widespread cortical areas. In a later time window, 500-800 ms after stimulus onset, there was a significant difference between high-frequency responses in the 30 Hz range. Difference maps obtained from both event-related potentials and high-frequency responses revealed strong between-category differences of signals recorded above motor and visual cortices. Behavioral data suggest that these different physiological responses are related to semantic associations (motor or visual) elicited by these word groups. Our results are consistent with a neurobiological model of language representation postulating cell assemblies with distinct cortical topographies as biological counterparts of words. Assemblies representing nouns referring to visually perceived objects may include neurons in visual cortices, and assemblies representing action verbs may include additional neurons in motor, premotor and prefrontal cortices. Event-related potentials and high-frequency responses are proposed to indicate two different functional states of cell assemblies: initial full activation ('ignition') and continuous reverberatory activity.     

Volatile Anesthetics Enhance Flash-induced γ Oscillations in Rat Visual Cortex

Background: The authors sought to understand neural correlates of anesthetic-induced unconsciousness. Cortical [gamma] oscillations have been associated with neural processes supporting conscious perception, but the effect of general anesthesia on these oscillations is controversial. In this study, the authors examined three volatile anesthetics, halothane, isoflurane, and desflurane, and compared their effects on flash-induced [gamma] oscillations in terms of equivalent concentrations producing the loss of righting reflex (1 minimum alveolar concentration for the loss of righting [MACLR]).

Methods: Light flashes were presented every 5 s for 5 min, and event-related potentials were recorded from primary visual cortex of 15 rats with a chronically implanted bipolar electrode at increasing anesthetic concentrations (0-2.4 MACLR). Early cortical response was obtained by averaging poststimulus (0-100 ms) potentials filtered at 20-60 Hz across 60 trials. Late (100-1,000 ms) [gamma] power was calculated using multitaper power spectral technique. Wavelet decomposition was used to determine spectral and temporal distributions of [gamma] power.

Results: The authors found that (1) halothane, isoflurane, and desflurane enhanced the flash-evoked early cortical response in a concentration-dependent manner; (2) the effective concentration for this enhancement was the lowest for isoflurane, intermediate for halothane, and the highest for desflurane when compared at equal fractions of the concentration that led to a loss of righting; (3) the power of flash-induced late (> 100 ms) [gamma] oscillations was augmented at intermediate concentrations of all three anesthetic agents; and (4) flash-induced [gamma] power was not reduced below waking baseline even in deep anesthesia.     

Hearing lips: gamma-band activity during audiovisual speech perception

Auditory pattern changes have been shown to elicit increases in magnetoencephalographic gamma-band activity (GBA) over left inferior frontal cortex, forming part of the putative auditory ventral "what" processing stream. The present study employed a McGurk-type paradigm to assess whether GBA would be associated with subjectively perceived changes even when auditory stimuli remain unchanged. Magnetoencephalograms were recorded in 16 human subjects during audiovisual mismatch perception. Both infrequent visual (auditory /ta/ + visual /pa/) and acoustic deviants (auditory/pa/ + visual /ta/) were compared with frequent audiovisual standards (auditory /ta/ and visual /ta/). Statistical probability mapping revealed spectral amplitude increases at approximately 75 and approximately 78 Hz to visual deviants. GBA to visual deviants peaked 160 ms after auditory stimulus onset over posterior parietal cortex, at 270 ms over occipital areas and at 320 ms over left inferior frontal cortex. The latter GBA enhancement was consistent with the increase observed previously to pure acoustic mismatch, supporting a role of left inferior frontal cortex for the representation of perceived auditory pattern change. The preceding gamma-band changes over posterior areas may reflect processing of incongruent lip movements in visual motion areas and back-projections to earlier visual cortex.     

Where vision meets memory: prefrontal-posterior networks for visual object constancy during categorization and recognition

Objects seen from unusual relative to more canonical views require more time to categorize and recognize, and, according to object model verification theories, additionally recruit prefrontal processes for cognitive control that interact with parietal processes for mental rotation. To test this using functional magnetic resonance imaging, people categorized and recognized known objects from unusual and canonical views. Canonical views activated some components of a default network more on categorization than recognition. Activation to unusual views showed that both ventral and dorsal visual pathways, and prefrontal cortex, have key roles in visual object constancy. Unusual views activated object-sensitive and mental rotation (and not saccade) regions in ventrocaudal intraparietal, transverse occipital, and inferotemporal sulci, and ventral premotor cortex for verification processes of model testing on any task. A collateral-lingual sulci "place" area activated for mental rotation, working memory, and unusual views on correct recognition and categorization trials to accomplish detailed spatial matching. Ventrolateral prefrontal cortex and object-sensitive lateral occipital sulcus activated for mental rotation and unusual views on categorization more than recognition, supporting verification processes of model prediction. This visual knowledge framework integrates vision and memory theories to explain how distinct prefrontal-posterior networks enable meaningful interactions with objects in diverse situations.     

