Tobias Egner

Emotional processing in anterior cingulate and medial prefrontal cortex

Negative emotional stimuli activate a broad network of brain regions, including the medial prefrontal (mPFC) and anterior cingulate (ACC) cortices. An early influential view dichotomized these regions into dorsal-caudal cognitive and ventral-rostral affective subdivisions. In this review, we examine a wealth of recent research on negative emotions in animals and humans, using the example of fear or anxiety, and conclude that, contrary to the traditional dichotomy, both subdivisions make key contributions to emotional processing. Specifically, dorsal-caudal regions of the ACC and mPFC are involved in appraisal and expression of negative emotion, whereas ventral-rostral portions of the ACC and mPFC have a regulatory role with respect to limbic regions involved in generating emotional responses. Moreover, this new framework is broadly consistent with emerging data on other negative and positive emotions.

Resolving Emotional Conflict: A Role for the Rostral Anterior Cingulate Cortex in Modulating Activity in the Amygdala

Effective mental functioning requires that cognition be protected from emotional conflict due to interference by task-irrelevant emotionally salient stimuli. The neural mechanisms by which the brain detects and resolves emotional conflict are still largely unknown, however. Drawing on the classic Stroop conflict task, we developed a protocol that allowed us to dissociate the generation and monitoring of emotional conflict from its resolution. Using functional magnetic resonance imaging (fMRI), we find that activity in the amygdala and dorsomedial and dorsolateral prefrontal cortices reflects the amount of emotional conflict. By contrast, the resolution of emotional conflict is associated with activation of the rostral anterior cingulate cortex. Activation of the rostral cingulate is predicted by the amount of previous-trial conflict-related neural activity and is accompanied by a simultaneous and correlated reduction of amygdalar activity. These data suggest that emotional conflict is resolved through top-down inhibition of amygdalar activity by the rostral cingulate cortex.

Cognitive control mechanisms resolve conflict through cortical amplification of task-relevant information

A prominent model of how the brain regulates attention proposes that the anterior cingulate cortex monitors the occurrence of conflict between incompatible response tendencies and signals this information to a cognitive control system in dorsolateral prefrontal cortex. Cognitive control is thought to resolve conflict through the attentional biasing of perceptual processing, emphasizing task-relevant stimulus information. It is not known, however, whether conflict resolution is mediated by amplifying neural representations of task-relevant information, inhibiting representations of task-irrelevant information, or both. Here we manipulated trial-by-trial levels of conflict and control during a Stroop task using face stimuli, while recording hemodynamic responses from human visual cortex specialized for face processing. We show that, in response to high conflict, cognitive control mechanisms enhance performance by transiently amplifying cortical responses to task-relevant information rather than by inhibiting responses to task-irrelevant information. These results implicate attentional target-feature amplification as the primary mechanism for conflict resolution through cognitive control.

Expectation (and attention) in visual cognition

Visual cognition is limited by computational capacity, because the brain can process only a fraction of the visual sensorium in detail, and by the inherent ambiguity of the information entering the visual system. Two mechanisms mitigate these burdens: attention prioritizes stimulus processing on the basis of motivational relevance, and expectations constrain visual interpretation on the basis of prior likelihood. Of the two, attention has been extensively investigated while expectation has been relatively neglected. Here, we review recent work that has begun to delineate a neurobiology of visual expectation, and contrast the findings with those of the attention literature, to explore how these two central influences on visual perception overlap, differ and interact.

Predictive Codes for Forthcoming Perception in the Frontal Cortex

Incoming sensory information is often ambiguous, and the brain has to make decisions during perception. “Predictive coding” proposes that the brain resolves perceptual ambiguity by anticipating the forthcoming sensory environment, generating a template against which to match observed sensory evidence. We observed a neural representation of predicted perception in the medial frontal cortex, while human subjects decided whether visual objects were faces or not. Moreover, perceptual decisions about faces were associated with an increase in top-down connectivity from the frontal cortex to face-sensitive visual areas, consistent with the matching of predicted and observed evidence for the presence of faces.

Congruency sequence effects and cognitive control

Congruency effects in selective attention tasks are subject to sequential modulation: They are smaller following an incongruent stimulus than following a congruent one. This congruency sequence effect has been interpreted as reflecting conflict-driven adjustments in cognitive control (conflict adaptation) or, alternatively, episodic memory effects of stimulus—response association (feature integration). The present article critically reviews support for these rival accounts in the experimental literature and discusses the implications thereof for assessing behavioral and neural signatures of cognitive control processes. It is argued that both conflict adaptation and feature integration contribute to the congruency sequence effect but that their respective contributions can be isolated experimentally. Studies that have pursued this isolation strategy have gained important insights into cognitive control processes. Finally, other factors, such as expectancies, may also contribute to the congruency sequence effect, and thus their potential role needs to be carefully examined and, if found significant, integrated into current models of cognitive control.