Functional neuroanatomy of anticipatory behavior: dissociation between sensory-driven and memory-driven systems

The ability to anticipate predictable stimuli allows faster responses. The predictive saccade (PRED) task has been shown to quickly induce such anticipatory behavior in humans. In a PRED task subjects track a visual target jumping back and forth between fixed positions at a fixed time interval. During this task, saccade latencies drop from approximately 200 ms to <80 ms as subjects anticipate target appearance. This change in saccade latency indicates that subjects' behavior shifts from being sensory driven to being memory driven. We conducted functional magnetic resonance imaging studies with 10 healthy adults performing the PRED task using a standard block design. We compared the PRED task with a visually guided saccade (VGS) task using unpredictable targets matched for number, direction and amplitude of required saccades. Our results show greater activation during the PRED task in the prefrontal, pre-supplementary motor and anterior cingulate cortices, hippocampus, mediodorsal thalamus, striatum and cerebellum. The VGS task elicited greater activation in the cortical eye fields and occipital cortex. These results demonstrate the important dissociation between sensory and predictive neural control of similar saccadic eye movements. Anticipatory behavior induced by the PRED task required less sensory-related processing activity and was subserved by a distributed cortico-subcortical memory system including prefronto-striatal circuitry.     

Electrophysiological and haemodynamic correlates of face perception,recognition and priming

Face perception, recognition and priming were examined with event-related functional magnetic resonance imaging (fMRI) and scalp event-related potentials (ERPs). Face perception was associated with haemodynamic increases in regions including bilateral fusiform and right superior temporal cortices, and a right posterior negativity (N170), most likely generated in the superior temporal region. Face recognition was associated with haemodynamic increases in fusiform, medial frontal and orbitofrontal cortices, and with a frontocentral positivity from 550 ms poststimulus. Face repetition was associated with a positivity from 400 to 600 ms and behavioural priming. Repetition of familiar faces was also associated with earlier onset of the ERP familiarity effect, and haemodynamic decreases in fusiform cortex. These data support a multi-component model of face-processing, with priming arising from more than one stage.     

Independence of visual awareness from the scope of attention: an electrophysiological study

Recent brain imaging studies have revealed that increased neuralactivity along the ventral visual stream and parietal and frontalareas is associated with visual awareness. In order to studythe time-course and temporal aspects of awareness, we examinedelectrophysiological correlates of conscious vision in two maskingexperiments. The differences in event-related potentials (ERPs)between unmasked (consciously recognized) and masked (unrecognized)stimuli were considered to be electrophysiological correlatesof awareness. Two attentional conditions (global, local) wereincluded to examine the relationship between the scope of attentionand awareness. Two ERP-deflections were found to correlate withawareness. First, awareness was associated with a posteriornegative amplitude shift 130–320 ms after the stimulus.This effect was present in both attention conditions, suggestingthat it emerges independent of the scope of attention. Second,ERPs to unmasked stimuli became more positive as compared withmasked stimuli around 400 ms, peaking at parietal sites. Thiseffect was attenuated in the local attention condition, althoughthe participants were aware of the stimuli, suggesting thatthe late positivity does not directly correlate with visualawareness. The results imply that the earlier negativity isthe earliest and most direct correlate of visual awareness.     

Linking hemodynamic and electrophysiological measures of brain activity: evidence from functional MRI and intracranial field potentials

We investigated the relation between electrophysiological and hemodynamic measures of brain activity through comparison of intracranially recorded event-related local field potentials (ERPs) and blood-oxygenation level dependent functional magnetic resonance imaging (BOLD fMRI). We manipulated the duration of visual checkerboard stimuli across trials and measured stimulus-duration-related changes in ERP and BOLD activity in three brain regions: peri-calcarine cortex, the fusiform gyrus and lateral temporal-occipital (LTO) cortex. ERPs were recorded from patients who had indwelling subdural electrodes as part of presurgical testing, while BOLD responses were measured in similar brain regions in a second set of subjects. Similar BOLD responses were measured in peri-calcarine and fusiform regions, with both showing monotonic but non-linear increases in hemodynamic amplitude with stimulus duration. In sharp contrast, very different ERP responses were observed in these same regions, such that calcarine electrodes exhibited onset potentials, sustained activity over the course of stimulus duration and prominent offset potentials, while fusiform electrodes only exhibited onset potentials that did not vary with stimulus duration. No duration-related ERP or BOLD changes were observed in LTO. Additional analyses revealed no consistent changes in the EEG spectrum across different brain sites that correlated with duration-related changes in the BOLD response. We conclude that the relation between ERPs and fMRI differs across brain regions.     

Carbachol-induced network oscillations in the intact cerebral cortex of the newborn rat

In mature cortex, activation of the cholinergic system induces oscillatory network activity and facilitates synaptic plasticity. We used an in vitro preparation of the intact cerebral cortex and cortical slices of the neonatal rat to study carbachol (CCh, >or=30 micro M)-induced network oscillations during the early postnatal period. Multi-site extracellular recordings revealed CCh-induced transient beta oscillations with an average duration of 4.6 +/- 0.2 s, amplitude of 123 +/- 7.4 microV and frequency of 17.7 +/- 0.5 Hz. These oscillations propagated uniformly at 0.5-1.5 mm/s over the cortex and were reversibly blocked by tetrodotoxin (TTX) and atropine, indicating that they depended on action potentials and activation of muscarinic receptors. The activity was not blocked by bicuculline methiodide or gabazine, but was reversibly abolished by kynurenic acid, indicating that activation of glutamate receptors, but not GABA-A receptors, was required. CNQX caused a significant decrease in the power of the Fourier frequency spectrum of the CCh-induced oscillations and CPP or MK-801 completely blocked the activity, indicating a contribution of AMPA/kainate receptors and an essential role of NMDA receptors. Oscillations were synchronized between sites separated horizontally by approximately 1 mm and for delays of 2-8 ms. Synchronized activity between neighboring recording sites was very stable over repeated applications of CCh. Whole-cell recordings from morphologically identified pyramidal neurons in the intact cortex revealed a close temporal correlation between CCh-induced membrane oscillations and local field potential recordings. In contrast, CCh-induced oscillations recorded in coronal neocortical slices were smaller in amplitude and frequency, suggesting that a widespread network of intracortical axonal connections is required for their generation.     

Distributed and associative working memory

This study explores the cortical cell dynamics of unimodal and cross-modal working memory (WM). Neuronal activity was recorded from parietal areas of monkeys performing delayed match-to-sample tasks with tactile or visual samples. Tactile memoranda (haptic samples) consisted of rods with differing surface features (texture or orientation of ridges) perceived by active touch. Visual memoranda (icons) consisted of striped patterns of differing orientation. In a haptic-haptic task, the animal had to retain through a period of delay the surface feature of the sample rod to select a rod that matched it. In a visual-haptic task, the animal had to retain the icon for the haptic choice of a rod with ridges of the same orientation as the icon's stripes. Units in all areas responded with firing change to one or more task events. Also in all areas, cells responded differently to different sample memoranda. Differential sample coherent firing was present in most areas during the memory period (delay). It is concluded that neurons in somatosensory and association areas of parietal cortex participate in broad networks that represent various task events and stimuli (auditory, motor, proprioceptive, tactile, and visual). Neurons in the same networks take part in retaining in WM the memorandum for each trial, whether it is encoded haptically or visually. The VH association by parietal cells in WM is analogous to the auditory-visual association previously observed in prefrontal cortex. Both illustrate the capacity of cortical neurons to associate sensory information across time and across modalities in accord with the rules of a behavioral task.     

Task-dependency of the neural correlates of episodic encoding as measured by fMRI.

Using event-related functional magnetic resonance imaging (fMRI), the neural correlates of memory encoding can be studied by contrasting item-related activity elicited in a study task according to whether the items are remembered or forgotten in a subsequent memory test. Previous studies using this approach have implicated the left prefrontal cortex in the successful encoding of verbal material into episodic memory when the study task is semantic in nature. In the current study, we asked whether the neural correlates of episodic encoding differ depending on type of study task. Seventeen volunteers participated in an event-related fMRI experiment in which at study, volunteers were cued to make either animacy or syllable judgements about words. A recognition memory test followed after a delay of approximately 15 min. For the animacy task, words that were subsequently remembered showed greater activation in left and medial prefrontal regions. For the syllable task, by contrast, successful memory for words was associated with activations in bilateral intraparietal sulcus, bilateral fusiform gyrus, right prefrontal cortex and left superior occipital gyrus. These findings suggest that the brain networks supporting episodic encoding differ according to study task.     

Tracking the temporal dynamics of updating cognitive control: an examination of error processing

In 2 experiments, we used event-related brain potentials (ERPs) to examine the temporal dynamics of neural processes related to adjustments of cognitive control following errors in the counting Stroop task. The ERPs elicited by errors revealed the error-related negativity (ERN) and error positivity consistent with a large body of literature. In addition, errors were associated with a frontal slow wave between 200 and 2000 ms after the response that was consistent with the activity of neural generators in the lateral frontal cortex. The ERN and frontal slow wave were correlated with posterror slowing of response time and positive affect (i.e., happiness and calmness) during task performance. These data are consistent with the idea that interactions between anterior cingulate cortex and lateral frontal cortex support adjustments of cognitive control and that this neural network is sensitive to the influence of affect experienced during task performance.