The effect of training distinct neurofeedback protocols on aspects of cognitive performance

The use of neurofeedback as an operant conditioning paradigm has disclosed that participants are able to gain some control over particular aspects of their electroencephalogram (EEG). Based on the association between theta activity (4-7 Hz) and working memory performance, and sensorimotor rhythm (SMR) activity (12-15 Hz) and attentional processing, we investigated the possibility that training healthy individuals to enhance either of these frequencies would specifically influence a particular aspect of cognitive performance, relative to a non-neurofeedback control-group. The results revealed that after eight sessions of neurofeedback the SMR-group were able to selectively enhance their SMR activity, as indexed by increased SMR/theta and SMR/beta ratios. In contrast, those trained to selectively enhance theta activity failed to exhibit any changes in their EEG. Furthermore, the SMR-group exhibited a significant and clear improvement in cued recall performance, using a semantic working memory task, and to a lesser extent showed improved accuracy of focused attentional processing using a 2-sequence continuous performance task. This suggests that normal healthy individuals can learn to increase a specific component of their EEG activity, and that such enhanced activity may facilitate semantic processing in a working memory task and to a lesser extent focused attention. We discuss possible mechanisms that could mediate such effects and indicate a number of directions for future research.

Learned self-regulation of EEG frequency components affects attention and event-related brain potentials in humans

Learned enhancement of EEG frequency components in the lower beta range by means of biofeedback has been reported to alleviate attention deficit hyperactivity disorder (ADHD) symptoms. In order to elucidate frequency-specific behavioural effects and neurophysiological mediators, this study applied neurofeedback protocols to healthy volunteers, and assessed impact on behavioural and electrocortical attention measures. Operant enhancement of a 12–15 Hz component was associated with reduction in commission errors and improved perceptual sensitivity on a continuous performance task (CPT), while the opposite relation was found for 15–18 Hz enhancement. Both 12–15 Hz and 15–18 Hz enhancement were associated with significant increases in P300 event-related brain potential amplitudes in an auditory oddball task. These relations are interpreted as stemming from band-specific effects on perceptual and motor aspects of attention measures.

Intentional false responding shares neural substrates with response conflict and cognitive control

The ability to deceive others is a high-level social and cognitive function. It has been suggested that response conflict and cognitive control increase during deceptive acts but this hypothesis has not been evaluated directly. Using fMRI, we tested this prediction for the execution of an intentional false response. Subjects were instructed to respond truthfully or falsely to a series of yes/no questions that were also varied in autobiographical and nonautobiographical content to further examine the influence of personal relevance when lying. We observed an interference effect (longer reaction times for false versus true responses) that was accompanied by increased activation within the anterior cingulate, caudate and thalamic nuclei, and dorsolateral prefrontal cortex (DLPFC), a circuit that has been implicated in response conflict and cognitive control. Behavioral and neural effects were more robust when falsifying autobiographical responses relative to nonautobiographical responses. Furthermore, a correlation between reaction time and left caudate activity supported the presence of increased response inhibition when falsifying responses. When presented with self-relevant (autobiographical) stimuli regardless of response condition, the mesial prefrontal and posterior cingulate cortices were recruited. Neural activity within these two regions and the anterior cingulate cortex (ACC) also showed correlations with self-report personality measures from the Psychopathic Personality Inventory (PPI). Overall, we conclude that the process of interference is inherent to the act of falsifying information and that the amount of conflict induced and cognitive control needed to successfully execute false responses is greater when dealing with personal information.

Separate conflict-specific cognitive control mechanisms in the human brain

To ensure optimal task performance, the human brain detects and resolves conflict in information processing via a cognitive control system. However, it is not known whether conflict resolution relies on a single central resource of cognitive control, or on a collection of independent control mechanisms that deal with different types of conflict. In order to address this question, we assessed behavioral and blood-oxygen-level-dependent (BOLD) responses during the simultaneous detection and resolution of two sources of conflict in a modified color-naming Stroop task: conflict stemming from incompatibility between the task-relevant and an irrelevant stimulus feature (stimulus-based or Stroop conflict), and conflict stemming from incompatibility between an irrelevant stimulus feature and response features (response-based or Simon conflict). Results show that control mechanisms recruited by stimulus-based conflict resolve stimulus-based conflict, but do not affect the resolution of response-based conflict, and vice versa. The resolution of response-based conflict was distinguished by modulation of activity in premotor cortex, whereas resolution of stimulus-based conflict was distinguished by the modulation of activity in parietal cortex. These results suggest that the human brain flexibly adopts, and independently controls, conflict-specific resolution strategies, biasing motor programming to resolve response-based conflict, and biasing stimulus representations to resolve stimulus-based conflict. We propose a non-centralized, modular architecture of cognitive control, where separate control resources operate in parallel, and are recruited in a context-sensitive manner.